;-jj5^]^

o

B

CATALOGUE

OF THE

MAMMALS

OF

WESTERN EUEOPE

(EUROPE EXCLUSIVE OF RUSSIA)

IN THE

COLLECTION

OF THE

BEITISH MUSEU])^^^?^

BY

GEREIT S. MILLER

/-/■

LONDON

PRINTED BY ORDER OF THE TRUSTEES OF THE

BRITISH MUSEUM

SOLD BY

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

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

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

AND at the

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

1912

All rights reserved

LONDON :

PRINTED BY WILLIAM CLOWES AND SONS, LIMITED,

DUKE STREET, STAMFORD STREET, S.B., AND GREAT WINDMILL STREET, W.

hh.y^i '^1^' '^'^-^ dhu't.

PEEFACE

Although the idea of a publicatiou ou the Mammals of
Europe was suggested many years ago by the late Lord
Lilford, who kindly contributed an annual sum towards the
collecting necessary for its realization, the possibility of
issuing the present Catalogue has mainly grown up from the
work which its author, Mr. Gerrit S. Miller, of the United
States National Museum at Washington, has for some years
been doing independently on the subject.

It is true that European Mammals had not been neglected
here, and that the collection had begun to grow, both by
the help of the Lilford Eund and by the efforts of Major
G. E. H. Barrett-Hamilton, who published many papers on
the subject, and of Mr. Oldfield Thomas, E.E.S., who devoted
a number of his vacations to collecting in various parts of
the Continent.

There was, however, no prospect of being in a position
to prepare a Catalogue until about 1905, when Mr. Miller
arranged to devote his entire time for a considerable period
to the study of European Mammals. The opportunity was
taken of having the results of this work published here
instead of in America, by inducing him to write a British
Museum Catalogue ; thus utilizing his knowledge, and com-
bining for the purposes of his studies the material of both
the American and the British National Museums.

Collections were then made in various selected areas,
partly by Mr. Miller himself and partly by trained collectors,
such as Messrs. A. Eobert, C. Mottaz, Eev. S. Gonzalez and
N. Gonzalez, the cost of whose services were contributed by
friends of the Museum, notably Mr. Oldfield Thomas, the
Hon. N. C. Itothschild and ^Ir. J, I. S. Whitaker. The
Catalogue could hardly have been contemplated if it had
not been for Mr. Thomas' unremitting efforts in developing

IV PREFACE

tlie collection. He has not merely regarded tliese efforts as
an official duty, but he has in addition been a generous
donor who has frequently supplied funds for the purpose of
obtaining specimens. Mr. Miller has thus had at his
disposal a collection fairly representative of all parts of
Western Europe, and immensely superior to anything that
had been thought of before he began his work.

Marine Mammals (Cetacea and Pinnipedia) are not
included in the present Catalogue. For a definition of
" Western Europe " reference must be made to page vii of
the Author's Introduction.

As Mr. Miller is on tlie staff of the United States
National Museum the special and cordial thanks of the
Trustees of the British Museum are due to the authorities of
the former Institution for the facilities granted to him for
carrying through the preparation of the Catalogue, a work
which involved a furlough of two years and a half from his
usual duties at Washington.

The thanks of the Trustees are also due to Mrs. Oldfield
Thomas and to Mr. E. C. Wroughton for their kindness
in undertaking the considerable labour of preparing and
verifying the lists of the specimens in the Museum collection
after Mr. Miller had made his scientific determinations.

SIDNEY E. HAEMER,

Keeper of Zoology.

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

October, 1912.

INTEODUCTION

The collection of European Land-mammals in the pjiitish
Museum consists of about five thousand specimens. One
hundred and twenty-four of these are types. It has for the
most part been brought together during the past thirty years
through the efforts of the late Lord Lilford, of Mr. Oldfield
Thomas, and of Major G. E. H. Barrett-Hamilton. The older
material, though not extensive, includes much that is of historic
interest, such as the numerous specimens received from the late
Baron E. de Selys-Longchamps, the types of various species
described by Gray and Bonaparte, and Darwin's Porto Santo
rabbits which have been the subject of so much groundless
speculation. It is, however, from the recently-obtained material
that the collection derives its true value. These specimens are
almost without exception carefully-prepared skins accompanied
by skulls and measurements, together with full records of sex,
date, and exact locality. The more important sources from
which they were obtained are as follows : collections brought
together by Professor W. Wolterstorff from central and eastern
(Jermany, and presented by the late Lord Lilford ; collections
made in Roumania by the late E. Dodson under the direction
of Major G. E. H. Barrett-Hamilton, and presented by the late
Lord Lilford ; material from southern Spaia presented by the
late Lord Lilford ; extensive collections made in south-western
France, in southern Italy and in Sicily by A. Pvobert and
presented by Mr. Oldfield Thomas; collections from south-
central France and the vicinity of Strassburg, Germany, made
by C. Mottaz and presented by Mr. O. Thomas ; small collections
from Denmark, Holland, Pas-de-Calais, Brittany, Portugal, the
Balearic Islands, Switzerland and northern Italy, made and
presented by Mr. O. Thomas ; collection from central and
northern Spain made by N. Gonzalez and presented by i\Ir. O.
Thomas ; a large collection from miscellaneous sources brought
together and presented by Major G. E. H. Barrett-Hamilton;
a collection from Greece made by C. Mottaz and presented by

VI INTRODUCTION

Mr. J. I. S. Whitaker and the Hon. N. C. Rothschild; a
collection from Spain and southern France made by G. S.
Miller and purchased by the Museum ; several collections
from Transylvania made by C. G. Danford ; collections from
Hungary made and presented by the Hon. N. C. Eothschild
and Mrs. Rothschild ; smaller collections and single specimens
have been received from many other persons,* whose names will
be found in the detailed lists of material in this Catalogue.

Although unquestionably forming the largest of all collections
of European mammals the material in the British Museum is
not sufficient to be made the basis of a monographic study of
the fauna. Free use has, therefore, been made, throughout the
preparation of this Catalogue, of specimens in other collections.
Chief among these are the United States National Museum in
Washington and the private collection of Charles Mottaz in
Geneva. The material at Washington, about 4000 specimens,
is mostly from the following sources : (a) Sweden, Germany,
Switzerland and Belgium, collected by J. Alden Loring ; (b)
Sicily, Italy and the region of Barcelonnette, Basses-Alpes,
France, by Dane Coolidge ; (e) south-western France, by
Robert T. Young ; (d) north-eastern Germany, the Riesengebirge
and Hai'z Mountains, by F. L. J. Boettcher. There are also
miscellaneous smaller collections from Switzerland (G. S. Miller,
L. Stejneger, E. H. Zollikofer), Belgium (de Selys-Longchamps),
Holland (G. S. Miller), Denmark (L. Stejneger), Norway (T.
Stejneger) and Sweden (Sundevall. Tullberg, Lonnberg). Finally,
the Merriam collection, now the property of the U.S. National
Museum but not yet catalogued, contains numerous European
specimens, for the most part received from de Selys-Longchamps.
The Mottaz collection, about 3000 specimens, is especially rich
in series of the smaller mammals of Switzerland and the adjoin-
ing portions of France ; it also contains useful material from
Italy and western France (Charente). Other supplemental
material to which I have been given free access, or which has
been sent for examination in London or Washington, is contained
in the museums of Madrid (types of Cabrera), Nimes (types of
Crcspon), Paris (types of Geoffroy and other historic speci-
mens), Genoa (Italian Bats, Microtines and Ungulates), Turin
(Italian mammals, especially Ungulates), Naples (type of Mi/otis
oxygnathus Monticelli), Geneva (types of Fatio, authentic Swiss

* This is particularly true of the many friends of the Museum who
have aided in procuring the large series of British mammals.

INTRODUCTION VU

specimens of Lynx), Lausanne Agricultural School (skull of
Ursus "formicnriiis " from the Alps), Munich (type of S/ialnx
(jrsecus Nehring), Berlin Agricultural High School (type of
Arvicola rattireps stimminiji Nehring), Breslau (skulls of foxes),
Leiden (co-types of Arvicola arenarius de Selys-Longchamps),
Copenhagen (Mus fseroensis and small carnivores), Christiania
(Sorex, Evotomys, etc), Stockholm (Swedish carnivores and
rodents), Cambridge {Mustela erminea ricinse, Lemmus lemmus
" n-nssidens") and Edinburgh (rodents from northern Scotland).
Private collections which have been in the same generous
manner placed at my disposal are those of Mr. Angel Cabrera,
of Madrid (Spanish mammals, including several types). Dr.
Enrico Festa, of Turin (Italian mammals), Mr. Angelo Ghidiui,
of Geneva (Swiss and north Italian mammals), and Dr. Fernand
Lataste, of Cadillac-sur-Garonne, France (carnivores and micro-
tines).

The total number of sj^ecimens on which this work is based
approximates 11,500. All those of which definite record has
been made are enumerated in the paragraphs headed : Specimens
examined* Absence of a note to the contrary indicates that
all the specimens from a given locality are in the British
Museum. Discrepancies frequently occur between the number
of " specimens examined " and the number tabulated in the final
paragraph as forming part of the Museum collection. These
result from the fact that under "specimens examined" are
included duplicates as well as registered specimens, while only
the latter appear in the final lists. f

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

* In these lists 11,372 specimens are recorded. They are distributed
as follows: Insectivora, 1,777 ; Chiroptera, 2,210; Carnivora, 877 ; Rodentia
Duplicidentata, 379, Rodentia Simplicidentata, 5,854 ; Ungulata, 28i.

t Certain duplicates have been transferred to the United States
National Museum since the lists of " specimens examined " were prepared.
No attempt has been made to alter the records in the lists on this account ;
but the U.S.N.]\I. numbers of such specimens are frequently to be found
in the Tables of cranial measurements. (All numbers above 10,000 indicate
specimens in Washington.)

X For use of this name in place of " Macacus inuus," see Thomas, Proc.
Zool. Soc. London, 1911, pp. 125-126, :March, 1911.

VIU INTRODUCTION

their presence to artificial introduction, are treated mono-
graphically on the basis of the material already enumerated.
This material has been found sufficient, in most of the groups, to
give what appears to be a fairl}' satisfactory idea of the essential
features of the fauna. In the ungulates and the larger car-
nivores, however, it is so totally inadequate that no attempt
could be made to revise the genera by which they are repre-
sented. This is especially to be regretted on account of the fact
that some of these larger mammals are nearly extinct, while
others are being modified by the introduction of foreign stock
to replenish exhausted game preserves. Immediate action is
necessary if the final opportunity to gain a clear understanding
of this part of the European fauna is not to he lost.

The litei'ature of European mammals is so voluminous,
particularly as regards local lists and special notes on distri-
bution, and it is for the most part based on conceptions of
species and local races so different from those underlying the
present work, that an amount of labour incommensurate with
the importance of the results would be required to prepare
extended bibliographical Tables for each form recognized. The
citations are, therefore, restricted to those which seem of impor-
tance in giving a clear idea of the systematic history of each
animal ; that is, to the specific and sub-specific names under
which it may have been described, to the first use of the actual
binomial or trinomial here adopted, to the names used in the
monographic works of Blasius, 18-57, and Trouessart, 1910, and
to any uthei- publication which might seem pertinent to a
particular case.

In deciding questions of nomenclature, an attempt has been
made to apply the International Code and the rulings of the
Commission strictly and consistently, even to the reluctant
acceptance of the terms applied to genera by authors who
f(jllowed a system different from that now in use.

With the exception of figure 121, lent by the Smithsonian
Institution, all the illustrations are original. The di'awings of
teeth were made in London by Mr. A. J. Engel Terzi ; part
of those of the skulls were made by Mr. Terzi ; the rest were
done in Washington by Mr. H. B. Bradford.

A few words in conclusion regarding the actual making of
the manuscript. I prepai'ed all the descriptions, synonymies,
lists of specimens examined, and Tables of cranial measurements.
The external measurements, which are not to be regarded as

INTRODUCTION IX

more thau approximately accurate, are mostly given as recorded
on the labels, though much verification and correction for ears
and hind feet has been done from the dried specimens. In order
to economize time, the records of registered material were made
directly from the specimens which I had identified. Mrs.
Oldfield Thomas and Mr. R. C. Wroughton carried out this
portion of the work.

G. S. M.

Washington,

July 1, 1912.

Order.

Number of

genera
recognized.

Number of

forms
recognized.

Number of

forms not

represented

in B.M.

Number of

recognized

forms not

seen.

Insectivora

7

45

5

0

Chieoptera

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

m

314

22

G

SYSTEMATIC INDEX

ORDER INSECTIVORA.

Family TALPID^ . .

Sub-Family 1. Talpin-tj: .
1. Talpa Linnaeus

1. europgea Linnfcus .

2. casca Savi

3. occidentalis Cabrera

4. romana Thomas

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

1. pyrenaicus Geoffroy
a. pyrenaicus Geoffroy
h. rufulus Graells .

Family SORICID^ . .

1. Sores Linnseus

1. araneus LiniuBUs .

a. araneus Liniueus

b. castaneus Jeoyns

c. santonus Mottaz

d. euronotus IMiller

e. bergensis Miller .
/. tetragonurus Hermann
g. pyrenaicus Miller
h. fretalis Miller
i. granarius Miller .

2. rainutus Linna»us .
a. minutus Liniia'us
h. lucanius Miller .

3. alpiuus Schinz .
a. alpinus Schinz .
h. hercynicus Miller

2. Neomys Kaup .

1. fodiens Schrebcr
a. fodiens Schreber.
h. bicolor Shaw .

2. milleri Mottaz .

3. anomalus Cabrera .

3. Pach>'ura de Selys-Long

champs ....
1. etrusca Savi.

4. Crocidura NVagler.

1. leucodou Hermann

2. mimula Miller .
a. mimula Jlillor .
h. iculisma Mottaz ,
c. cantabra Cabicra

2
3
3
15
15
18
20
20
21
26
2G

28
29
31
35
37
40
41
41
42
44
45
52
53
55
GO
CO
62
63
65
66
69
73
78
81

81
83
86
88
94
95
98
99

PAGE

8. russula Hermann ... 99

a. russula Hermann . . 101

h. pulchra Cabrera . . . 103

c. cintrre Miller. ... 108

4. sicula :Miller .... 108

5. cauere Miller .... 109

6. caudata Miller .... 110

7. cyrnensis Miller . . . Ill

8. balearica Miller ... 112

Family ERINACEID.-E ... 114

1. Erinaoeus Linnaeus . . . 114

1. europseus Linna'us. . . 115
a. europreus Linna'Us . . 120
h. hisiaanicus Bariett-

Hamilton . . . . . 122

c. italicus Barrett-Hamil-
ton 123

d. consolei Barrett-Hamil-
ton 126

2. roum aniens Barrett-

Hamilton 127

3. nesiotes Bate .... 129

4. algirus Duvernov and

Lereboullet . ". . . 130

a. algirus Duvernoy and
Lereboullet .... 131

b. vagans Tliomas . . . 133

ORDER CHIROPTERA.

Sob-Ordee MICROCHIROPTERA.

Family RHINOLOPHID.^ . . 136

1. Rhiuolophus Lac6pede . . 137

1. ferruin-oquinum Schrober 189

a. ferrum-equinum Schre-
ber 142

b. i n s u 1 a n u s Barrett-
Hamilton 147

2. hipposideros Bcchstein . 147

a. hipposideros liechstein . 149

b. minimus Heuglin . . 151

c. minutus Montagu . . 154
8. euryalc Blasius .... 155

4. meholyi Matschie . . . 159

5. blasii Peters 162

SYSTEMATIC INDKX

Order Chikopteka — Sub - Order

MlCBOCHlKOPTEBA— aw^i.

Family VESPERTILIONlD.t: . 165

Sub-Family Vespkktiuo:sis.t: . 165

1. ilyotis Kaup 166

1. mystaciuus Kohl . . 169

2. nattereri Kohl .... 17-i

3. emargiuatus Geoffroy . . 177

4. bechsteimi Kuhl . . 179

5. daubentonii Kuhl . . 1S4

6. capaccinii Bouaparce . . 1S7

7. dasycneme Boie . . . 189

8. myotis Borkhauseu . 192

9. oxvgnathus Monticelli . 199

2. Pipistrellus Kaup ... 202

1. pipistrellus Schrebor . . 201

2. uatkusii Keyserliug aud

Blasius 213

3. kuhlii Kuhi 215

4. savii Bonaparte .... 219

3. Eptesicus Rafinesque . . 224

1. serotinus Schreber . . 226

2. sodalis Barrett-Hamiltou . 231

3. uilssonii Kevserling and

Blasius. .".... 234

4. Vespertilio Linnaeus . . . 238
1. murinus Lin me u-^ . . . 238

5. Nyctalus Bowdieh ... 242

1. maximus Fatio. . . . 244

2. noctula Schreber . . . 245

3. leisleri Kuhl .... 252

4. azoreuni Thomas . . . 254

6. Plecotus GeofEroy. . . 256
1. auritus Linnwus . . . 256

7. Barbastella Gray .... 263
1. barbastellus Schrebt-r . . 263

Sub-Family Minioptesisj. . . 266

1. Miniopterus Bonaparte . . 268

1. schreibersii Kuhl . . . 269

Family MOLOSSID.E ... 276

1. Nyctrnomus Geoflroy . . 276

1. teniotis Rafinesque . . 277

ORDER CARNIVORA.

Family URSID.E 2S4

1. Ursus Linnseus .... 285
1. arctos T.inn.f n< . . . 285

2. Thalarctos Gray .... 297
1. maritimus Fhipp> . . . 298

Family CAXID.F 303

1. Canis Linnfeus .... 304

1. lupus Linnseus .... 305
a. lupus LinnKUs . . . 313
6. signatus Cabrera . . 314
c. deitanus Cabrera . . 315

2. aureus Liunieus . . . 315

VAGK

2. Alopex Kaup 318

1. lagopus Liiuuvus . . . 319

2. spitzbergenensis Barrett-

Hamilton aud Bouhote . 324

3. Vulpes Oken 325

1. vulpes Linn*us . . . 326
a. vulpes Liuuams . . . 330
6. crucigera Bechsieiu. . 331
c. silacea Miller. . . 333

2. ichnusse Miller .... 336

Family MUSTELID.E . . . 340

Sub- Family Mklix.?; .... 341

1. Meles Brissou 341

1. meles Linnaeus .... 343

a. meles Linnseus . . . 348

b. mariauensis GraeUs. . 352

2. arealus Miller .... 352
Sub-Family LuxRrs.t: .... 354

1. Lutra Brissou 354

1. lutra Linmvus .... 355

Sub-Family Mustehn.*; . . . 364

1. Martes I'inel 365

1. martes Linnieus . . . 366

a. martes Linnseus . . . 372

b. latinorum Barrett-Ha-
milton 373

2. foina Erxleben .... 374
(I. foina Erxleben . . . 375
h. mediterranea Barrett-
Hamilton 360

3. bunites Bate .... 380

2. Mustela Linnseus. . . . 381
Sub-geiius Mustela Linnseus . 384

1. erminea Linnseus . . . 385

a. erminea Linuseus . . 387

b. sBstiva Kerr .... 389

c. stabilis Barrett-Hamil-
ton 390

d. riciuifi MiUer. ... 397

2. hibernica Thomas and

Barrett-Hamilton . . 398

3. nivalis Linnseus . . . 401
a. nivalis Linnseus . . 402
6. boccamela Bechstoiii . 405
c. iberica Barrett-Hamilton 407

4. africana Desmarest . . 412

5. galinthias Bate . . . 414
Sub-Genus Lutreola Wagner . 415

6. lutreola Linnseus . . . 415
Sub-Genus Putorius Cuvier . 418

7. putorius Linnseus . . , 419

a. putorius Linnseus . . 423

b. aureolus Barrett-Hamil-

ton 425

3. Vormela W. Blasius ... 428
1. peregusna Gueldenstaedt . 429

Sub-Family Gulonls-E . . . 432

1. Gulo Storr 433

1. ?ulo Linnseus .... 434

KTSmUTIC IKDEX

PAGK

(jrder Cabsivoea — conlinu^l.

Family VIVERKID>iE . .

440

1. Mungofi Geoff roy and Cuvier 440

1. widdringtonii Gray

441

2. Genetta Oken ....

44e

1. genetta LinnasuB .

447

a. genetta Ltinnmnn

451

b. balearica Thomasj .

452

c. rhodanica Matschie .

452

Family FBLID>E ....

1-5.5

1. Felk Linrijeus.

456

1. Hilvestrii-; Schrel>er .

457

a. silvestris Schreter .

462

fc. grarapia Miller .

464

c. tartesBia Miller .

465

2. sarda Lataste .

468

3. agrius Bate ....

470

2. Lynx Kerr

470

1, lynx Linnajus .

471

2, pardellus Miller . .

475

ORDER RODENTIA.
Sub-Okdkk duplicidentata.

Family LEPORID^ .... 484

1. OryctoLagus Lilljeborg . . 484
1. cuni/.-uluH Linrijeiis . , 48-5

o. cxiniculu.8 Linrijeus . . 490

6. huxleyi Haeckel . . . 491

2. Lepus Linnaeus .... 495

1. europfftUB PaJlas . . . 498

a. europaius Pallas. . . 502

b. occidentalis de Winton. 50^

c. pyrenaicus Hilzheimei* . 506

d. meridiei Hilzheimer . 506

e. corsicanus de Winton . .507
/. hybriduB Desmarest. 508
g. tranBsylvanicus Mat-
schie 509

2. creticus Barrett-Hamilton 512

3. mediterraneus Wagner . 513

4. granatensis Flosenhauer . 515
a. granat^jnsis Pi^jsenhauer 516
6. gaUiecius Miller . . 517

c. iturisBiuB Miller . 518

d. pamasBius Miller . . 519

5. timidus Linnaeus . . . 522

a. timidus Linnaeus . 526

b. varronis Miller . . . 528

c. scoticus Hilzheimer. . 529

6. hibemicus Bell. . . . 531

Scb-Obdee SIMPLICIDENTATA.

Family ZAPODIDiE .... 5-35

Scb-Family S1CIETIH.S; . . . 536

1. Sicista Gray 536

1. loriger Nathusius . . . 537

2. trizona Pet^nyi .... 539

J-ACE

Family HYSTBICIDiE . .

.542

1. Hyetrix Linnaeus .

542

1. crigtata Linnseus

543

Family MUSCARDINID^^ .

549

1. Eliomys 'SVagner .

5.50

1. quercinus LinnseuB

5-51

2. gymnesicus Thomas .

558

3. paUiduB Barrett-Hamilton 559

4. sarduB Barrett-Hamilton

500

5. lusitanicus Pi^uvens .

560

2. DyromyB Thomas.

.566

1, nitedula Pallas . . .

567

a. nitedula Pallas .

568

6. intemiediuB Nehring

569

c. wingei Nehring .

570

2. robustus Miller.

572

3. Glis BrisBOn ....

572

1. glis Linnaeus

.573

a. glis Linnaeus .

.577

t. italicus Barrett-Hamil

ton

.578

c. melonii Thomas .

579

d. pyrenaicus Cabrera .

582

4. Muscardinus Kaup .

583

1. aveDanarius Linnaeus .

583

2. pulcher Barrett-Hamilton 590

Family MUEID.S: . . .

591

Scb-Family Ceicetot^ . .

592

1. CriMjtultiB Milne-Edwards

.593

1. atticus Nehring

593

2. Cricetus Leske

.596

1. cricetus Linnaeus .

597

a. cricetus Linn3?UB

602

6. canesoens Nehring .

603

c. nehringi Matschie .

605

3. ilesocricetuB Nehring .

605

1. newtoni Nehring .

606

Sub-Family Micbotis^ . .

610

1. MyopuB Miller

611

1. Bchisticolor Lilljeborg.

611

2. LemmusLink.

614

1. lemmuB Linnaeus .

615

3. Evotomys Coues .

623

1. glareolus Schreber .

626

a. glareolus Schreber .

a32

b. britannicus Miller .

6-34

c. suecicuB JliUer .

6-36

d. istericus iliUer .

6-37

e. norvegicns ililler

6.38

/. vasconiae Miller .

639

g. helveticus Miller

640

k. nageri Schinz

641

i. hallucalis Thomas .

643

2. skomerensis Barrett -Ha

milton ....

. 644

3. caesariuB ililler .

. 645

4. rutilus Pallas .

. 646

5. rafocanus Sundevall .

. 648

SYSTEMATIC INDEX

and

Order Rodentia — Sub - Order
SiMPLiciDENTATA — Family
MuEiD.E — Sub-Family ili-
CBOTIN.E —continued.

4. Microtus Schrank
Sub-Genus Microtus Schrank

1. agrestis Linuieus .

a. agrestis Linnseus

b. exsul Miller .

c. levernedii Crespon

d. bailloni de Selys-Long-
champs

e. hircus Bellamy .
/. neglectus Jenyns .
g. rozianus Bocage .

2. arvalis Pallas

a. arvalis Pallas

b. meridianus PaUas

c. duplicatus Rorig
Borner

d. levis Miller .

3. incertus de S61ys - Loug-

champs

4. asturianus Miller .

5. orcadensis Millais .

6. sandayensis Millais

a. sandayensis Millais

b. westrse Miller

7. sarnius Miller .

8. cabrerge Thomas

9. dentatus Miller.

10. hartingi Barrett-Hamilton

11. angularis Miller

12. ratticeps Keyserling and
Blasius .

Sub-Genus Chionomys Iililler

13. nivalis Martins

a. nivalis Martins .

b. aquitanius Miller

14. lebrunii Crespon .

a. lebrunii Crespon.

b. leucurus Gerbe .

15. ulpius Miller .
5. Arvicola Lac^pede

1. amphibius Linnaeus

a. amphibius Linnaeus

b. reta Miller

2. sapidus Miller .

a. sapidus JMiller

b. tenebricus Miller

3. terrestris Linnaus.

4. italicus Savi.

5. illyricus Barrett-Hamilton

6. musignani de Selys-Long-

cbamps

7. scherman Shaw

a. scherman Shaw .

b. exitus Miller .

c. monticola de
Lougchamps .

Selys-

658
659
662
668
669
671

672
673
675
680
681
683
686

686
687

690
693

694
696
697
698
700
701
703
704
706

708
712
713
716
717
718
719
722
723
723
725
730
732
732
733
735
738
740
741

744
744
745
746

749

6. Pitymys MclMurtrie .

1. subterraneus de Selys-

Longchamps ....

a. subterraneus de Selys-
Longchamps ....

b. capucinus Miller

2. dacius ^Miller ....

3. drueutius Jliller

4. fatioi Mottaz ....

5. multiplex Fatio

6. savii de Selys-Longchamps

7. nebrodensis Mina-Palumbo

8. pyrenaicus de Selys-Long-

champs

a. pyrenaicus de Selys-
Longchamps ....

b. brunneus Miller .

9. planiceps Miller .

10. gcrbii Gerbe ....

11. lusitanicus Gerbe.

12. marite Major ....

13. pelandonius MiUer .

14. depressus Miller .

15. ibericus Gerbe

a. ibericus Gerbe .

b. centralis Miller .

c. pascuus Miller .

d. regnlus Miller

16. duodecimcostatus de

Selys-Longchamps

17. provincialis Miller

18. thomasi Barrett-Hamil-
ton

19. atticus Miller ....

Sub-Family IMurin.e ....

Apodemus Kaup ....

1. epimelas Nehring .

2. sylvaticus Linnaeus

a. sylvaticus Liunaus .

b. callipides Cabrera

c. dichrurus Rafinesque .

d. creticus Miller .

3. hebridensis de Winton

4. hirtensis Barrett-Hamil-

ton

5. fridariensis Kinuear .

6. liavicollis jNIelchior

a. flavicoUis Melchior .

b. wiutoni Barrett-Hamil-

ton

7. agrarius Pallas . . . .
. Micromys Dehne . . . .

1. minutus Pallas . . . .

a. soricinus Hermann .

b. xjratensis Ockskay
. Epimys Trouessart .

1. rattus Linnteus . . . .

a. rattus Linnaeus .

b. alexandrinus Geoft'roy .

2. uorvegicus Ersleben .

PAGE

752

755

758
760
760
762
763
764
768
770

770

771
772
772
773
776
777
778
779
780
782
782
783
784

784
785

786
787

791
791
794
797
803
809
810
818
824

825
825
828
829

831
836
840
841
844
846
848
849
853
854
858

SYSTEMATIC INDEX

Order Rodentia — Sub - Order
SiMPLiciDENTATA — Family
MuBiD.E — Sub-Family Mu-
KiNyE — continued.

4. Mus LimisBus ....

1. musculus Linua3us
fl. musculus Linna3US .
b. aiioricus Schinz .

2. muralis Barrett-Hamilton

3. fteroensis Clarke

4. spicilegus Pet6nyi .

a. spicilegus Petlnyi .

b. hispanious Miller

c. lusitanicus Miller

5. Acomys Geoffrey .
1. miiious Bate

Family SPALACID^ .
1. Spalax Gueldenstaedt

1. dolbrogeae Miller .

2. hungaricus Nehring

3. greecus Nehring

Family SCIURID^ . .

1. Sciurus Linnaeus .

1. vulgaris Linnajus .

a. vulgaris Linnaeus

b. varius Gmelin

c. leucourus Kerr .

d. russus Miller.

e. fuscoater Altum .
/. italicus Bonaparte
g. lilajus Miller .

h. alpinus Desmarest
i. numantius Miller
j. infuscatus Cabrera
A;, segurse Miller
I. bffiticus Cabrera .

2. CitcUus Oken . . .

1. citellus Linuajus .

2. suslica Gueldenstaedt

3. Marmota Blumenbach

1. marmota Linnoeus .

2. bobak Miiller . .

Family PETAURISTID^

1. Sciuropterus F. Cuvier

1. russicus Tiedemann

Family CASTORID^

1. Castor Linnaeus .

1. fiber Linnaeus .

863
865

869
871
874
875
877
878
879
882
883
883

887
887
889
894
895

897
898
898
905
906
907
909
910
912
913
914
914
916
917
923
924
924
929
931
932
937

940
941
941

947
947
947

ORDER UNGULATA

page

Family SUID^ 956

1. Sus Linnteus 956

1. scrofa Linnaeus. . . . 957
[attila Thomas] .... 960

2. meridionalis Major . . 960

Family CERVID^ .... 962

1. Cervus Liunteus .... 963
1. elaphus Linnaeus . . . 964

a. germanicus Desmarest . 965

b. elaphus Linnaeus . . 967

c. atlanticus Lonnberg . 967

d. scoticus Lonnberg . . 968

e. hispanicus Hilzheimer . 969
/. corsicanus Erxleben. . 969

2. Dama Hamilton Smith . . 970
1. dama Linuffius .... 970

3. Capreolus Gray .... 972
1. capreolus Linnaeus. . . 973

a. capreolus Linnaeus . . 974

b. transsylvanicus Matschie 975

c. canus Miller .... 975

d. thotti Lonnberg. . . 975

4. Alces Gray 976

1. alces Linnaius .... 978

5. Rangifer Hamilton Smith . 979

1. tarandus Linnaeus . . . 980

2. fennicus Lonnberg. . . 981

3. platyrhynchus Vrolik . . 985

Family BOVID.E 986

1. Ovis Linnaeus .... 986
1. musimon Pallas . . . 987

2. Capra Linnaeus .... 988

1. ibex Linnffius .... 989

2. pyrenaica Schinz . . . 989

a. pyrenaica Schinz . . 1)90

b. liisitanica Franca . . 991

c. victoriae Cabrera. . . ',)91

d. hispanica Schimper. . 991

3. segagrus Erxleben . . 992

3. Rupricapra BlaiuviUe . . 992

1. rupicapra Linnasus . . 993

2. ornata Neumann . . . 994

3. pyrenaica Bonaparte . . 995

4. parva Cabrera .... 995

APPENDIX.
Crocidura ichnusas Festa . . . 998
Evotomys glareolus norvegicus —

cranial measurements . . . 999
Spalax — Forms recognized by

M6hely. . 1000

CATALOGUE

OP THE

LAND-MAMMALS OF WESTERN
EUROPE.

Order INSECTIVORA.

1827. Insectivora Gray, Griffith's Cuvier, Aoim. Kingd., v, -p. 100.

Gcoijraphical distribution. — Africa (iucluding Madagascar),
Europe, Asia (including the Malay Archipelago), North America,
^^ Greater Antilles, and extreme north-western portion of South
America.

Characters. — Terrestrial, non volant, placental mammals with
low development of brain, the cerebral hemispheres without
convolutions ; teeth of a primitive or modified tuberculo-sectorial
type, the postei'ior upper premolar and anterior lower molar
never specially modified as carnassials.

Bemarhs. — The mammals of this order present such diversity
of form and structure that it is difficult to frame any definition
by which all members of the group may be invariably recognized
by skeletal or external characters. Ten families of Insectivora
are currently recognized, three of them occurring in Europe.

KEY TO THE EUROPEAN FAMILIES OP INSECTIVORA.

Dentition of a semi-crushing type : first and second
upper molars with four subequal cusps, their styles
and commissures rudimentary ; sutures in skull
persistent ; a large external pterygoid plate ; zygo-
matic arch complete, heavy. (In European species

back covered with spines.) (Hedgehogs) Erinaceidie, p. 114.

Dentition of a strictly sectorial type : first and second
upper molars with three or four cusps strongly
contrasted in size, their styles and commissures
highly developed and forming an important func-
tional part of tooth ; sutures in skull mostly dis-
appearing early in life ; no external pterygoid
plate ; zygomatic arch slender or incomplete. (Back
covered with soft, dense fur.)
Anterior lower incisor greatly elongated in axis of jaw;

mandibular articulation double ; zygoma absent ;

floor of brain-case with largo lateral vacuities ;

no auditory bulla; general form mouse-like, the

neck evident ; external ear present (Shrews) Soricidie, p. 28.

B

-a INSECTIVOEA

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

Desmans) Talpidie, p. 2.

Teeth in front of molars sharply differentiated by
form into incisors, canines and premolars, the
upper incisors small, sub-equal ; front feet
highly modified for burrowing, the palms
everted ; tail scarcely as long as head (Moles) Talpinx, p. :2.
Teeth in front of molars not differentiated by
form into incisors, canines and premolars, the
inner upper incisor greatly enlarged, vertical,
trenchant ; front feet not modified for burrow-
ing (habits aquatic), the palms in normal
position ; tail (in European members of the
group) longer than head and body (Desmans) Desmauinie, p. 20.

Family TALPID^.

1826. Talpidx Gray, Thomson's Annals of Philosophy, xxvi, p. 339.

Geographical (listribuiion. — North temperate portions of Old
and New "Worlds ; in Europe south to the Mediterranean coast
and west to England.

Characters. — Skull long and narrow, strongly tapering an-
teriorly, most of its sutures disappearing early in life ; zygomatic
arch complete, slender ; floor of brain-case completely ossified ;
tympanic bone attached to skull, forming a ilattened bulla ;
mandible with single articulation, the glenoid surface normal ;
no external pterygoid plate ; crowns of upper molars low, much
narrower internally than externally, the paracone and metacone
near middle of crown, the commissures and styles well developed
and forming with corresponding portions of lower teeth an
effective cutting apparatus ; body heavy, cylindrical, the short
neck concealed between the greatly developed shoulders ; eye
minute, often covered by the integument : snout much elongated,
tei'ete or depressed ; no external ear.

Beinarhs. — The members of the family Talpidie are at once
recognizable among European mammals by the great development
of the shoulder girdle and apparent absence of neck, the auditor}-
orifice seeming to lie at the shoulder. Though excessively modified
in general form the Talpidse are much less specialized than the
Soricidse in the more fundamental characters of skull and teeth.
The family is divisible into several very distinct groups or sub-
families, two of which are represented in Europe. In one of
these the animals are specially adapted to subterranean habits :
in the other they are modified for aquatic life.

Sub-Family TALPIN.-E.

Geot/raphical distribution. — Temperate portions of Europe and
Asia, from England to Jajian : in Europe south to the Meditei-
ranean coast.

Characters. — Teeth in front of molars sharply differentiated
by form into incisors, canines, and premolars, the incisors, both
above and below, small, sub-equal, chisel-shaped, the upjaer
canine large, strongly trenchant ; external form highly moditied
for subterranean life, the greatly enlarged orbicular front feet
with palms permanently turned outward, the hind feet much
smaller, not peculiar in form ; tail short ; muzzle terete.

Remarks. — This group, comjwsed of the true moles of the
Old World, and specially characterized by the relatively primi-
tive condition of the anterior teeth, is represented by four or five
genera, one of which occurs in Europe.

Genus TALPA Linn;eus.
1758. Talpa Linnaeus, Sysfc. Nat., i, 10th ed., p. 52.

Type species. — Talpa europsea Linnteus.

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

Characters. — Dental formula: i 3Z3, CjEti, pm xEr^, »h 3=3 = 44.
Up23er premolars small, distinctly spaced, showing no tendency
to become imbricated. Lower canine slightly but evidently
differentiated from incisors in size and form ; auditory bulla very
slightl}^ inflated, its outlines usually indistinct, the meatus small,
sub-circular ; external form strictly talpine ; ear-conch absent ;
eye minute, often covered by the skin.

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

KEY TO THE EUROPEAN SPECIES OF TALPA.

Greatest diameter of «i' about 4 mm. ; three lower
molars together about 8 mm. (vicinitj' of

Rome) T. romana, p. 18.

Greatest diameter of 5k' about 3 mm. ; three lower

molars together about 7 mm. or less.
. Condylobasal length of skull 33 to 37 mm. (dis-
tribution general) T. curopxa, p. 3.

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

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

Posterior border of anteorbital foramen over

middle of .second molar (Iberian) T. occidentalis, p. 15.

TALPA EUROP^A Linnaeus.

1758. [Talpa] europxa Linnseus, Syst. Nat., i, 10th ed., p. 52 (Sweden).
17G6. Taljici /n',sn(.s- P. L. S. Miiller, Natursyst. Suppl. u. Regist.-Band,

p. 36 (Ostfriesland).
1772. [Talpa] caudata Boddaert, Kortbegrip van het zaraenstel der Natuur,

1, p. 50 (Renaming of europxa).

B 2

4: INSECTIVORA

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

p. 117 (Ostfriesland).
1785. [Talpa] vulgaris Boddaert, Elenohus Anim., i, p. 126 (Europe).
1789. [Taliya europsea] 0 variegata Gmelin, Syst. Nat., i, 13th ed., p. 110

(Sweden).
1789. [Talpa europma'\ y alba Gmelin, Syst. Nat., i, 13tli ed., p. 110

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

Germany) .
1792. Talpa etirop[iea] nigra Kerr, Anim. Kingd., p. 200 (Renaming of

euro2)iea).
1797. Talpa curopxa riifa B[orkhause]n, Der Zoologe (Compendiose Biblio-

thek gemeinntitzigsten Kenntnisse fiir aUe Stande, pt. xxi),

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

Siiugeth. des In- und Auslandes, fig. 473 (Saxony).
1852. Talpia europsea albida Reichenbach, Vollstiindigste Naturgesch. des

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

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

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

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

(Renaming of albo-maculata).
1869. Talpa europxa 3 grisea Fitzinger, Sitzunsgber. kais. Akad. Wissensch.

Wien, JNIath.-Naturwiss. Classe, lix, pt. i, p. 403 (synonym of

cinerea wrongly attributed to Zimmermann, Geogr. Gesch., ii,

p. 390, 1780, where vernacular name only is used).
1897. ITaliM] scalops Schulze, Abh. u. Vortr. Gesammtb. Naturw. iv,

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

Type locality. — Upsala, Sweden.

Geographical distribution. — Europe from Great Britain and
the Channel Islands eastward, and from the Mediterranean coast
to Scotland and central Sweden.

Diagnosis. — Size medium (hind foot about 18 mm., condylo-
basal length of skull rarely less than 33 mm.) ; skull with orbit
rather long, so that distance from posterior border of orbit to
posterior border of anteorbitai foramen is about equal to that
from latter point to front of hist j^remolar or back of canine ;
teeth of moderate size, the length of upper tooth-row (exclusive
of incisors) less than 14 mm., the greatest diameter of wi^ about
3 mm. ; mesostyle of m- and m^ entire or with apex slightly
notched.

E.itrrnal rliararters. — General form highly moditied for sub-
terranean burrowing habits, the neck so short that the conical
head with much produced snout ajapears to be joined directly,
between the very large, everted front feet, to broad, powerful
shoulders, behind which the unusually long, nearly cylindrical
body tapers gradually to somewhat weak and narrow pelvic
region. Fur dense and velvety, the hairs all of the same length,

about 12 mm. long throughout body, shorter on head. Muzzle
pad well developed, slightly wider than high, its entire margin
free, the upper edge folded back and with a deep nari-ow median
emargination, the whole surface finely and evenly rugose.
Behind pad the skin of the muzzle is naked and wrinkled along
median line above, the exact size and form of the naked area
varying in different individuals, but its length usually about
7 mm., its breadth anteriorly equal to greatest diameter of pad,
that posteriorly somewhat less. Under side of upper lip with
deep median groove extending forward to muzzle-pad. Eyes
excessively minute, less than 1 mm. in diameter, in some
individuals completely covered by the integument, in others with
a pin-hole aperture.* No external ear, the meatus about 2 • 5 mm.
in diameter. Legs so short that scarcely more than the feet
project beyond general integument of body. This is particularly
true of the front legs, which are entirely concealed to wrist.
Front feet very large, the palms orbicular, permanently turned
outward, their surface naked, finely and evenly tuberculo-reticu-
late, without trace of pads. Toes, five, each armed with a long,
slightly-curved nail broadly grooved along under surface, that of
fifth digit not so large as the others. Third digit longest, second
and fourth sub-equal and slightly shorter, first and fifth slightly
shorter than second and fourth. Hind foot short, somewhat
triangular in outline, broad through base of toes but narrowing
rapidly toward heel, the five toes with well developed, curved,
but not flattened claws, much less enlarged than those of front
foot ; second, third and fourth digits sub-equal and longest, fifth
and first successively shorter, soles naked, reticulate, with
five rudimentary tubercles at bases of digits, and a fifth neai-
middle of inner margin, its anterior extremity projecting so as to
suggest a supplemental (clawless) toe. Dorsum of pes thinly
clothed, the hairs at its edge forming a slight fringe. Tail
thickened and fleshy, about 1^ times as long as hind foot,
subterete but somewhat compressed, much constricted basally ;
scales arranged in rather irregular rings, of which there are
about twelve to the centimeter at middle ; hairs of tail sparse,
not concealing rings, sometimes forming a thin pencil. Mamma? :
/> 2 - 2 ; i2-2 = 8.

Colour. — Fur everywhere dark slaty grey, sometimes almost
l>lackish, at others more neai'ly a dark smoke-grey, the hairs
every wliere with a noticeable metallic or purplish iridescence ;
underparts sometimes a little less dark than back, and occa-
sionally with a yellowish brown sufl'usion : tail concolor with body.

Skull. — The skull is long and narrow, tapering gradually
forward from middle of brain-case, the widest region, to just
behind canines, the extreme tip of rostrum widening a little, the
zygomatic arches not standing out beyond general contour.

* JTuoh confusion has been caused by the supposition that the presence
or absence of this minute aperture is a specific or racial character.

IXSECTIVOKA

Outline when viewed from the side long wedge-shaped, rounded
oti' posteriori". Surface of skull smooth, except for a slight
sagittal crest, present in old individuals along course of parietal
suture, and a crest-like elevation extending ftirward and outward
from autero-external angle of interparietal along edge of slightly
intlated mastoid region, and terminating anteriorly in a slightly
jxiinted in'ojection. Brain-case with posterior mai'gin nearly semi-
circular to projecting points already mentioned, then abruptly
conical to back of interorbital region, its outline when viewed
from behind fusiform, a little less than half as deep as wide :
condyles slightlv projecting, but not noticeably breaking general
posterior outline. Interparietal large, strap-shaped, slightly
convex in front, similarly concave behind, the lateral extremities

squarely truncate, its antero-pos-
terior diameter about one-third
transverse diameter. Its anterior
and lateral sutures remain visible,
but the lambdoid suture is nearly
obliterated early in life, though
its position is usually marked by
a slight ridge representing the
lamlxloid crest. Base of brain-
ca.se smooth, without conspicuous
ridges, furrows or open spaces,
the bones for the most part some-
what inHated : two ill-tielined pits
in surface of basioccipital in front
of foramen magnmu ; a shallow,
broadlv triangular median furrow
between the low, flattened bulke,
the sub-circular, slightly triangu-
lar outline of which is sometimes
distinctly indicated, but more
often very obscui-e : auditory
meatus small, nearly circular.
Iinier pterygoid plate small but well developed, the hamulars
short, turned slightly outward : mesopterygoid space much
narrower posteriorly than anteriorly, its greatest breadth less
than half length, its length much more than width of palate
bctN\ ccn posterior molars, its anterior border broadly rounded :
region outside pterygoid plate inflated and with low but distinct
longitudinal ridge, convergent anteriorly with that forming edge
of mesopterygoid space. Interorbital region sulvcylindrical, dis-
tinctly expanded at middle. Rostrum narrower and somewhat
abruptly lower than interorbital region, its narrowest point just
behind canines : nares with evident posterior emargiuation.
Anteoi-bital foramen moderately large, its jxisterior border OAcr
metastyle of */('-, the plate forming outer wall of canal usually
nnich nari-ower than foramen Lachrymal foramen above anteor-

YW. 1.
Titlpii eur,tpK.t. y-.n. size.

r

bital foramen and slightly in front of its middle ; its orifice over
metastj'le of j/ji. Temporal fossa rather large, the distance from
its posterior' border to posterior border of anteorbital foramen
equal to that from latter point to front of first premolar or back
of canine. Palate moderately broad, without special peculiarities
of form, terminating posteriorly, a little behind tifi, in a slightly
raised crescentic ridge ; a vacuity about as large as the minute
incisi\e foramen on each side of palate opposite space between
//(' and »;«'-. Mandible rather slender, the ramus curved downward
near middle and upward posteriorly, the angular process nearly
on level with alveolar line. Coronoid process large, its broadly
rounded off extremity rising considerably above articular level,
its height above alveolar line about eijual to least breadth of
l)Osterior segment of mandible. Articular process slender, the
single articular surface small, normal in position. Angular
process slightly longer than articular process, and distinctly
broader, forming an obliquely-set plate directed almost horizon-
tally backward, the extremity slightly hooked upward.

Ti^eth.—In proportion to the size of the skull the teeth are
moderately large, their general
asjject noticeably trenchant. Upper
incisors simple, chisel - shaped,
IXirpendicular, forming a strongly
convex row between canines, their
height equal, but their breadth
diminishing regularly from first to
third ; crowns of first and second
usually in contact, that of third
separated by a slight space from
second and from canine.
Lower incisors projecting
somewhat obli(juelv for-
ward, essentially similar to
the upjier teeth in form,
but smaller and narrower.
Upper canine large, two-
rooted,* the height of the
shaft greater than that of an v
of the othei' teeth, and fully
e(jual to anterior breadth of
palate ; shaft wider ante-
riorly than posteriorly, with
anter-o-internal longitudinal
groo^•e, and highly deve-
loped, slightly concave pos-
terior cuttirrg edge. Lower canine very small, resembling a
fourth incisor, but with shaft conical instead of chisel-shaped,

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

h'lO. 2.
Talpa europtea. Teeth x o.

INSECTIVORA

and set in the jaw at a slightly different angle posterior
surface of shaft with well developed longitudinal ridge. First,
second and third upper premolars two-rooted, small, their
points on level with those of incisors and inner cusps of
molars, their crowns separated from each other as well
as from canine and large premolar by narrow equal spaces ;
crowns rather higher than long, compressed, with slightly
developed posterior cutting edge, narrowly triangular in outline
when viewed from the side, the first more slender than the
others, the second and third with slight though evident postero-
external angle on cutting edge. Fourth premolar essentially
like the other three in form, but much larger and three-rooted,
the length of crown greater in proportion to height ; no secondary
cusps. Lower premolars two-rooted, similar to the upper teeth
in form, the second and third the same size as smaller upper
premolars, the first nearly as large as pm*, but narrower,
owing to absence of third root, the fourth intermediate. Upper
molars with crowns much wider externally than internally,
and completely divided into two unequal sections by a deep
longitudinal groove jjassing between protocone and bases of
paracone and metacone. Protocone large, its posterior com-
missure extending in line parallel to sagittal plane, and ending
abruptly at posterior edge of crown, near which it is slightly
thickened, its anterior commissure similar though shorter, but
usually showing some trace of thickening,* particularly in m-.
Paracone and metacone sub-equal in iir and vi^, the metacone
the larger in the former, the paracone in the latter. In «<' the
metacone is about doulile the size of paracone, and is the
largest and highest cusp of the upper molar series, its posterior
cutting edge and long commissure functioning with similarly
enlarged protoconid of m.,. Styles and outer commissures well
developed in m- and forming a distinct VV-p^-ttern ; mesostyle
entire or with apex slightly notched. In m^ the parastyle is
reduced to a minute though usually evident cusplet on the
cingulum., and the mesostyle to a thickening or angle in
commissure connecting the two main cusps ; metastyle well
developed.f In m^ the parastyle, mesostyle and their commis-
sures are well developed, metastyle and fourth commissure
absent ; mesostyle with apex usually bifid. Middle lower molar
largest, its protoconid the highest cusp in the series. First lower
molar broader posteriorly than anteriorlj-, second and third
slightly broader anteriorly than posteriorly, the two triangles
essentially alike in form. Metaconid of 7/*i low, scarcely more

* The thickenings vary considerably iu different individuals. Occasion-
ally they arc obsolete, but more frequently they are so well developed as
to form an evident protoconule and metaconule, the latter always the
larger of the two.

t In this tooth there is no anterior y, the outer surface of paracone
essentially resembling that of ^ww' except for its smaller size.

than a slight thickening of the cingulum. In the other teeth
it is a well developed cusp nearly equal to entoconid. In all
three teeth the outer cusps are noticeably higher than those of
the inner row.

Menmrements. — Average and extremes of five males from
Borrohol, Sutherland, Scotland: head and body, 148 "4 (145-
152); tail, 26-8 (26-28); hind foot, 18-8 (18-19). Average
and extremes of four females from the same locality : head and
body, 135 (133-138) ; tail, 25-3 (24-28) ; hind foot, 17-5 (17-18).
Average and extremes of five males from Solferino, Landes,
France : head and body, 138 (134-142) ; tail, 26-2 (25-28) ; hind
foot, 17-8 (17-18). Five females from the same locality: head
and bodjs 128-6 (126-133) ; tail, 27-4 (25-29) ; hind foot, 17-2
(17-18). Average and extremes of six males from Lucinges,
Haute-Savoie, France: head and body, 136-6 (132-140); tail,
24-8 (23-26); hind foot, 18-1 (17-19). Average and extremes
of three males from Turin, Italy : head and body, 138-3 (123-
147) ; tail, 31 (27-34) ; hind foot, 19-2 (18-20). Average and
extremes of eight males from the Dehesa de Valencia, Spain :
head and body, 144-3 (135-165) ; tail, 26-6 (26-28); hind foot,
18-1 (18-19). For cranial measurements see Table, p. 12.

Specimens examined. — Three hundred and ninety, from the following
localities : —

Scotland: Borrohol, Sutherland, 9 (Wilson); Black Isle, Cromarty, ;3;
Cromarty, 1 ; Gordonstown, Elgin, 1 ; Grantown-on-Sisey, Elgin, 13 (Wilson) ;
Cortach)-, Forfar, 1 (Wilson) ; Stockbriggs, Lanarkshire, 1.

England : Bowdon, Cheshire, 1 ; Altriucham, Chester, Cheshire, 1 ;
Parsop, Hereford, 1 ; Lavenham, Suffolk, 1 (Wilson) ; Barrow, Suffolk, 1
(U.S.N.M.); Arley, Staffordshire, 1 (Wilson); Rugby, Warwickshire, 2;
Warwickshire, no exact locality, 2 ; Pulbourn, Cambridge, 1 ; HoUoway,
Somersetshire, 2; Somersetshire, no exact locality, 1 ; Banstead, Surrey, 2 ;
Coonibe, Surrey, 1; Cobham, Surrey, 2; Egham, Surrey, 2; Knockholt,
Kent, 2; Bromley, Kent, 2; Devonshire, no exact locality, :3.

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

Belgium: Waremrae, Li6ge, 3 (U.S.N.M.).

France : Guines, Pas-de-Calais, 1 ; Pont-de-Briques, Pas-de-Calais, 1
Trinity, Jersey, 2 ; St. Lawrence, Jersey, 1 ; Barbizon, Seine-et-Marne, 2
Melun, Seine-et-Marne, 1 (Mottaz) ; Lignieres, Charente, 1 (Mottaz)
Huelgoat, Brittany, 1 ; Cadillac-sur-Garouue, Gironde, 1 (U.S.N.M.)
Foret de Bouconne, Gers, 13 ; Solferino, Landes, 10 ; Calerille, Haute-
Garonne, 10; Legouvin, Haute-Garoune, 8; Luchon, Haute-Garonne, 3;
Bareges, Hautes-Pyr6n6es, 5; I'Hospitalet, Ariege, 1: Port6, Pyreuees-
Orientales, 3 ; St. Gilles, Gard, 1 ; Valescure, Var, 1 ; Agay, Var, 1 ;
Etupes, Doubs, 3 (Mottaz) ; Lucinges, Haute-Savoie, 9 ; Montaubau,
Haute-Savoie, 15 ; Cranves-Sales, Haute-Savoie, 12.

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

Gerjuany : Konigsberg, 4 (U.S.N.IM.) ; Moritzburg, Saxony, 8 (U.S.N.M) ;
Ummerstadt, Thiiiingen, 3; Brunswick, 15 (U.S.N.M.); Aachen, 9
(U.S.N.M.); Bremen, 1 ; Ingelheim, Rheinhessen, 4; Strass, near Burg-
heim, Bavaria, 7; Niesky, Silesia, 1; Kalbe, Saale, 2; Magdeburg,
Saxony, 1 ; Strassburg, 3.

.VusTRi A-HuNGARY : Csallokoz-Somorja, Pressburg, Hungary, 2 ; Hatszeg,
Huuyad, Transylvania, 2.

10

INSECTIVORA

RouMANiA : Bustenari, Prahova, 1 ; Comana, Vlasca, 1 ; Bucharest, 1
(Genoa) .

Bulgaria: Sofia, 1 (Andersen).

Switzerland : Geneva, 15 (U.S.N.M. and Mottaz) ; Lausanne, Vaud, 1
(U.S.N.M.) ; Les Plans, Vaud, 2 (U.S.N.M.) ; Chesieres, Vaud, 1 (Mottaz)
Andermatt, Uri, 2 (U.S.N.M.) ; Miirreu, 1 ; Tliurgau, 1 ; Oberhasli VaUey
1 ; St. Galleu, 3 ; Rheinthal, St. Gallen, 2; Degersheim, St. Gallen, 3
Gossan, St. Gallen, 3; Untervatz. Grisons, 1; Breganzona, Ticino, 1
Cortivallo, Ticino, 6 (B.M. and U.S.N.M.) ; Comano, Ticino, 2 (U.S.N.M.)
Lugano, Ticino, 4 (B.M. and U.S.N.M.); Muzzano, Ticino, 2 (B.M. and
U.S.N.M.) ; Sorengo, 1 (U.S.N.M.) ; Cremignone, Ticino, 1.

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

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

2 6. Black Isle, Cromarty, Scot-
land.
9. Gordonstowu, Elgin.

6. Stockbriggs, Lanarkshire.

9. Parsop, Herefordshire, Eng-
land.

2 6. Rugby, Warwickshire.

2. Warwickshire.

6. Fulbourn, Cambridgeshire.

2 al. Holloway, Somerset.

2. Somerset. [Hiigcl.)

2 al. Cobham, Surrey.

1 al. 1. Eghani, Surrey.

?. Knockholt, Kent.

2 (J.al. Bromley, Kent.

3 6. Devonshire.

2 6. Trinity, Jersey, Channel
Islands. (/?. H. Bunting.)
?. St. Lawrence, Jersey.
6. Guines, Pas-de-Calais, 80 ft.

France.
i. Pont - de - Briques, Pas - de-
Calais.
6. Huelgoat, Brittany, 600 ft.
2 6,29. Foret de Bouconue, Gers,

2.50 m. {A. Robert.)
2 6,29. Solferino, Landes. (.4. Fw-

hrrt.)
2 6,2 9. Caterille, Haute - Garonne,
9U0-1000in. {A. Bobcrr.)
2 6, 9. Luchon, Haute - Garonne.
(A. Bohcrt.)
2 6,19. Bareges, Hautes-Pyr6nees.
c5. L'Hospitalet, Ariege. (A.

Bobert.)
6. Port6, Pyren6es-0rientales.
2 6 Porte, Pyren6es - Orientales,
1600-1700 m. (.1. Bobert.)
6. Valescure, Var.

W. R. Ogilvie-

Grant (p).
W. R. Ogilvie-

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

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

H. King (p).

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

F. Heiss (p).

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

0. Thomas (p).

Mrs. Power (p).
O. Thomas (c&p).

11. 1. 3. 62-G3.

11. 1. 3. 66.

79. 9. 25. 3.
11. 1. 3. 67.

11. 1. 3. 64-65.

7. 1. 1. 17-18.
11. 1. 3. 71.
58. 4. 22. 1-2.
79. 9. 25. 5-6.

58. 1. 2. 1-2.
11. 1. 3. 61.
81. 4. 2. 1-2.
11. 1. 3. 68-70.

8. 9. 2. 3-4.

8. 12. 17. 1.
94, 6. 6. 19.

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

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

0. Thomas (p).

0. Thomas (p).

O. Thomas (p.)

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

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

92. 9. 5. 1.
G. 4. 1. 28-31.

6. 4. 1. 32-.35.

6. 4. 1. 24-27.

G. 4. 1. 21-23.

8. 8. 4. 129-132.
8. 9. 1. 38.

8. 8. 4. 133.
8. 9. 1. 36-37.

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

TALPA 1 1

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

1. Montauban, Haute - Savoie, A. Eobort (c & p). 97.1.9.1.

1000 m.

3 <i, 1 9. Cranves-Sales, Haute-Savoie, A. Eobert (c & p). 5. 4. 4. 1. 5. 4,

900 m. 9. 3-5.

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

909-1200 m. France. [A.
Robert.)
6, 9. Pajares, Leon, Spain. {N. O. Thomas (p). 8. 2. 9. 26-27.
Gomalez.)

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

Spain.
2 (J, 5 9. Gastrillo de la Reina, Burgos. N. Gonzalez (c.) 8.7.7.1-7.

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

6(J, 6 9. Barraoas, Castellon. {N.Gon- 0. Thomas (p). 8.2.9.29-40.

zalez.)
3 £, 9. Gatarroja, Valencia. {N. Gon- O. Thomas (p). 8. 2. 9. 12-15.
zalez.)

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

encia. (N. Gonzalez.)
3 6, 9. Ingelheim,Rheinhessen,Ger- C. Hilgert (c). 8. 11. 2. 11-14.

many.
(J, 9. Ummerstadt, Thiiringeu. Lord Lilford (p). 11.1.1.115-116.
[Schuchardt.)
6. Ummerstadt, Thiiringen. Lord Lilford (p). 11.1.1.137.
(Schuchardt.)
6,S9. Strass, Burgheim, Bavaria. Lord Lilford (p). 11.1.1.1-4.

(Korbitz.)
2 <i, 9. Strass, Burgheim, Bavaria. Lord Lilford (p). 11.1.1.132-134.
{Kiirbifz.)
6. Niesky, Silesia. (W. Bacr.) Dr. E. Hamilton 97.12.4.18.

9. Kalbe, Saale. (TT'. Bauer.) Lord Lilford (p). 11. 1. 1. 5.
6. Kalbe, Saale. {W. Bauer.) Lord Lilford (p). 11.1.1.136.
i. Magdeburg, Saxony. (IloZ^er- Lord Lilford (p). 11.1.1.135.

storff.)
6, 9. Strassburg, Alsace. (C. Mot- 0. Thomas (p). 8. 8. 10. 10-11.

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

Hungary. (p).

2 6. Hatszeg, Hunyad, Transyl- C. G. Dauford (c). 3.2.2.20-21.

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

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

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

(£. H. Zollikofer.)
1. Oberhasli Valley, Switzerland. Tomes Collection. 7.1.1.130.

(Kocscrma7i.)
2 6,9 St. Gallen, 650 m. Switzer- O. Thomas (p). 4. 4. 5. 23-25.

land. {E. H. Zollikofer).
6. Breganzona, Ticino, Switzer- 0. Thomas (p). 2. 8. 4. 17.

land. (£■. H. Zollikofer).
6. Cortivallo, Ticino. (E. H. O. Thomas (p). 2. 8. 4. 15.

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

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

Zollikofer).
9. Cremignono, Ticino. (E. H. O. Thomas (p). 2. 8. 4. 20.

Zollikofer).

12

INSECTIVOKA

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14

INSECTIVORA

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15

TALPA C.ECA Savi.

1822. Talpa cxca Savi, Nuovo Gioru. de' Letterati, Pisa, i, p. 265.
1857. Talpa csaca Blasius, Siiugethiere Deutschlands, p. 115.

1906. Talpa cxca Camerano, Boll. Mus. Zool. ed Anat. Comp. della K.

Univ. di Torino, xxi, No. 530, p. 1, June 22, 1906.
1910. Talpa cxca Trouessart, Faune INIamm. d'Europe, p. 63.

Type locality. — Mountains near Pisa, Italy.

Geographical distribution. — Mediterranean region, eastward
into Asia Minor ; details of distribution not known.

Diagnosis. — Smaller than Talpa europwa (condylobasal length
of skull, 29-32 mm. instead of 33-37 mm.) and .skull with
narrower rostrum and palate (breadth of rostrum over canines
about 4 mm. instead of 4 • 6-5 mm.) ; temporal fossa shortened, the
distance from its posterior margin to posterior margin of anteor-
bital foramen about equal to distance from latter point to second
premolar ; anteorbital foramen large, its posterior border over
front of last molar, the plate forming outer wall of canal a
slender, terete thread ; teeth small, the length of upper tooth-row
(exclusive of incisors), 11-12 mm. ; mesostyle of m- and m^ with
apex deeply notched (this character not visible in specimens with
much worn teeth).

Measurements. — Average and extremes of five specimens from
Cortivallo, Ticino, Switzerland : head and body, 133 • 8 (125-142) ;
tail, 25-2 (21-30) ; hind foot (dry), 15-6 (15-16). Average and
extremes of three specimens from Reggello, Tuscany, Italv (in
alcohol) : tail, 23-6 (22-25) ; hind foot, 14-7 (14 -4-15 -5). " For
cranial measurements see Table, p. 17.

Specimens examined. — Twenty-five, from the following localities : —
Switzerland : Agmizzo, Ticino, 1 ; Bt-llinzona, Ticino, 1 (U.S.N.M.) ;
Breganzona, Ticino, 1 (U.S.N.M.) ; Cauabbio, Ticino, 1 (U.S.N.M.) ; Corti-
vallo, Ticino, 5 (U.S.N.M.); Lonvico, Ticino, 2 (U.S.N.M.); Lugano,
Ticino, 3 (U.S.N.M.) ; Muzzano, Ticino, 1 (U.S.N.M.) ; Origlio, Ticino 1
(U.S.N.M.) ; Sorengo, Ticino, 1 (U.S.N.M.) ; Stabio, Ticino, 1.

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

6. Agmizzo, Ticino, Switzerland. 0. Thomas (p). 2. S 4 18

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

likofvr.)
c5, 9 al. Italy.' Dr. Riippell (c). 45.7.22.32-3.3.

TALPA OCCIDENTALIS Cabrera.

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

XX, p. 212, September, 1907. Type in Cabrera collection.
1907. Talpa cxca occidentalis Cabrera, Bol. Real. Soc. Espafi. Hist. Nat.,

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

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

Type locality. — La Granja, Province of Segovia, Spain.
Geographical distrihtiiion. — Iberian Peninsula.

16

IXSECTIVOKA

Ding)tosis. — Like Talpa cseca but skull rathtn- I'obust, the
breadth of rostrum over roots of canines usually more than
4 mm. ; anteorbital foramen contracted, its posterior border ovei'
mesostyle of vi'^, the plate forming outer wall of canal usually as
wide as transverse diameter of foramen ; crowns of molars
appreciably enlarged (length of upper tooth-row, exclusive of
incisors, frequently more than 13 mm.), but dentition otherwise
as in Tnlpa csern.

Skull. — The skull is somewhat larger and more robust than
that of Talpa cseca, a character particularly noticeable in the
greater width of rostrum and palate. In details of structure,
however, it shows no striking peculiari-
ties except in the form and position of
the anteorbital foramen. This foramen
i.s noticeably smaller than in Talpa
cseca, and its posterior boi'der lies over
middle or front of second molar instead
of over front of third. The plate form-
ing outer wall of canal is wider and less
thread-like than in TaJpa cseca. Zygoma
about as long as in Talpa cseca, but
owing to the diflerent position of pos-
tei'ior border of anteorbital foramen, the
distance from this poiiat to posterior
edge of temporal fossa equals that from foramen to canine, or
even in one exceptional instance, to outer incisor.

Teeth. — The upper molars are distinctly larger than those of
Talpa cseca, but otherwise the teeth show no peculiarities.

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

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

Talpa caeca (upper figure), and
T. occidentalis (lower tigure).
Nat. sizj.

lal.

Galicia, Spain.

Prof.Seoane (c&p).

94. 1. 1.

23.

2^,29al.

La Granja, Segovia,
Spain.

M.delaEscalera(c).

8. 7. 30.

19-22.

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La Granja, Segovia.

M.delaEscalera(c).

6. 11. 4

2.

2<f,lal.

Cintra, Portugal.

0. Thomas (c&p).

98. 2. 2.

8 9,58

1 skull.

59.

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TALPA ROMANA Thomas.

1902. Talpa romana Thomas, Ann. and Mag. Nat. Hist., 7tli ser., x, p. 517,

December, 1902. Type in British Museum.
1904. Talpa romana Camcrano, Mem. Reale Accad. Sci. di Torino, Qnd ser.,

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

Type locality. — Ostia, Eome, Italy.

Geographical distribution. — Vicinity of Rome, Italy.

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

Skull and teeth. — Except that it is rather more robust the
skull does not differ appreciably from that of Talpa europsea,
though posterior base of zygoma appears to be usually situated
somewhat further back. Teeth as in Talpa europsea, but larger
throughout, a diffei'ence particularly noticeable in the first upper
molar and in the large lower cheek-teeth (the combined length of

Fig. 4.

Crown of niol.ars in Talpa europsea (a), and T. romana (b).

these four teeth about 10 mm. instead of about 8 mm.). In form
the teeth are .similar to those of the related animal, but the
mesostyle of m^ is relatively as well as actually larger, and its tip
is distinctly bifid, though not so deeply as in the succeeding
teeth. Cingulum between outer bases of main cusps of lower
molars better develojied than in T. europsea, its edge frequently
forming an evident cusp, especially in m.y

Measurements. — Extei'nal measurements of type (from
Thomas) : head and body, 126 ; tail, 29 ; hind foot, 19 (all
measurements from skin). Four males from the vicinity of
Rome (in alcohol) : head and body, 138-3 (130-14-5) ; tail, 28-6
(27-30) ; hind foot, I'J -6 (19-20). Three females from the same

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INSECTIVOEA

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

Specimens examined. — Fourteen, from the vicinity of Rome (B3I. and
U.S.N.M.) ; also about fifty from the same region in Turin Museum.

Remarks. — Talpa romana is a well characterized species,
readily distinguished from other European moles by its unusually
lai-ge teeth.

1. Ostia, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 8.

{Type of species.)
1. Frascati, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 9.

S, ? al. Rome. Genoa Museum (e). 3. 1. 31. 1-2.

2 6. Rome. [C. Coli.) G. Barrett-Hamilton (p). 11.1.2.1-2.

Sub-Family DESMANIN.'E.

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

Characters. — Teeth in front of molars not differentiated by
form into incisors, canines, and premolars ; anterior upper
incisor greatly enlarged, canine-like, directed downward, its
outer edge highly trenchant, the two anterior lower incisors
slightly elongated, projecting obliquely forward ; external form
less evidently mole-like than in the Talpime, modified for
aquatic life, the much enlarged hind feet with completely webbed
toes, the front feet smaller, not peculiar in form ; tail long ;
muzzle depressed.

SemnrJcs. — The sub-family Desmanlnx contains the genera
Desmana of Russia and .Siberia, and GaJeinys of the Iberian
Peninsula and south-western France, strictly aquatic animals
not distantly related to the American moles. The anterior
teeth present a much higher degree of specialization than that
met with in the TaJphiie.

Genus GALEMYS Kaup.

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

p. 118 [Mygale pyrcnaica Geoffroy).
184G. Galoniys Agassiz, Nom. Zool., Index Univ., p. 149 (Emendation of

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

(Mygale pyrenaica Geoffroy).

Type species. — Mygale pyrenaica< Geofiroy.

Geographical distribution. — Northern half of Iberian Peninsula,
and Pyrenean region of south-western France.

Characters. — Tail flattened laterally at distal extremity, else-
where terete ; unicuspid teeth slender, their width conspicuously
less than height of crown ; main cusp of large premolar trenchant
anteriorly ; .brain-case without unusual ridges.

GALEMYS 21

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

GALEMYS PYRENAICUS Geoffroy.

(Synonymy under subspecies.)

Geographical disti-ibnfion. — South-western France (Pyrenees
and their immediate neighbourhood) and northern half of Iberian
Peninsula.

Diagnosis. — General characters as in the genus ; head and
body, 110 to 135; tail, 130 to 155; hind foot, 32-5 to 38;
condylobasal length of skull, 33 to 35-5 ; mandible, 22 to 24.

External characters. — Form somewhat intermediate between
that of a mole and rat, the body less elongated than in Talpa,
but the neck almost equally short (so that auditory oritice
appears to be at shoulder) and the muzzle similarly produced ;
legs less shortened than in the moles and front feet not
specially enlarged ; tail rat-like, longer than head and body.
Fur less dei:ise and velvety than in the moles, and of the peculiar
quality characteristic of acjuatic mammals, the hairs of two
kinds, the longer, coarser ones about 12 mm. in length, those of
shorter under fur about half as long. Head conical, as in the
moles, but the much elongated snout (length about 20 mm. from
incisors) greatly flattened instead of terete, its breadth at middle
about 7 mm., dejDth in same region 2 mm. At tip the muzzle
broadens rather abruptly to about 10 mm., the broadened
portion divided by shallow notch at middle of anterior border
into two slightly indicated lobes ; at middle of each lobe and
about 1 mm. back from anterior edge is situated one of
the rather large, transversely elongated nostrils. | The median

* The synonymy of this genus is as follows : —
1777. Dcsmana Gueldenstaedt, " Beschiift. Berliner Gesellsch. Naturforsch.
Freunde, iii, p. 108." (Castor moscliatus Linnaeus.)

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

1800. ikfyjaZc Cuvier, Lee. d'Anat. Comp., I, Tabl. 1. Same type. (Described

in Tabl. E16m. d'Hist. Nat. des Anim., p. 109.)
1815. Desmanus llafinesque, Analyse de la Nature, p. 59 (Emendation of
Desman).

1829. Mi/ogalea Fischer, Synops. ^lamm., p. 250 (Substitute for Mijgalr).

1830. Caprios Wagler, Nat. Syst. Amphibien, p. 14 (Substitute for Mi/galc).
1836. Myogale Brandt, Wiegmauu's Archiv fiir Naturgesch., 1S34, i, p. 17(3.

t The actual narial opening is situated at extreme inner portion of
nostril, and is capable of complete closure bj- the combined action of
antoro-interual border of nostril, a narrow semilunar membrane at upper
edge of inner narial aperture, and a large wart-like thickening of upper
border of nostril.

22

INSECTIVORA

notch is continued backward on upper surface as a narrow
groove still further dividing the two lobes ; on under side
it is continuous with a similar though much longer groove
extending to middle of upper lip. Each lobe is further
marked by a narrow groove extending backward and slightly
inward from near outer edge of nostril. Surface of lobes
very iinely rugose and pitted, that of rest of muzzle coarsely
rugose and without pits. Behind lobes the muzzle is naked
along median region, thinly haired at sides and beneath.
Eye minute, essentially as in Taipa, though probably never
covered by the integument. No external ear, the meatus about
4 mm. in diameter. Front feet rather large and broad, readil}'

turned outward but not perma-
nently in this position, the five
short fingers joined by a narrow
web, and armed with strong,
slightly curved claws 4 to 5 mm.
in length ; fourth digit longest,
tifth and third sub-equal and
slightly shorter, second and first
still shorter. Palms naked, their
surface finely tuberculo - rugo.se^
without trace of larger tubercles,
though the surface is marked
by three deep wrinkles ; balls of
digits projecting conspicuously
beneath bases of claws. Dorsal
surface of fi'ont foot covered
with minute hairs, these lengthen-
ing along edges to form distinct
fringes. Hind foot much larger
than front foot, the toes webbed
to base of claws, the claws
similar to those on front foot
but lai'ger ; fourth digit longest,
third, fifth, second and first suc-
cessively shorter, tlie first extending nearly to end of first
phalanx of second ; surface of sole hke that of palm, the
three longitudinal wrinkles at bases of digits well developed,
the large inner tubercle present in the moles very slightly
developed, its extremity not projecting like a supplemental
digit. Upper surface of hind foot naked, somewhat more
coarsely tuberculate than sole. A fringe of stiffened hairs
along outer edge of outer toe and continuing along foot
nearly to heel. Tail longer than head and body, terete except
at tip, where it is flattened laterally. Scales arranged in some-
what irregular rings, of which there are about nine to the
centimeter at middle of tail. Hairs of tail short, not conceal-
ing scales except on flattened terminal portion, where they form

}"lfi. 5.
Gdlemys i->ijreiiaicits

GALEMYS 23

a rudimentary dorsal and ventral keel. Mamniiv :pl — l,al — 1,
i2-2 = 8.

Colour. — Back and sides dark brown varying somewhat in
exact shade, but never a distinct slaty as in Talpa, the longer
hairs lighter than the under fur and sometimes producing a
slight eliect of coarse " lining," particularly on posterior third
of back and along sides. Under parts buffy in rather strong
contrast with back, but without true line of demarcation.
Hairs of tail and feet buffy. Claws whitish.

Skull. — In general the skull resembles that of Talpa europsea,
but the brain-case is shorter and squarel)' truncate jwsteriorly,
the interorbital region is shorter and narrowed instead of
widened at middle, and the rostrum is longer. Surface of skull
smooth except for the same ridges as in Talpa, those at sides of
brain-case not unusually developed. Brain-case decidedly moi-e
than half as deep as wide, its outline when viewed from behind
vaguely pentagonal, its posterior margin nearly straight, though
with slight median swelling ; condyles not projecting posteriori}^,
completely hidden when viewed from above. Interparietal
projecting further forward than in Talpa, its antero-posterior
diameter nearly equal to its width. Base of brain-case with
deep but broad median furrow, the surface of the bones more
angular and less inflated than in Talpja ; no pits in basioccipital
in front of foramen magnum. Bullse low and flattened, less
perfectly formed than in Talpa, the tympanic bone annular and
retaining its distinctness, though joined with surrounding parts ;
meatus large, occupying about one-half surface of bulla.
Mesopterygoid space short, its length scarcely equal to width of
palate between posterior molars, its width about one-third length,
its anterior border double rounded, encroached on by slight
median spine. Outer pterygoid plate i-educed to a small but
evident ridge. Interorbital region hour-glass shaped, widening
more rapidly posteriorly than anteriorly, its narrowest region
slightly behind middle. Zygoma straight, flattened posteriorly,
compressed anteriorly, its length (measured from posterior border
of orbit to posterior border of anteorbital foramen) equal to
distance from posterior border of anteorbital foramen to front of
canine. Rostrum about as wide as in Talpa, but relatively
longer, the distance from posterior border of anteorbital foramen
to gnathion about equal to greatest breadth of brain-case instead
of much less, its dorsal surface on level with that of interorbital
region, its outer margins nearly parallel to the squarely truncate
anterior extremity. Xares scarcely emarginate posteriorly.
Anterior portion of border of alveolus of large incisor distinctly
thickened, the thickened region terminating laterally in a small
but evident wart-like nodule. Lachrymal foramen over middle
of anteorbital foramen and midille of m^. Posterior border of
anteorbital foramen o^er parastyle of tir. Palate essentially as
in Talpa, but vacuities smaller and incisive foramina large, their

24

IXSECTIVORA

longitudinal diameter about equal to width of palate in same
region. Posterior palatal ridge much as in TaJpa, but the
extremities produced as distinct backward-curved processes.
Mandible lather robust, the ramus neai'ly straight, the angular
I^rocess much below alveolar line Coronoid process high and
narrow, slightly recurved at tip, its height above alveolar line
considerabl)' greater than least breadth of posterior segment of
mandible. Articular process short and robust, the single
articular surface rather large, normal in position. Angular
process longer than articular process, its form essentially as
in Talpa.

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

in general aspect. An-
terior upper incisor much
the largest of all the
teeth, the two together
closing entire front of
palate ; shaft triangular
in cross section, and all
three faces sub-triangular
in outline, the posterior
and antero-external faces
widest, the former slightly
concave, the latter
slightly convex, the two
forming a perpendicular
external cutting edge
about 4 mm. in length
along their line of con-
tact ; a much shorter but
well developed cutting
edge along line of contact
of posterior and antero-
internal faces, extending
from acutely triangular-
pointed apex of tooth to
point of contact with
tooth of opposite side,
a distance of about
0 ■ 6 mm. ; line of contact
between antero-external
and anterointernal faces
marked by a slight thougli evident ridge ; height of shaft
about equal to width of palate ; tirst and second lower incisors
small, chisel-shaped, strongly imbricated, the second about
twice as large as first, their shafts directed forward in line
with upper portion of symphysis menti, the tips of the four
teeth together forminsc a straio;ht transverse cutting edge which

Fig. 6.
Galemys j'urenaicus.

I

acts iu oi^position to combined posterior surface of large ujjper
incisors. Upper unicuspids forming an unbroken row continuous
posteriorly with series of cheek teeth, but separated anteriorly
from large incisoi' by distinct space into which the apex of
second lower incisor tits when jaws are closed. Two anterior
unicuspids (?"" and v') minute, terete, single-rooted, their axes
directed backward and falling in same line with those of two
anterior lower unicuspids when jaws are closed. Third unicuspid
(canine) two-rooted, its crown perpendicular, compressed, larger
than those of first and second combined, and distinctly greater
in height. Fourth unicuspid (pm^) single-rooted, subterete,
scarcely larger than second. Fifth and sixth unicuspids (pnr
and 2>m^) essentially like canine, their crowns distinctly higher
than wide, compressed obliquely to the tooth-row, with slightly
developed anterior and posterior cutting edge. Height of third
and fifth sub-equal, greater than in the others, their tips about
on level with main cusps of molars ; sixth lower than fifth but
with crown longer and posterior ridge better developed. Lower
unicuspids not unlike jj?»^ and jv?«^ but with crowns lower and
longer, slanting a little forward, each with a faintly developed
antero-internal lobule. These teeth are slightl}' imbricated and
their form approximates that of the unicuspids of the Soricidie.
First and second {i^ and ,) higher than third, their form suggest-
ing that of anterior lower incisor, all three single-rooted.
Fourth larger than any of the first three, obscurely two-rooted ;
fifth slightly smaller, single-rooted ; sixth (pm^) largest of the
series, distinctly two-rooted, its cusp nearly on level with
main cusps of molars, its an tei'O external lobule (rudiment of
parastyle) more evident than in the others. Large upper pre-
molar three-rooted, its crown area about equal to that of third
molar, its main cusp with well de^ eloped anterior and posterior
cutting edges, its antero-internal cusp small but evident, its
posterointernal cusp about equal to protocone of m^. Upper
molars with crowns wider and less oblique than in Talpa
europsea, and main cusps not so high. Transverse groove
between bases of main cusps converted into a median pit by
better development of commissures of protocone and larger size
of protoconule and metaconule. Paracone and metacone about
equal in height, the latter slightly the more robust. Styles well
developed, except the reduced parastyle of m^, the mesostyle in
each tooth completely divided into two cusps, the W-pattern thus
changed into two V-shaped figures. Third upper molar with
crown area about two-thirds that of second, its metastyle and
fourth commissure absent. Lower molars essentially as in
TaJpn etirop»a, but contrast in height of outer and innei'
cusps very slight.

26

INSECTIVORA

Galemys pyrenaicus pyrenaicus Geoffroy.

1811. Mijgale pyrenaica GeoSroy, Ann. Mus. d'Hist. Nat., Paris, xvii,

p. 193.
1910. Mijogale pyrenaica Trouessart, Faune Mamm. d'Europe, p. 60.

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

Geogrnphical distrihtition. — Pyrenees and adjacent portion of
southern France ; probably also north-eastern Spain to the
Ebro ; Asturias ?

Diagnosis. — Hind foot, 32 -4 to 3-1: '6 mm.; condylobasal
length of skull about 33 to 34 mm.

Colour. — Back and sides intermediate between prouts-bi'own
and seal-brown, the longer hairs not so dark as under fur and
with a conspicuous lustre ; a small ill-defined buffy area around
eye. Underparts varying from ochraceous-buff to a dull light
cream-buli', clouded by slaty under colour. Front feet dull
ochraceous-buft' tinged with dark brown. Hairs of tail and
fringe on hind foot light buffy.

Measurements. — Two males from Ax-les-Thermes, Ariege,
France: head and bodj^, 115 and 130; tail, 134 and 137 ; hind
foot, 34 "6 and 34. Average of three females from the same
locality: head and body, 117 (110-133) ; tail, 131-6 (126-137) ;
hind foot, 32 '9 (32-4-34). For cranial measurements see Table,
p. 27.

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

Spain : Pajdres, Leon, 2.

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

720 m. France. (p).

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

2. Pyrenees. (Yeircaiix.) Tomes Collection. 7. 1. 1. 15-lC.

1. Unknown. F.Maxwell Lyte (p). 62.1.13.2.

1 al. Pyrenees. (No history.)

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

daki).

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

6, ?. Pajares, Leon, Spain. O. Thomas (p). 8. 2. 9. 47-48.
(N. Gonzalez.)

Galemys pyrenaicus rufulus Uracils.

1897. Myogalea riifida Graells, ilom. Real. Acad. Sci., Madrid, xvii, p. 460.
1910. Myogale pyrenaica rufula Trouessart, Faune Mamm. d'Europe, p. 61.

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

Geograpliical distribution. — Central Spain, south of the Ebro
Valley.

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

27

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28

INSECTIVORA

of skull iibout 34' 5 to 35 • 5 mm. ; colour apparently not so dark
as in the Pyrenean race.

Colour. — Upper parts essentially as in G . pyrenaicus pyrenaicus,
but slightly less dark, the general hue somewhat leaden. Longer
hairs on rump distinctly buffy. Pale area around eye larger and
more noticeable than in the p3'renean foi'ui.

Measurcvienfs. — Three males from Silos, Burgos, Spain : head
and body, 123, 131 and 134 ; tail, 135, 145 and 156 ; hind foot,
36, 38 and 38. For cranial measurements see Table, p. 27.

Specimens examined. — Six, five froin Silos, x'rovince of Burgos, Spain,
and one from Buitrago, province of Madrid (U.S.N.M.).

Hemarhs. — The central Spanish form of Galemys appears to
be well differentiated from true pyrenaicus by its greater size,
a character which is particularly noticeable in the larger,
more massive skull. Two specimens from the Asturias (Xos. 8.
2. 9. 47-48, Pajares, Leon, N. Gonzalez, collector) are apparently
identical with the Pyrenean animal.

When in the water this animal shows much less agility
than the water-rat and watei'-shi-ew, probably because, though in
appearance the most perfectly adapted of the three to aquatic
life, it retains too much of its Talpine inheritance of shortness of
limb and heaviness of gener'al form to be an active swimmer.
Its defective vision, inherited from the same source, would also
tend to a like result.

2 ? sks. Silos, Burgos ; Spain. Eev. S. Gonzalez (c). 8. 7. 7. 8-9.
6, 2 sks. Silos. G. S. Miller (c). 8. 8. 4. 43-45.

Family SORICIDyE.

1821. Soricidx Gray, London Med. Eepos., xv, p. 300, April 1, 1821.

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

Characters. — Skull long and narrow, strongly tapering an-
teriorly', most of the sutures disappearing early in life ; zygomatic
arch incomplete, represented by a slight though usually evident
I'udiment of the zygomatic ^^rocess of maxillary ; floor of brain-
case with median longitudinal bridge of bone and wide lateral
fenestrate area on each side, in which auditory parts are sus-
pended ; tympanic bone annular-, not attached to skull ; basi-
sphenoid without auditory process ; no external pterygoid plate :
mandible with complete double articulation ; anterior teeth not
differentiated by form into incisors, canines and premolars, the
first upper incisor very large, strongly projecting forward, its
tip hooked downward, its base with a secondary lobe, the anterior
lower incisor nearly straight, much i)roduced in axis of mandible,
the other anterior teeth foi'ming a series of small " unicuspids,'
differing from each other chiefly in size : crowns of upper molars

SOREX 29

low, sub-(iuadi'ate in outline (except the much reduced third), the
paracone and metacone near middle of crown, the styles and
commissures well developed and forming an important functional
part of the cutting apparatus ; form mouse-like, but snout always
pointed and much produced beyond incisors, eyes small, and ears
often partly or entirely hidden in the fur.

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

KEY TO THE EUROPEAN GENERA OF SOBICIDjE.

Posterior lower molar with five cusps ; teeth pigmented

at tips ; tail without sprinkliug of elongated hairs.

Upper unicuspid teeth 6-5 ; cutting edge of anterior

lower incisor with more than one lobe ; feet not

fringed Sorex, p. 29.

Upper unicuspid teeth 4-4 ; cutting edge of anterior
lower incisor with one lobe; feet fringed. (Water

Shrews) Neomys, p. 65.

Posterior lower molar with four cusps ; teeth white
throughout ; tail with noticeable sprinkliug of
elongated hairs.

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

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

Genus SOREX Linn.'eus.

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

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

p. 119 (Included the undeterminable Sorcx constrictus Hermann

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

II, p. 28 (hcrmanni = Neo7nys fodiens skull -f Sorex araneus

tetragomrus animal) Part.
1838. C&rsira Gray, Proc. Zool. Soc. London, 1837, p. 123. June 14, 1837

(vulgaris = araneus).
1842. Otisorex De Kay, Zool. of New York, i, Mamm., p. 22 (j)latyr]iinus

= personafus).
1857. Sorcx Blasius, Siiugethiere Deutschlands, p. 124.
1890. Homalurus Schulze, Schriften Naturwiss. Vereins Harzes in Werni-

gerode, v, p. 28 (nlpinus).

Type species. - Sorex araneus Linnanis.

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

30

INSECTIVOEA

Characters. — Ujjper unicuspid teeth 5-5 (dental formula :
i ?=?, cti ^Jm'^, iii'^ = 32) ; posterior lobe of anterior upper
incisor fully half as high as main cusp ; anterior lower incisor
with three well developed lobes on cutting edge ; third lower
molar with hypoconid and entoconid small but distinct, so that
the form of the tooth differs from that of first and second molars
in the reduced size of the second triangle only, its crown, like the
others, 5-cusped ; second lower unicuspid with rudimentary
second cusp and commissure ; points of all the teeth pigmented
(the coloured portion wearing away in extreme old age) ; skull
lightly built, with slender weak rostrum and abruptly wider
brain-case ; rudimentary zygomatic process of maxillary evident ;
no special modifications in external form ; tail covered with
hairs of uniform length (except that those of pencil are elon-
gated) ; ear nearly concealed by the fur, the meatus closed by
a large valvular outgrowth from the antitragus supplemented
by a fold on inner surface of conch ; habits terrestrial.

Bemarhs. — This is the most widely distributed genus of
Insectivora. It contains about sixty described forms, fourteen
of svhich occur in Europe.

KEY TO THE EUROPEAN FORMS OP SOBEX.

Anterior mandibular incisor with low, sometimes ill-
defined lobes on cutting edge ; first lower uni-
cuspid two-pointed; lachrymal foramen over
point of contact between m' and m- ; tail about
as long as head and body ; colour uniform darlj

slaty grey. (Alpine Shrews) .S'. alpinus, p. 60.

Condylobasal length of sliuU 19-4 to 20-6 mm.;
upper tooth-row 8 • 6 to 9 ■ 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 mandiljular incisor with high, distinct lobes
on cutting edge ; first lower unicuspid single-
pointed ; lachrymal foramen in front of point of
contact between ?»' and m" ; tail shorter than
head and body ; colour never uniform dark slaty
grey, the under parts usually much lighter than
back.
First, second and third upper unicuspids sub-equal;
condylobasal length of skull 14 '8 to 16 ■6 mm.;
head and body usually about 60 to 60 mm.

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

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

Molars and anterior upper incisor enlarged

(Southern Italy) ,S'. m. hicanius, p. 60.

First and second upper unicuspid much larger than
third; condylobasal length of skull 17-4 to
20 mm. ; head and body usually about 65 to
80 mm. (Common Shrews) S. arancus, p. .31.

SOREX 31

Palate broad anterioi'ly, its width at level of first
unicuspid nearly equal to that of tooth.
Condylobasal length of skull 18--1 to 19-2 mm.;

anterior toeth enlarged (Island of Jersey). . . S. a. fretalis, p. 45.
Condylobasal length of skull about 17 • 5 mm. ;
anterior teeth not enlarged (Jlountains of

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

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

Hind foot 13 • 0 to 14 mm. ; colour in summer
pelage frequently light and brownish.
Back frequently blackish in summer pelage

(Alps and neighbouring regions) S. a. tetra^ontirus, -p. 4:2.

Back rarely if ever blackish in summer

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

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

back (Charente, Prance) S. a. aantonus, p. 40.

Underparts greyish or brownish, contrasted

with back except when latter is also

brown.

Colour of sides not distinctly contrasted

with that of back ; belly heavily

washed with wood-brown (Plains of

south-western France) S. a. euronotus, p. 41.

Colour of sides usually forming distinct
contrast with that of back ; belly
lightly washed with wood-brown.
Average colour darker, the back ranging
from bister to a deep blackish
brown (Central Europe and Scan-
dinavia, except south - western

Norway) ,S'. a. araneus, p. 35.

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

SOREX ARANEUS Linnajus.

(Synonymy under subspecies.)

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

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

32

IXSECTIVORA

compressed latei'all\', the length of its base equal to that of base
of anterior cusp ; colour brown or blackish, the underparts
never as dark as back (usually much lighter), and the sides often
contrasted with both.

External characters. — Fur moderately dense, its depth at
middle of back about -i mm. in summer, 8 mm. in winter, its
texture not specially modified ; no elongated hairs on Hanks and
across rump. Eyes small and inconspicuous ; ears nearly
concealed in the fur. Feet not peculiar in form, thinly clothed
with very fine hairs on dorsal sui-face ; fore foot with third and
fourth digits sub-equal and longest, fifth extending just beyond
base of fourth, first barely reaching base of second, the claws
small but well developed ; palm so conspicuously rugose reticulate
that tubercles are not very distinct ; tubercles 6, sub-equal, the
three at bases of main digits well-defined, that at base of thumb
sometimes confluent with that at inner side of wrist (so that the
number is ajjparently reduced to 5), the two wrist-pads separated
from each other in median line by a noticeable space ; extreme
posterior edge of palm covered with ordinary integument. Hind
foot with third and fourth digits sub-equal and longest, second
slightly shorter, fifth reaching base of fourth, and first extending
to base of fifth ; surface of sole as in palm, but tubercles better
defined, four at bases of digits and two situated more posteriorly,
all six about equal in size ; sole finely haired fi-om hinder
tubercles to heel, the middle of which is bare ; claws like those
of fore-foot. Tail terete or somewhat 4-sided, rather more than
half as long as head and body, its hairs minute, ratlier closely
appressed, and nearly concealing the annulation ; pencil usually
well developed, 4-6 mm. in length, but occasionallj^ in aged
individuals much reduced or absent, together with the rest of
the hairy covering of the tail. Caudal annulations rather
indistinct, about 24 to the centimeter at middle. jNIammfe :
/ 3-3 = 6.

Colour. — Dorsal area, extending from base of tail to crown
cheeks and muzzle, brown, the exact shade varying much both
seasonally, racially and individually, but the normal extremes
falling between hair-brown or light bister and a very dark,
blackish seal-brown. Sides wood-brown, usually forming an
evident contrast with dorsal area, though this contrast is
occasionally inconspicuous in dull light specimens in summer
coat, or wholly obliterated in the general darkening of entire
animal in S. araneus sanfonus. Underparts smoky grey washed
with wood-brown, or occasionally sufiused with slaty (particularly
in the dark »S'. a. santonus). Between colour of back and sides
there is usually an evident line of demarcation : between sides and
belly the contrast is less marked and the transition less abrupt.
Feet a dull indefinite light brown, often with a faint dark shade
along outer edge. Tail dark brown above and at tip, light brown
below, especially near base, sometimes bicolor throughout.

SOREX 33

While there is no invariable rule, the colour in winter is
usually darker than in summer, and the tricolor pattern of
dark back, yellowish brown sides and greyish belly is more
pronounced. A trace of tliis pattern is often the most convenient
character by which to recognize shrunken ill-prepared specimens,
which might otherwise be mistaken for Sorex minutus.

SJcuIl~The skull is slender and lightly built, with no special
peculiarities of form as compared with that of other shrews
Brain-case well marked off from interorbital region, its surface
smooth except in extreme old age, its main sutures remainincr
open until late in life. It is sub-circular in °

general outline when viewed from above,
but with antero-external portion of border
noticeably flattened, so that at point of
greatest breadth there is usually an evident
angle ; condyles scarcely visible, causing no
break in posterior outline. Depth of brain-
case at middle slightly more than half
greatest width ; no sagittal crest except in
extreme old age; lambdoid crest at tirst
confined to lateral portions of occiput, rarely
extending to median line. Dorsal profile
usually with evident concavity in inter- ^^''" ""

orbital region (more marked than in Sorex ^"wat S!"'"

minutus and ,S'. alpiniis). Nares broadly
rounded posteriorly, the lateral margin obtusely angled near
middle. Anteorbital foramen moderately large, not very con-
spicuous when skull is viewed from in front. Lachrymal
foramen over middle of m\ Mesopterygoid space nearly parallel-
sided, less than half as wide as long.

Teeth. — Anterior upper incisor with basal lobe relatively
larger than in any other European shrew, the length of its
base nearly equal to diameter of anterior lobe at level of angle
between the two cusps. When tooth is viewed from below the
posterior lobe appears nearly as large as anterior cusp ; in
lateral view it approximates the size and form of first and
second unicuspid, and in height falls a little short of anterior
cusp. The two teeth come in contact anteriorly slightly below
tips, which do not diverge conspicuously. Anterior lower incisor
robust, its shaft very slightly tapering, its cutting edge with
three well-developed lobes, the lengths of bases of which
fhminish regularly from first to third, the first lobe more
distinctly separated from succeeding lobe than from anterior
point of tooth. Upper unicuspids robust, their crowns squarish
in outline when viewed from below, bluntly triangular when
viewed from the side ; height about e.jual to length ; anterior
and posterior borders straight or faintly concave, upper border
convex, more strongly posteriorly than anteriorly. The highest
point of cusp lies slightly in front of middle of crown. Upper

D

34

INSECTIVORA

riG. 8.

Siirex araneits: Anteriur

teeth in prolile. X 5.

and posterior borders sub-equal, slightly longer than anterior

border. Cusp rounded on antero-external face, squarely truncate

postero-internally atong line connecting antero-internal and
postero-external corners of crown. The
region behind this line is occupied by a
flattened or somewhat concave, nearly
horizontal crushing surface opposed
during mastication to second and third
lobes of lower incisor (1st and 2nd upper
unicuspids) and to points of the two
lower unicuspids (3rd and 4th upper
unicuspids). In size the first and second
unicuspids are sub-equal and decidedly
larger than third, which in turn some-
what exceeds fourth. Fifth still smaller

than fourth and closely crowded between it and antero-external

cusp of large premolar, its cusp relatively lower than in the

other teeth of the series. First lower iniicvispid essentially

similar to i3rst and second upper,

but crown longer than broad and

without distinct crushing surface.

Second larger than first, not

conspicuously different from it in

general form when viewed from the

side, but cutting edge much longer

and better developed, distinctly

angled behind middle, the angle

clearly i-epresenting a rudimentary

second cusp, and transverse portion

behind it a second commis-
sure, the tooth thus contain-
ing the modified elements

of one of the triangles of a

molar. Large upper pre-

molar with protocone

smaller than in the first

and second molars, though

well developed and of

essentially the same form ;

paracone smaller than pro-
tocone, therefore much

smaller than in first and

second molars ; hypocone

small but well developed ;

posterior margin of crown

more deeply emarginate

than that of molars.

Crowns of first and second molars squarish in outline, though

somewhat wider posteriorly than anteriorly. Outer re-entrant

angles deeper in second than in first. Pi-otocone long and rather

i'lG.
Sorex aranetis.

SOREX 35

low. Hypocone small but well developed. Third upper molar
with about half the crown area of second, all the elements of the
tooth present except hypocone. First and second lower molars
alike in size and form, the posterior triangle slightly larger than
anterior. Third molar smaller than the others, the relative size
of the triangles reversed, but elements of tooth all present.

Measuretiients. — While there is some variation in size among
the different races the head and body in full-grown individuals
is seldom if ever less than 65 or more than 80 mm. The tail
falls short of head and body by from 20 to 35 ram., being thus
relatively shorter than in either of the other European species.
The hind foot ranges from 11 to about 14 '5 mm. and the
condylobasal length of skull from 17 '8 to 20 mm. Detailed
measurements are given under each of the subspecies.

Bemarlcs. — Sorex araneus, the most widely distributed and
best known of the European shrews, is so easily recognized that
it needs no special comparison with the other species. Specimens
of the dark race from Charente, France, have a superficial
likeness to Sorex alpinus, but are at once distinguishable
externally by their blackish instead of slaty coloration and
relatively short tail. Immature, dull coloured individuals of
the other races may occasionally be confused with S. minutus.
The same is true of shrunken, badly prepared skins. But some
trace of the tricolor pattern can almost invariably be found in a
common shrew no matter what its condition, while a glance at
the size of the teeth and form and proportions of the unicuspids
will always serve to determine the identity of any specimen in
hand.

SoBEX AKANEUS AKANEUS Linnseus.

1758. [Sorex^ araneus Linnseus, Syst. Nat., i, 10th ed., p. 53 (Sweden).

1828. Sorex coronatus Millet, Faune de Maine-et-Loire, i, p. 18 (Blou,
Maine-et-Loire, Prance).

1828. Sorex iiersonatus Millet, Faune de Maine-et-Loire, I, p. 18, foot-
note (Rejected MS. name for coronatua). Not Sorex personatus
I. Geoffroy, 1827.

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

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

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

1838. Sort's; vulgaris Nathusius, Wiegmann's Archiv fiir Naturgesch.

IV, I, p. 45.

1839. Skirex] macrotrichus de S61ys-Longchamps, Etudes de ^licromamm.,

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

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

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

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

T[ipc locality. — Upsala, Sweden.

Geographical distribution. — Western Continental Europe,

36

INSECTIVORA

except Atlantic watershed of south-western Norway, from
Finland to central France, central Germany, and northern Hun-
gary. Exact southern and eastern limits of I'ange not known.

Diagnosis. — Size small (condylohasal length of skull usually
17 "8 to 19 mm., hind foot usually 11 to 13 mm.) ; colour rather
dark, the back ranging from bister to a deep blackish brown ;
sides distinctly lighter than back except in specimens representing
the pallid extreme of colouration ; teeth moderately pigmented,
the hypocone of m^ and m'^ usually white to tip.

Teeth. — The teeth show no special peculiarities of form. In
pigmentation they represent the extreme of restriction of the dark
areas. While the area of j^igoientation on all of the cusps is less
extensive than in the Alpine and Pyrenean races, the ditierences
are best seen in the hypocones of the three lai'ge upper cheek
teeth and }:)rotocone of m^, as these small cusps, unlike the larger
ones, may be completely without brown colour. A comparison
of seventy-five topotypes of Sorex araneus with eighty Swiss
specimens of *S'. araneus tetragonmus gives the following results : —

araneuK. tetragonurug.

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

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

Second molar with pigment on hypocone . 21'2 % ... 88' 7 %

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

None of the small cusps pigmented . . 54 • G % ... " 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, centi'al Gudbrandsdal, Norway
(dry): hind foot, 12 '3 (12 "2-1 2 '6). Average and extremes of
four specimens from Holaaker, upper Gudbrandsdal, Norway :
head and body, 70-2 (69-71); tail, 36-5 (34-39); hind
foot, 12 "2 (12-13). Average and extremes of twenty specimens
from Brunswick, Germany : head and body, 78 " 9 (68-85) ;
tail, 39 ■ 7 (36-45) ; hind foot, 1 2 • 7 (1 2 • 2-1 3 -^O). Average and
extremes of ten specimens from Waremme, Liege, Belgium : head
and body, 68-9 (66-72); tail, 42-3 (38-47)'; hind' foot (dry),
12 '2 (11 '8-1 2 '8). For cranial measurements see Table, p. 46.

Specimens examined. — Two hundred and ninety-eight, from the following
localities : —

Norway : Molmen, Upper Gudbrandsdal, 2 ; Holaaker, Upper Gud-
brandsdal, 4 ; Lesjevark, Middle Gudbrandsdal, 1 ; Lillehammer, Middle
Gudbrandsdal, 9 (U.S.N.M.) ; Eggedal, Buskerud, 8 (U.S.N.M.) ; Spjosod,
Telemarken, 4 (U.S.N.M.) ; Asker, near Ohristiania, 4 (U.S.N.M.) ; Holme,
Mandal, 7.

Sweden: Upland,!; Upsala, 97 (U.S.N.M.) ; Skaane, 3 (U.S.N.M.).

Denmabk : HillerOd, Zealand, b ; Nystad, LoUand, 3 (U.S.N.M.) ;
Skansen, Lolland, 1 (U.S.N.M.).

Holland: Oosterbeek, Guelderland, G; Leiden, 4 (U.S.N.M.).

Belgium : Hastiere, Namur, 1 ; Waremme, Li6ge, 10 (U.S.N.M.).

Fbance : Guines, Pas-de-Calais, 4 ; Manonville, Meurthe-et-Moselle, 2 ;
Barbizon, Seine-et-Marne, 3.

37

Gebmany: Brunswick, 35 (BM. and U.S.N.M.) ; Bodethal, Harz Mts.,
15 (U.S.N.M.); Mcauseklippe, Harz Mts., 2 (U.S.N.M.); Bahrenberg, Harz
Mts., 9 (U.S.N.il.) ; Tharaud, Saxony, 1 ; Magdeburg, Saxony, 5 ; Moritz-
burg. Saxony, 1 (U.S.N.M.); Ingeiheim, Rheinhessen, 3; Nuremberg,
Bavaria, 10 (U.S.N.M.) ; Marxheim, near Monheim, Bavaria, 15 ; Strass,
near Burgheim, Bavaria, 1 ; Eulengrund, Riesengebirge, Silesia, 2
(U.S.N.M.) ; Wolfshau, near Sneekoppe, Riesengebirge, Silesia, 8 (U.S.N.M.);
Niesky, Silesia, 5 ; near Konigsberg, 6 (U.S.N.M.) ; no exact locality, 1.

AusTRiA-HuKGAEY : Haida, Arva, Bohemia, 9.

6, 9.
2(5,2 9.

6.

6, 5 9.

dal.

9.

4:6,2 9.

4 d,2 9.

9.

3 6,19.

2 6,2 9.

6, 3 9, 2.

9.
5 al.

<5, 2 9.

3 6, 4 9.

2 6.

6.

3 6, 2 9.

2al.

1.

3 6. 4 9.

jNIolmen, Gudbrandsdal.
Norway.

Holaaker, Gudbrands-
dal, 1900 ft.

Lesjevark, Gudbrands-
dal.

Holme, Mandal, 200 ft.
Norway.

Upland, Sweden.
(G. Kolthojf.)

Hillerod, Zealand, 10 m.
Denmark.

Oosterbeek, Guelderland ,
10-15 m. Holland.

Hastiere, Namur, Bel-
gium.

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

Barbizou, Seine - et-
Marne.

Auerum Forest, Bruns-
wick, Germany.

Tharandt, Saxony.

Magdeburg, Saxony.

Ingelhoim, Rheinhessen.
Marxheim, Bavaria.
Bayreuth, Bavaria.

Strass, Burgheim, Ba-
varia.

Niesky, Silesia, 200 m.
(W. Bacr.)

No exact locality.

Germany.

Haida, Bohemia.
Haida, Bohemia.

R. J. Cuuinghame

R. J. Cuninghame

(P)-
Miller Collection.

K. J. Cuninghame

Lord Lilford (p).

O. Thomas (c & p).

O. Thomas (c & p).

G. A. Boulenger

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

G. S. Miller (c).

G. Barrett-Hamil-
ton (c & p).
Lord Lilford (p).
Dr. W. Wolterstorff

(P)-
C. Hilgert (c).
Lord Lilford (p).
IMiller Collection.

Lord Lilford (p).

Lord Lilford (p).

Zool. Soc. Collec-
tion.
Stockholm ^Museum

(E).

Lord Lilford (p).
Lord Lilford (p).

98. 5. 2. 1-2.
98. 2. 28. 1-4.

7. 7. 7. 4452.

8. 8. 9. 1-6.
8. 8. 9. 40.
8. 9. 8. 19.

98. 6. 7. 2-7.

98. 2. 1. G-8.
94. 7. 9. 1.
94. G. 6. 4-7.

8. 8. 4. 155-158.
11. 1. 2. 75-80.

99. 1. 9. 14.
92. 12. 1. 3-7.

8. 11. 2. 6-3.
^. 9. 8. 7-13.

7. 7. 7. 2363-
2869.

8. 9. 8. 20.

99. 1. 9. 9-13.

55. 12. 26. 300-

301.
46. 6. 2. 36.

8. 9. 8. 14-18.
8. 9. 8. 21-22.

SOREX ARANEUS CASTANEUS JenyiLS.

1838. S[oc(?a-] tctragonums var. ;8 Slorcx] castaneus Jenyns, Ann. Nat.

Hist., I, p. 424, August, 1838 (Burwell Fen, Cambridgeshire,

England).
1857. Sorex vulgaris Blasius, Saugethiere Deutschlands, p. 129 (part)
1910. Sorex araneus Trouessart, Faune Mamm. d'Europe, p. 51 (part).

Type locality. — Burwell Fen, Cambridgeshire, England.
Geographical distribution. — Great Britain. Usually confined

38

INSECTIVORA

to the mainland, though occurring on Bardsey Island, Carnar-
vonshire.

Did'jiwsis. — Similar to Sorex araneus araneiis, but colour in
series of skins averaging less dark, that of dorsal area ranging
from hair-brown slightly tinged with bister to seal-brown, and
seldom if ever attaining the deep blackish brown frequently
seen in the typical race.

Skull mid teeth. — The skull and teeth resemble those of true
Sorex araneus.

Measureiiients.--Avevd.ge and extremes of ten specimens from
Cromarty, Scotland: head and body, 70*6 (62-78); tail, 39 "8
(38-43); hind foot, 12 '9 (12- 5-13). Average and extremes
of eight sjaecimens from Aberia, Merioneth : head and body,
67-5 (58-73); tail, 38-7 (36-41); hind foot, 12 (11-13).
Average and extremes of eight specimens from Grimsby, Lincoln-
shire : head and body, 65-2 (58-72); tail, 42 (38-44); hind
foot, 12 '5 (12-13). Average and extremes of six specimens
from Northlew, Devonshire : head and body, 66 '6 (65-70) ; tail,
36 '8 (35-39); hind foot, 13 (13). For cranial measurements
see Table, p. 48.

Specimens examined. — Two hundred and fifty-two, from the following
localities : —

Scotland: Black Isle, Cromarty, IG ; South Sutor, Cromarty, 5;
Nairn, Morayshire, 4 ; Dunphail, Elgin, 2 ; Gordonstown, Elgin, 3 ; Lhan-
bride, Elgin, 1; Lossiemouth, Elgin, 3; Grantown-on-Spey, Elgin, 26
(Wilson) ; Kennordy, 1 (Wilson) ; Cortachy, Forfar, 6 (Wilson) ; Cromlix,
Stirling, 10 ; Islay, 3 ; Dunkeld, Perthshire, 2 ; Loch Earn Head, Perth-
shire, 1; Stoclibriggs, Lanarkshire, 2; Kirtle Bridge, Dumfriesshire, 4;
Wyseby, Dumfriesshire, 3.

Wales : Aberia, Merionethshire, 8 ; near Bridgend, Glamorganshire, 6
Bardsey Island, Carnarvonshire, 1.

England : Berwick-on-Tweed, Northumberland, 2 ; Riding Mill-ou-
Tyne, Northumberland, 3 ; Newby Bridge, Lake Windermere, Cumber-
land, 1 ; Grimsby, Lincolnshire, 17 ; Whitnash, Warwickshire, 1 ; Rugby,
Warwickshire, 2 ; Filey, Yorkshire, 3 ; Wellersey Hill, Broadway, Wor-
cestershire, 1 ; W^est Cheshire, 1 ; Shropshire, 1 ; StaSordshire, 1 ; Swith-
land, Leicestershire, 10 ; Bishopstoke, Herefordshire, 1 ; Leominster, Here-
fordshire, 1 ; Graftonbury, Herefordshire, 19 ; Lilford, Northamptonshire,
2 ; Drinkstone Park, Bury St. Edmunds, Suffolk, 2 ; Lowestoft, Suffolk, 6 ;
Wormsley, Oxfordshire, 1 ; Stokenchurch, Oxfordshire, 4 ; Cambridge-
shire, 1; Kensington Gardens, London, 1; Hillingdon, Middlesex, 3;
Bletchingley, Surrey, 4; Godalming, Surrey, 2; Merstham, Surrey, 1;
Richmond Park, Surrey, 1; Crowborough, Sussex, 3; St. Leonard's,
Sussex, 1 ; Tunbridge Wells, Sussex, 1 ; Eastwell, Kent, 3 ; Lyndhurst
Road, Hampshire, 2; New Forest, Hampshire, 17; Basingstoke, Hamp-
shire, 1 ; Alum Bay, Isle of Wight, 3 ; Cliftou Bridge, Gloucestershire, 2 ;
Leigh Woods, Clifton, Gloucestershire, 3 ; Blandford, Dorsetshire, 1 ;
Combmartin, Devonshire, 4 ; Chagford, Devonshire, 5 ; Northlew, Devon-
shire, 6 ; no exact locality, 3.

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

SOREX 39

race the dark extreme is less dark and less frequent, while the

light extreme is more light and more frequent. The constantly
small size of this animal as compared with the large Continental

races (benjensls, ietragonurus and pi/renaicus), is shown by the
fact that among 102 British specimens measured by many
different collectors the hind foot exceeds 13 '2 mm. in only six
instances.

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

9 i. Black Isle, Cromarty. W. R. Ogilvie-Grant 11. 1. .3. 103-

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

2(5. South Sutor, Cromarty. W.R. Ogilvie-Grant 11. 1. 8. 108,

(c & p). 113.

2 (5, 2 ?. Nairn, Morayshire. W.R. Ogilvie-Grant 11. 1. 3. 99-102.

(c & p).

5. Morayshire. E. R. Alston (c & p). 79.9.2.5.79.
9. Dunphail, Elginshire. W. R. Ogilvie-Grant 11.1.3.97.

(c & p).

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

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

shire. (c & p).

6. Gordonstown, Elgin- W. R. Ogilvie-Grant 11.1.3.98.

shire. (c & p).

3(5,7 9. Cromlix, Stirlingshire. W. R. Ogilvie-Grant 11.1.3.114-128.

(c & p).

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

6,9. Stockbriggs, Lanark- E. R. Alston (c & p). 79.9.25.11-12.

shire.

6, 3 9. Kirtle Bridge, Dumfries- Miss D. Bate (c <fep). 11. 1. 3. 88.

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

3 9. Wyseby, Dumfriesshire. Miss D. Bate (c & p). 11. 1. 3. 89-91.
5 (5, 3 9. Aberia, Merionethshire, G. H. Caton Haigh 11. 1. 3. 156-

Wales. (c & p). 163.

3(5,3 9. Bridgend, Glamorgan- R. I. Pocock (c & p). 11. 1. 3. 164-

shire. 169.

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

Northumberland,
England.

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

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

(c & p). 131.

(5,6 9. Swithland, Leicester- P. A. Butler (c & p). 11. 1. 8. 132-

shire. 188.

1 al. Shropshire. T. 0. Eyton (c & p). 68.10.12.4.

9 al. Bishopstoke, Hereford- S. 0. Ridley (c&p). 84. 10. 6. 1.
shire.

8 c?, 8 9. Graftonbury, Hereford- W. de Winton 11.1.3.72-87.
shire. (c & p).

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

shire. (c&p).

2al. Bury St. Edmunds, J. H. Powell (c & p). 80.5.22.1-2.
Suffolk.

3 (5, 3 9. Lowestoft, Suffolk. 0. Thomas (c & r). 11. 1. 8. 189-

144.

4 9. Stokcnchurch, Oxford- W.R. Ogilvie-Grant 11. 1. 8. 145-

shire. (c & p). 148.

3 9. Hillingdon, Middlesex. MiUer Collection. 7. 7. 7. 8595-

(O. Thomas.) 3597.

40

IXSECTIVORA

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

Loudon.

4 f . Bletchingley, Surrey. W.R. Ogilvie-Grant 11. 1. 3. 149-

_(p). 152.

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

{W.li.Ogilvic-Grant.)

9 al. Richmond Park, Surrey. Prof. Owen (p). 75.9.17.1.

7 6, 10 9. New Forest, Hampshire. Miller Collection. 7. 7. 7. 2851-

2867. 3023.

9. Basingstoke, Hamp- Miller Collection. 7. 7. 7. 3611.

shire. {W. P. Stark.)

6, 2 9. Alum Bay, Isle of Wight. 0. Thomas (c & p). 11. 1.8.153-155.

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

6, 4 9. Nortblew, Devonshire. R. C. Wroughton 11. 1. 3. 170-

(c & p). 174.

2 c5, 3 9. Chagford, Devonshire. Miller Collection. 7. 7. 7. 3606-
(W. P. Stark.) 3610.

1 al. England. Dr. J. E. Gray (p). 46. 5. 2. 7.

80RE.\ ARANEUS SANTONUS Mottaz.

1908. Sorex santonus Mottaz, Bull. Soc. Zool. de Geneve, i, p. 118,

April 30, 1908. Type in Mottaz Collection.
1910. Sorex ,';a)itonus Trouessart, Faune Mamm. d'Europe, p. 54.

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

Geographical distribution. — Known only from the vicinity of
the type locality.

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

Colour. — Upper parts dai'k sepia anteriorly, deepening to
blackish posteriorly, the sides essentially similar to the back.
Underparts a slaty drab washed with wood-brown and forming
no evident contrast with sides and back, though a well-detined
line of demarcation is present. Tail blackish throughout. Feet
scantily clothed with inconspicuous brownish hairs.

Skull and teeth. — The skviU and teeth do not differ appreciably
from those of the other small races.

Measurements. — Type (female), from Mottaz : head and body,
75 ; tail, 42 ; hind foot, 13. For cranial measurements see
Table, p. 51.

Specimens examined. — Ten, all from the type locality (Mottaz).

BemarJcs. — The Charente shrew is so different in aspect from
tlie other European races of Sorex araneus that it needs no
special comparison with any of them. Its jseculiar colour is,
however, almost exactly duplicated by that of a larger, longer -
tailed form from Asia Minor.

41

SOKEX ARANEUS EURONOTUS Miller.

1901. Sorex araneus curonotus Miller, Proc. Biol. Soc, Washington, xiv,
p. 44, April 25, 1901 (IMontrejeau, Haute-Garonne, Prance). Type
iu U.S. National Museum.

1910. Sorcx araneus curonotus Trouessart, Paune Mamm. d'Europe, p. 53.

Ti^je locality. — Montrejeau, Haute-Garonne, France.

Geographical distribution. — Probably the i^lains between the
Pyrenee.s and the Garonne. At present known from the type
locality only.

Diarjnotiis. — Similar to Sorcx araneus araneus, but colour in
.summer (winter pelage not known) more dull, the sides scarcely
if at all contrasted with the back, and underparts more heavily
washed with wood-brown. 8kull with slightly narrower less
elevated brain-case, and teeth just perceptibly smaller than in
true araneus.

Measurements. — External measurements of type specimen :
head and body, 78; tail, 44; hind foot (dry), 12 '6. Average
and extremes of nine specimens from the tyjje locality : head and
body, 71-4 (67-78); tail, 42 (37-44); hind foot (dry), 12-5
(12-2-13'0). For cranial measurements see Table, p. 51.

Specimens examined. — Niue, all from the type locality (U.S.N.M.).

SoREX ARANEUS BERGENSIS Miller.

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

Ill, p. 416, May, 1909. Type iu U.S. National Museum.

1910. Soi-ex araneus bergensis Trouessart, Paune Mamm. d'Europe, p. 52.

Tijpe locality. — Graven, Hardanger, Norway.

Geographical distribution. — Western Norway, from region of
Bergen, north at least into Nordland.

Diagnosis. — Larger than Sorex araneus araneus (hind foot,
13-6 to 14-4 mm. ; condylobasal length of skull, 19 to 20 mm.),
and colour in summer pelage darker, the dark brown or blackish
dorsal area sharply defined from the yellowish brown of sides.

Colour. — Dorsal area well defined, ranging from a blackish
seal-brown to bister, usually darker on rump and lumbar region
than on head, and almost invariably with sharp lateral line
of demarcation. Sides yellowish isabella-colour. Underparts
greyish, washed with light wood-brown. The contrast between
colour of sides and underparts, though less marked than that
between sides and back, is usually evident. Feet scantily clothed
with inconspicuous Isabella-coloured or dusky hairs. Tail obscurel)*
bicolor, brownish above, dull yellowish below.

Sliull and teeth. — The skull and teeth do not differ appreciably
from those of Sorex araneus araneus, except in their larger size,
as shown by the detailed measurements. The pigmentation of

42 INSECTIVORA

the teeth is scarcely, if at all, more extensive than in the typical
race.

Measurements. — External measurements of type : head and
body, 80-5; tail, 44-5; hind foot, 13-6. Average and extremes
of eleven specimens from the Bergen district : head and body,
78-7 (76-83) ; tail, 49-3 (44-56) ; hind foot, 13-8 (13 -6-14 -4).
For cranial measurements see Table, p. 51.

Specijnens examined. — Twenty-seven, from the following localities: —
Noeway: Vefsen, Nordland, 1; Skjserdal, Nordfjord, 7; Opheim,

Bergen, 4; Graven, Bergen, 8 (U.S.N.M.) ; near city of Bergen, 7 (B.M.

and U.S.N.M.).

Bemarlcs. — This large race of Sorex araneus, closely resembling
the large Alpine and Pyrenean forms, but decidedly darker in
colour, appears to be strictly confined to the Atlantic slope of
western Norway. On the eastern watershed it is replaced by
true araneus, even so far north and west as the upper portion of
the Gudbrandsdal. I have not seen specimens in full winter
pelage, but an adult female taken at Graven on June 10, 1898
(No. 84,663, U.S.N.M.), is moulting, the winter fur remaining on
posterior half of body. Others taken at the same place and
about the same date have completed the change to the short,
velvety summer coat.

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

way. Waldo (c & p).

6, 4 9. Skjserdal, Nordfjord. C. H. Stephenson 8. 1. ,5. 1-5.

(c & P).

■2 6,29. Opheim, Bergen. Miller CoUection. 7. 7. 7. 44.53-

445G.

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

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

ton (p).

SoREX ARANEUS TETRAGONURUS Hermann.

1780 Sorc.v tetragonurus Hermann in Zimmermann, Geogr. Gesch., ii,
p. 383 (Strassburg, Germany).

1792. Sorcx quadricaudatus Kerr, Auim. Kingd., p. 208 (Strassburg,
Germany). (Based on Pennant's account of S. tetragomirus
Hermann.)

1835. Sorcv fodiens Duvernoy, Mem. Soc. du I\Ius. d'Hist. Nat., Stras-
bourg, II, p. 17. Part : skull, not animal.

1835. Sorex hermanni Duvernoy, M6m. Soc. du Mus. d'Hist. Nat., Stras-
bourg, II, p. 23 (Near Strassburg, Germany). Part: animal, not
skull.

1857. Sarex vulgaris Blasius, Saugethiere Deutschlands, p. 129 (part).

1868. ? Sorex aranetis, pallidus Fitzinger, Sitzungsber. kais. Akad. Wis-
sensch., Wieu, Math. Naturwiss. Classe, lvii, pt. I, p. 488
(Based on specimen from unknown locality, probably in Italy,
figured by Bonaparte, Iconogr. Faun. Ital., fasc. xxis, pi., fig. 5.)

18G9. [Sorex vulgaris'] var. nitda Fatio, Faune Vert. Suisse, i, p. 127
(Bernese Oberland, Switzerland).

SOEEX 43

1869. ISorcx vulgaris] vkr. 7ugra Fatio, Faune Vert. Suisse, i, p. 127
(Lucerne, Switzerland).

1900. Slorcx} vulgaris var. vel subsp. mollis Fatio, Rev. Suisse de Zool.,

VIII, p. 471 (Substitute for nigra).

1901. Sorex araneus alticola ililler, Proc. Biol. Soc, Washington, xiv,

p. 43, April 25, 1901 (Meiringen, Switzerland). Tj^pe in U.S.
National Museum.

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

1905. Crossojjus ou Sorcx ignotus Fatio, Arch. Sci. Phys. et Nat., Geneve,

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

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

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

Type in British Museum.

1910. Sorex araneus tetragonurus and S. aranezis carpathicus Trouessart,
Faune Mamm. d'Europe, pp. 52, 54.

Type locality. — Strassburg, Germany.

Geographical distribution. — Alp.s and neighbouring portions of
Germany, France and Italy ; eastward through Tirol to the
mountains of Transylvania. "

Diagnosis. — Similar to Sorex araneus hergensis, but colour in
summer pelage not so dark, the contrast between back and
sides often not consj^icuous, though back is frequently blackish ;
pigmentation of teeth more extensive than in the northern races,
the hypocone of m^ and m- usually brown at tip (see tabular
comparison with true araneus on p. 36).

Measurements. — Average and extremes of eight specimens
from the type locality : head and body, 75-8 (71-79) ; tail, 47-5
(45-50); hind foot, 13-5 (13 •2-14 -2). Average and extremes
of twenty specimens from Andermatt, Uri, Switzerland : head
and bod}', 73-1 (70-81); tail, 51-1 (50-55); hind foot, 13-3
( 1 3-14). Measurements of an adult female from Zermatt, Valais,
Switzerland (in alcohol) : * head and body, 74-6 ; tail, 51 ; hmd
foot, 13-6. Average and extremes of three specimens from
Vitznau, Switzerland : f head and bodv, 67-3 (62-72); tail,
46-3 (44-49); hind foot, 13-2 (12 -8-13 -6). Type oi Sorex
araneus alticola Miller (Meiringen, Switzerland,' No. 85,930
U.S.N.M.) : head and body, 76 ; tail, 55 ; hind foot, 14. Average
and extremes of ten specimens from Meiringen, Switzerland : t
head and body, 74-0 (70-77); tail, 52-5 (47-57); hind foot,
lo'3 (13-14). Average and extremes of four specimens from
Locarno, Ticino, Switzerland: head and body, 76-7 (76-82):
tail, 48-5 (45-50): hind foot, 13-6 (13-4-13"'-8). For cranial
measurements see Table, p. 49.

* Cotype of Sorex vtdgaris crassicaudatus Fatio.
t Essentially topotypes of Sorex vulgaris nigra and itwllis Fatio.
X Topotypes of Sorer araiieus alticola ililler and essentially topo'types
of Sorex vulgaris nuda Fatio.

44

INSECTIVORA

Specimens examined. — Two hundred and sixty-two, from the following
localities : —

Feance : Etupes, Doubs, 10; Barcelonnette, Basses-Alpes, 8; Cha-
monix, Haute-Savoie, 10 (U.S.N.M.).

Germany : Strassburg, 8.

SwiTZEBLAKD : Geneva, 17 (U.S.N.M. and Mottaz) ; St. Cergues, Vaud,
19 (U.S.N.M. and Mottaz) ; Chesieres, Vaud, 13 (Mottaz) ; Bioux-Dessus,
Vaud, 4 (Mottaz) ; Les Plans, Vaud, 4 (U.S.N.M.) ; Zermatt, Valais, 7 (B.M.,
U.S.N.M. and Geneva) ; Stalden, Valais, 2 (Geneva) ; Grindelwald, Bern, 4
(U.S.N.M.) ; Briinig, Bern, 9 (U.S.N.M.) ; Meiringen, Bern, 17 (U.S.N.M.) ;
Vitznau, Lucerne, 6 (B.M. and U.S.N.M.) ; Goschenen, Uri, 5 (U.S.N.M.) ;
Audermatt, Uri, 49 (U.S.N.M.) ; Hospenthal, Uri, 1 (U.S.N.M.) ; Ziirich, 1 ;
Murgsee region, St. Gallen, 16 (U.S.N.M.); Degersheim, St. Gallen, 3
(U.S.N.M.); Uzwil, St. Gallen, 1 (U.S.N.M.); Sitterwald, St. GaUen, 5
(U.S.N.M.) ; Ziiberwangen, St. Gallen, 5 (U.S.N.M.) ; Wildlnrchli, Appen-
zell, 1 (U.S.N.M.) ; Albulapass, Grisous, 1 (U.S.N.M.) ; Untervatz, Grisons,
5 (U.S.N.M.); Grisons, no exact locality, 4 (U.S.N.M.); Faido, Tioino, 3
(B.M. and U.S.N.M.); Lugano, Ticino, 1 (U.S.N.M.); Gentilino, Ticino, 1
(U.S.N.M.) ; Locarno, Ticino, 4 ; no exact locality, 1.

AusTKiA-HuNGARY : Hatszeg, Hunyad, Transylvania, 10 ; Csalloku'z-
Somorja, Pressburg, Hungary, 3 ; Schwaz, Tirol, 2 (U.S.N.M.).

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

8.^,2 9.

Etupes, Doubs, 350 m.
France. (C. Mottaz).

0. Thomas (p).

8. 8. 10. 14-24.

3 6.

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

0. Thomas (v).

8. 8. 10. 25-27.

4 6, 1 9.

Strassburg, Alsace.
(C. Muttaz.)

0. Thomas (p).

8. 8. 10. 28-32.

lal.

Zermatt, Valais, Swit-

Dr. J. Anderson

91. 10. 15. 2G.

zerland.

(c & P).

3 6.

Vitznau, Lake Lucerne.

0. Thomas (c & p).

5. 8. 2. 82-34.

1 al.

Zurich.

C. Mosch (p).

89. 11. 8. 3

9.

Faido, Ticino.

0. Thomas {c & p).

5. 8. 2. 19.

2 i5' 2 9.

Locarno, Ticino.-

0. Thomas (c & p).

5. 8. 2. 2-5.

9.

Switzerland.

E. E. Alston (p).

79. 9. 25. 10.

8 6, 1 9.

Hatszeg, Hunyad, Tran-

C. G. Dauford (c).

3. 2. 2. 8-10.

sylvania, 1500-2000£t.

3. 11. 8. 15.

Hungary.

3.

Csallokdz-Somorja,

Budapest ^Museum

94. 3. 1. 22-25,

Pressburg, 400 ft.

(E).

Hungary.

2al.

Vallombrosa, Florence,

Dr. G. Cecconi

1. 8. 2. 2-3.

Italy.

(c & P).

80REX ARANEUs pYREXAicus Miller.

1909. Sorex araneus injrenaicns Miller, Ann. and JIag. Nat. Hist., 8th ser.,

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

1910. Sorex araneus pyrenaicus Trouessart, Faune Mamm. d'Europe, p. 53.

Type localiti/. — L'Hospitalet, Ariege, France. Altitude
4,700 feet.

Geographical disirihution. — Pyrenees. At present known
from France only, though occurring on the south slope of the
mountains in the Department of Pyrenees-Orientales.

Diagnosis. — Very similar to Sorex araneus tetragonurus, but
distinguishable by the duller, less evidently tricolored summer

SOREX 45

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

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

Skull and teeth. — As in S. araneus tetragonurus.

Measurements. — External measurements of type : head and
bod\', 72; tail, 51 ; hind foot, 14. Average and extremes of six
specimens from the type locality : head and body, 70*6 (69-72) ;
tail, 47 (44-4-51); hind foot, 13-3 (13-14). Average and
extremes of nine specimens from Bareges, Hautes-Pyrenees : head
and body, 72-5 (69-75); tail, 45 (42-49); hind foot, 13-3
(13-14). For cranial measurements see Table, p. 50.

Specimens examined. — Thirty-two, from the following localities in the
French Pyrenees : —

Porte, Pyren^es-Orientales (Spanish watershed), 9 ; I'Hospitalet, Ariege,
12 ; As-les-Thermes, Ariege, 2 ; Bareges, Hautes-Pyr6n6es, 9.

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

c$, ?. Port6, Pyr6n6e8-Orien- G. S. Miller (c). 8. 8. 4. 142-143.

tales, 1600m. France.
2 i, 9. Porte, Pyrenees-Oricn- 0. Thomas (p). 8. 9. 1. 49-51

tales, lGOO-1700 m.

{A. Bobcrt).
6,4:9. L'Hospitalet, Ariege, G. S. Miller (c). 8.8.4.151-154.

4700 ft. 301.

(8. 8. 4. 301. Type of subspecies.)
?. Ax-les-Thermes, Ariege, G. S. Miller (c). 8. 8. 4. 141.

2600 ft.
6, 6 9. Bareges, Hautes-Pyr6- G. S. Miller (c). 8. 8. 4. 144-150.

uees, 1300-1500 m.

SOREX ARANEUS FRETALIS Miller.

1909. Sorex araneus frctalis Miller, Ann. and Mag. Nat. Hist., 8th ser., iii,

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

1910. Sorex araneus fretalis Trouessart, Faune Mamm. d'Europe, p. 52.

Type locality. — Trinity, Jerse\', Channel Islands.

Geographical distribution. — Known only from the island of
Jersey.

Diagnosis. — Like Sorex araneus araneus, but skull with
rostral portion shortened, broadened and deepened, and anterior

46

INSECTIVOKA

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48

INSECTIVORA

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Germany : near Sneekoppe, Riesen
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S. araneus castaneus.

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Cromlix

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52

INSECTIVOEA

teeth (incisors both above and below, and first and second ujjper
unicuspids) enlarged.

Colour. — Siiramer pelage about like that of S. araneus araneus,
but upper parts perhaps not so dark. Sides apparently averaging
lighter than in true araneus, and underparts in some specimens
a very pale, almost whitish, bufiy grey, decidedly lighter than in
any skins of the other races yet examined.

SJchU. — In size and general form the skull does not difier
appreciably from that of true Sorex araneus, but on comparison
of specimens the rostral portion in front of large premolar is
seen to be relatively shorter, broader and deeper. The greater
bi'eatlth is especially noticeable from below, the space between
the anterior unicuspids about equalling diameter of these teeth,
while in ail the other races (except (S'. o. granarius) it is evidently
less.

Teeth. — While in general the teeth resemble those of Sorex
araneus araneus, the large incisors both above and below are
appreciably more robust, and the first and second upper uni-
cuspids are wider. The difference is particularly noticeable in
the lower incisor, the shaft of which is thickened and the lobes
on cutting edge distinctly enlarged.

Measurements. — External measurements of tyjje (adult female) :
head and body, 63; tail, 48-2; hind foot, 13. For cranial
measurements see Table, p. 51.

Specimens exmnined.— Five, all from the island of Jersey.

Bemarhs. — The Jersey shrew is easily distinguishable fi'om
the other races by its enlarged anterioi' teeth. It is probable
that a sufficient series of skins will show that there is an average
difference in colour as well.

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

{I!. H. Bunting.) (8. 9. 2. 1. Type of subspecies.)

Sorex araneus oranarius Miller.

1910. Sorex araneus granarius Miller, Ann. and Mag. Nat. Hist., 8th ser.,
VI, p. 459, November, 1910. Type in British Museum.

Ti/pc localiiij. — La Gi'anja, Segovia, Spain.

Geographical dislrihuiion. — At present known from the type
locality only, and probably confined to the mountains of central
Spain.

Diagnosis. — Smallest known European race of Sorex araneus,
the condylobasal length of skull onlj^ about 17 "5 mm., upper
tooth-row about 7 ■ 5 mm. ; palate wide anteriorly as in S. araneus
fretalis, but anterior teeth not enlarged.

External characters and colour. — Externally the animal shows
no special peculiarities, though the ear and the plantar tubercles

SOREX 53

seem to be relatively smaller than usual. Colour of specimens
in alcohol apparently as in true aranctts.

Skull and teeth. — Apart from its small size the skull does not
differ noticeably from that of the other races, except in the
relative shortness of the rostral j^ortion and tooth-row as com-
pared with the breadth of palate. Palatal breadth between
anterior unicuspids as great in proportion to width of teeth as
in S. araneus f fetalis. Teeth small, normal in form, the pig-
mentation apparently less extensive than usual.

Measurements. — Type (adult male), and an older male, also
from La Granja : head and body, 62 and 66 ; tail, 36 and 37 ;
hind foot, 11 '6 and 11*6; ear from meatus, 6'6 and 7. For
cranial measurements see Table, p. 51.

Specimens examined. — ^Two, both from the tj^pe locality.

Hemarlcs. — ^In its broad palate the (Tuadarrama shrew bears
a remarkable likeness to the form inhabiting the island of
Jersey. It is readily distinguishable from the Jersey animal
by its small size, and by the absence of all tendency to enlarge-
ment of the anterior teeth.

2 al. La Granja, Segovia, M.dela Escalera(c). 6.11.4.3-4.
Spain. (6.11.4.4. TyjJC of subspecies.)

SOREX MINUTUS Linnajus.
(Synonymy under subspecies.)

Geographical distribution. — Northern portion of Eurasia from
Ireland eastward (exact eastern limits of range not known). In
Europe south to the Pyrenees and southern Italy.

Diagnosis. — Size small, head and body usually about
50-60 mm., the tail 10-15 mm. shorter, condylobasal length
of skull 14 '8-16 "6 mm.; posterior lobe of anterior upper
incisor sub-terete, the length of its base about half that of
anterior cusp ; colour brown, the underparts always lighter
than back, and sides never specially contrasted.

E.iternal characters. — In general external characters Sorex
minutus agrees with S. araneus, except for its smaller size and
relatively longer tail.

Colour. — The colour rather closely resembles that of Sorex
araneus in dull summer pelage, except that the back usually has
a peculiar greyish cast not easy to describe, but by which it is
possible to recognise skins with much certainty. There is never
any indication of a specially differentiated colour area along sides.
U})per parts between sejjia and wotxl-brown in summei', more
nearly hair-brown in winter, the hairs slate-grey at ba.se and
with faint .silvery sub-terminal annulations more visible in some
lights than in others, and giving rise to the grej'ish effect already
alluded to. Underparts smoke-grey of varying depth, some-

54

INSECTIVORA

times almost whitish, the line of demarcation along sides usually
evident though not very conspicuous. Feet pale wood-brown
with a silvery gloss or suffused with drab, sometimes noticeably
lighter than back. Tail concolor with back above, not so dark
below.

SlcuU. — Apart from its conspicuously smaller size (condj'lo-

basal length, 14 '8 to 16 mm. instead of 17 "8

to 20 mm.) the skull differs notably from

that of Sorex araneus and S. nlpinus in the

narrower, more elongate brain-case, the outline

of which when viewed from above is distinctl)'

oval instead of sub-circular. Antero-external

jwrtion of border flattened, though less notice-

abh' than in Sorex araneus. Owing to its

different form the brain-case is less abruptly

Fio. 10. marked off" from interorbital region than in

Sorex minutus. *^^® other European species. Depth of brain-case

Nat. size. usually less relatively to breadth than in

S. araneus. Lachrymal foramen opening over

posterior half of first molar.

Teeth. — Aside from their very much smaller size (upper
tooth-row 6 to 7 mm. instead of 8 to 9-6 mm.) the teeth of
Sorex minutits differ in numerous details of form from those of
S. araneus. Anterior upper incisor with basal lobe nearly as
high as anterior cusp, so that the two points of the tooth are
essentially in line with the tips of the first three unicuspids.
AVhen viewed from below the posterior lobe appears, howe\er,
distinctly smaller than anterior cusp, while from the side its
outline is conspicuously higher and narrower than that of first
unicuspid. Anterior lower incisor with
first lobe on cutting edge as well defined
from anterior point of tooth as from second
lobe, and lengths of bases of all three lobes
approximately equal. Upper unicuspids not
essentially different in form from those of
Sorex araneus, but crowns perceptibly longer
than wide and less abruptly rounded off'
anteriorly, the resulting form less nearly ^"^" "'

T 1 J- 1 • J.1 J.T ■ -L Sorex miniitus. Anterior

square. In lateral view the outline is cjuite teetii in prutiie. x 5.
as in the larger animal. The relative size

of the unicuspids differs markedly from that in the larger
animal. The first, second and third are sub-equal, with the
second usually a trifle smaller than the third, and the first with
distinctly the greatest crown area. Fourth slightly smaller than
third, and fifth equally smaller than fourth'"', between which and
large premolar it is tightly crowded. First lower unicuspid
differing from that of S. araneus in its greater length along
cingulum and less height of cusjj, its form when viewed from
* Sometimes equal to fourth or slightly larger.

SOREX S5

the side thus noticeably different from that of first upper
unicusisid. Second unicuspid, together with other mandibular
teeth, essentially as in S. aranem. Maxillary cheek-teeth as in
S. araneus, except that hypocones are less developed, that on
large premolar obsolete.

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

Sorex minutus minutus Linnaeus.

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

Siberia).
1789. [6'orea-] exilis Gmelin, Syst. Nat., i, 13th ed., p. 115 (Yenesei River,

Siberia).
1806. Soi'ex canaliculatus Ljungh, Kongl. Vetensk. Akad. Nya Handl.,

XXV3I, p. ,263 (Lommaryd Vicarage, northern Vedbo district,

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

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

(Accidental renaming of minutus).
1832. Swex puviilio Wagler, Isis, p. 54 (Bavaria).
1838. S[orex] rusticus Jenyns, Ann. Nat. Hist., i, p. 423, August, 1838

"(England).
1838. Siorex] rusticus var. S S{orex} hihernicus Jenyns, Ann. Nat. Hist., i,

p. 423, August, 1838 (Dublin, Ireland).
1844. Sorex immilus Nilsson. Ofversigt af Kongl. Vetensk.-Akad. Forhandl.,

Stockholm, I, p. 33, March 20, 1844 (North-eastern Skaane,

Sweden).
1857. Sorex pygmmus Blasius, Siiugethiere Deutschlands, p. 133.
1895. Sorex minutus Thomas, The Zoologist, 3rd ser., xix, p. 63, February,

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

Type locaJiiij. — Vicinity of the Yenesei River, Siberia.

Geoijraphiral distribution. — The entire European range of the
species, except southern Italy.

Diagnosis. — Teeth normal in size, the molars and anterior
upper incisor not enlarged.

Measurements. — .Average and extremes of thirteen specimens
from Grantown-on-Spey, Elgin, Scotland : head and body, 52 • 3
(49-55); tail, 36 (32 -5-39 -5); hind foot, 10-4 (10-11).
Average and extremes of eight specimens from the Isle of ^lan :
head and body, 59-6 (52-64); tail, 40-2 (36-43): hind foot,
11-1 (10-12). Average and extremes of five specimens from
Ariege, France: head and body, 55-8 (51-62); tail, 44*2
(42-46): hind foot, 11-4 (11-12). Average and extremes of

56

IN'SECTIVORA

four specimens from the Harz Mountains, Germany : head and
body, 57-3 (51-63) ; tail, -40 -5 (39-42) ; hind foot, 11-5 (11-12).
Average and extremes of four specimens from Stalden, Valais>
Switzerland: head and body, 55-6 (50-61); tail, 42-6 (42-43);
hind foot, 11 "0 (10 "8-11 '2). Average and extremes of seven
specimens from Hatszeg, Hunyad, Transylvania : head and body,
51-3 (47-55): tail, 40-6 (32-46); hind foot, 11-2 (10-12).
For cranial measurements see Table, p. 58.

Specimens examined. — One hundred and twenty-five, from the following
localities ; —

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

Wales : Aberia, Merionethshire, 2 ; near I3ridgend, Glamorganshire, 1
no exact locality, 1.

England : Spurn Head, Yorkshire, 1 ; Grimsby, Lincolnshire, 3
Waltham, Lincolnshire, 2 ; Thornhaugh, Northants, 2 ; Swithland, LeiceS'
tershire, 1 ; Graftonbury, Herefordshire, 2 ; Crippetts, Gloucestershire, S
Clifton, Gloucestershire, 1 ; Loughton, Essex, 2 ; Wilbraham, Cambridge-
shire, 1 ; Dartford, Kent, 1 ; New Forest, Hampshire, 1 ; Alum Bay, Isle
of Wight, 1 ; Chagford, Devonshire, 2 ; Combmartin, Devonshire, 1 ; South
Molton, Devonshire, 1 ; Lundy Island, Devonshire, 1 ; Isle of Man, 8.

Ireland : Clonbroch, Co. Galway, 1 ; Caragh Lake, Co. Kerry, 1 ; Co.
Longford, 1 ; Knock, Co. Down, 1 ; North Esk, Dunkettle, 2 ; Duncannon,
3 ; Kilkenny, 1 ; no exact locality, 2.

Norway : Holaaker, upper Gudbraudsdal, 1 ; Graven, Hardanger, 1
(U.S.N.M.).

Sweden : Upsala, 1 (U.S.N.M.) ; near Stockholm, 1 ; Svarta, Orebro, 1
(U.S.N.M.).

France ; Barbizon, Seine-et-^Iarne, 1 ; Montr^jeau, Haute-Garonne, 1
(U.S.N.M.); Ax-les-Thermes, Ariege, 2; I'Hospitalet, Ariege, 3; Porte,
ipyren^^es-Orientales, 1.

Germany: Brunswick, 3 (B.M. and U.S.N.M.) ; Bodethal, Harz Mts., 3
(U.S.N.M.); Mauseklippe, Bodethal, Harz Mts., 1 (U.S.N.M.); Frankfort-
on-Main, Hessen-Nassau, 1 ; Niesky, Silesia, 1 ; Marxheim, near Mannheim,
Bavaria, 2 ; near Konigsberg, 2 (U.S.N.3M.).

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

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

Italy: Vallombrosa, near Florence, 1.

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

9. Dunblane, Stirlingshire. W. R. Ogilvie-Grant 8.9.0.1.

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

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

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

(0 & P).
1 al. North Uist, Hebrides. J. A. Harvie Brown 79. 9. 18. 1.

(c & p).
1. Barra Island, Hebrides. W. E. de Winton 8. 9. 20. 1.

(c & P).

57

6. Stockbriggs, Lanark-
shire.

1. Bridgend, Glamorgan-

shire, Wales.

1 al. Wales.

9. Grimsby, Lincolnshire,

England.

2. Waltham, Lincolnshire.

2 st. Graftonbury, Hereford-
shire.

9. Wilbraham, Cambridge-

shire. (S. F.Harmcr.)

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

6. Lundy Island, Devon-
shire.

2 6. Chagford, Devonshire,

85Gft. (W.P.Stark.)
6, 4 9. Ramsay, Isle of Man.

6. Sulby Glen, Isle of Man.

9. Clonbroch, Galway, Ire-

land.

1 al. Longford.

6 St. Knock, Down.

2 al. Dunkettle, Cork.

2 6,19. Duncannon, Wexford.
(Rev. Dr. Martin.)

1 al. Ireland.

6. Holaaker, Gudbrandsdal,
1900 ft. Norway.

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

2 9. Ax-les-Thermes, Ari^ge.
6. V Hospitalet, Ariege,

4700 m.
6. r Hospitalet, Ariege,

1450 m. (A. Robert.)
9. Port6, Pyr6n6es-Orien-

tales, 1600 m.

2. Querum Forest, Bruns-

wick, Germany.
1 al. Frankfort - on - ]\Iaiu,
Nassau.

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

6. IMarxheim, Bavaria.

1. Csall6k6z-Somorja, Press-

burg, 400 ft. Hungary.

1. Hatszeg, Hunyad, Tran-
sylvania, 2000 -
6500 ft.

3. Hatszeg, Transvlvania,

2000-6500 ft."

3 6. Hatszeg, Hunyad, 1600-

2000 ft.

E. R.Alston (c&p).

R. I. Pocock (c & p).

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

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

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

(c & p).
Miller Collection.

Miller Collection.

N. H. Joy (c & p).

Miller Collection.

C. H. B. Grant
(c & p).

C. H. B. Grant
(c & p).

H. P. Witherby
(c & p).

Dr. G. E. Dobson
(c & p).

Hon. N. C. Roths-
child (c&p).

Miss O'Keefe (c&p).

G. Barrett-Hamilton

Zoological Society's

Collection.
R. J. Cuuinghame

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

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

O. Thomas (p).

G. S. Miller (c).

G. Barrett-Hamilton

Tomes Collection.

Lord Lilford (p).

Lord Lilford (p).
Budapest Museum

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

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

79. 9. 25. 13.

8. 9. 6. 1.

48. 9. 24. 3.
8. 9. 16. 1.

11. 1. 2. 81-82.

96. 4. 28. 20-21.

7. 7. 7. 2850.

7. 7. 7. 3604.

6. 5. 16. 1.

7. 7. 7. 4472,
4474.

8. 9. 7. 1-5.

8. 9. 8. 4.
8. 9. 19. 1.

80. 12. 14. 6.

I. 9. 3. 7.

80. 11. 20. 4.

81. 6. 29. 1.

II. 1. 2. 83-85.

52. 9. 13. 288.

98. 2. 28. 5.

8. 8. 4. 160.

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

8. 9. 1. 45.

8. 8. 4. 161.

11. 1. 2. 86-87.

7. 1. 1. 46.

99. 1. 9. 15.

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

3. 2. 2. 11.

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

58

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60

IXSECTIVOKA

1. St. Gothard, Uri, Swit- Purchased (Brandt). 46. 2. 13. 14.
zerland.

1 al. Vallombrosa, Florence, Dr. G. Cecconi (p.) 1. 8. 2. 4.
Italy.

SoREx MixuTUS LUCANius Miller.

1909. Sorex niinutus Incaniits Miller, Ann. and Mag. Nat. Hist., 8tli ser.,

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

1910. Sorcx minutiis lucanms Trouessart, Fauna Mamm. d'Europe, p. 55.

Type locaUty. — Monte Sirino, Lagonegro, Italy.

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

Diagnosis. — Similar to Sorex minutus minufus, but with molars
and anterior upper incisor noticeably enlarged

Teeth. — As compared with specimens of Sorex minutus minutus
from a wide range of European localities and also with specimens
from Scalipa and Sumela, Asia Minor, the type of S. minutus
Iwanius is at once recognizable by its enlarged teeth The actual
size is so small that it is impossible to express the differences by
measurements, but to the e3'e the greatei- area of the molar
crowns and the larger more projecting anterior upper incisor
and more robust unicuspids and anterior lower incisor are at
once apparent.

Measurements. — Tail, 42; hind foot, 10 '4. For cranial
measurements see Table, p. 59.

Specimen examined. — The type.

1. IMonteSirino, Lagonegro, 0. Thomas (p). 8. 9. 1. 5.

S. Italy. (.4. Robert.) (Ti/pe of subspecies.)

SOREX ALPINUS Schinz.

(.Synonymy under subspecies.)

Geographical (Jistrihution. — Pyrenees, Alps, Harz Mountains,
Riesengebirge and Carpathians.

Diagnosis. — Size essentially as in Sorex aranvus (head and
body usually 65 to 70 mm.), but tail about equal to head and
body ; colour uniform dark slaty grey, the underparts nearly as
dark as back ; basal lobe of anterior upper incisor sub-terete, the
length of its base about half that of antei'ior lobe.

External characters. — Except for the dark, slaty colour, and
the much greater relative and actual length of the tail, Sorex
alpinus does not differ markedly from Sorex araneus in general
external characters. The fur of the back varies from 5 mm. to
8 mm. in de^^th according to season, and the tail may either be
well covered with closely appressed hairs and provided with a
pencil 4 to 5 mm. long, or practically naked at tip and ver}-

SOREX 61

scantily haired elsewhere, a variation not wholly dependent on
season.* At middle of tail there are about 18 annulations to the
centimeter. Feet relatively larger than in Sorex araneus, but
not peculiar in form.

Colour. — Entire upper parts a deep slaty grey produced by
the combination of blackish-slate under colour and short seal-
brown tips, lightened to a varying degree by silvery sub-terminal
annulations. Underparts slightly washed with sepia, but the
difference in colour of the two surfaces not conspicuous, and sides
entirely without line of demarcation. Feet scantily clothed with
silvery hairs. Tail sharply bicolor, nearly black above and at
tip, buffy whitish below, the light area always less extensive than
the dark, and sometimes reduced on distal half of tail to a
narrow median line.

SJcidl. — The skull, though of the same general dimensions as
that of Sorex araneus, is recognizable by its
less elevated brain-case, the outline of which
when viewed from above is more nearly
circular, owing to the very slight flattening
of antero-external portion of border. Inter-
orbital region less tapering than in the
related species, a character due to the
slightly though appreciably greater lachrymal
breadth. The diameter of the tympanic ring
is usually though not constantly less than in
Sorex araneus. Anteorbital foramen slightl}'
larger than in S. araneus. Lachrymal fora-
men opening over point of contact between .
first and second molars.

Teeth. — Kelatively to size of skull the teeth
are noticeably smaller than in Sorex araneus; this is particularly
evident in the anterior upper incisor when viewed from the side,
and the upper unicuspids when viewed from below. Anterior
upper incisor with basal lobe even smaller than in S. miniitus, its
height slightly less than that of first unicuspid and much less than
that of anterior lobe of incisor. Viewed from below it appears
to occupy decidedly less than half of tooth, while from side it
appears scarcely half as large as first unicuspid, from which it
further diflers in its simple peg-like form. The two teeth come
in contact at about the same level as in S. araneus, but their tips
diverge more noticeably. Anterior lower incisor with shaft
distinctly tapering, its cutting edge with three low, sometimes
ill-defined lobes. Upper unicuspids less robust than in Sorex
araneus, their crowns distinctly longer than broad. The cusp
occupies more than half area of crown, at expense of crushing
area, most of which, except in fifth unicuspid, lies obliquely
instead of horizontally. Viewed from the side the unicuspids

* A specimen in full winter coat, but with the tail bare at tip (B.M.
3. 2. 2. 2), was taken at Hatszeg, Hunyad, Hungary, on December 12.

62

INSECTIVOEA

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

, . ■ ' . peculiarities of form. The first, how-

Sorex ahnnus. Anterior ^ . ..... ,.,,, , i. , i

teeth. X 5. ever, IS strikingly dinerent irom the

corresponding tooth in S. araneus.
Its general outline when viewed from side is irregularly
elliptical, with longest axis parallel to that of mandible, the
upper edge with two low, rounded cusps, the anterior of which
is about as large as lobes on cutting edge of incisor, the posterior
smaller. Upper cheek-teeth not essentially difl^erent from those
of Sorex araneus, except that hypocones are less developed.
Lower cheek-teeth as in the related species. Pigmentation of
teeth slightly less extensive than usual in Sorex araneus.

Sorex alpinus alpinus Schinz.

1837. Sorex alpinus Schinz, Neue Denkschr. Allgem. Scbweiz. Gesellsch.

Naturwiss., Neuchatel, i, p. 13 (St. Gothard Pass, Switzerland).
1840. ? Sorex antinorii Bonaparte, Iconogr. Faun. Ital., i, fasc. 29 (No exact

localit}-, and probably not a European species).
1857. Sorex alpinus Blasius, Saugetbiere Deutscblands, p. 126.
1870. ? Sorex intermedins Cornalia, Catal. Descrit. IMamm. Ital., p. 27 (Hills

of Brianza, Uomo, Italy). Part : body ; see Sordelli, Atti Soc. Ital.

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

Mus. Civ. Stor. Nat., Milano, xx^viii, p. 363 (MS. .synonym of

intermedius) .
1910. Sorex alpinus Trouessart, Faune Mamm. d'Europe, p. 50. ^

Type locality. — St. Gothard Pass, Uri, Switzerland.
Geoijrapliical distrihution. — From the Jura and Alps through
Tirol to Transylvania ; Pyrenees. f

* In an adult female from Briinig, Switzerland (No. 85830, U.S.N.M.),
tbe tbird and fourtb unicuspids on rigbt side are fused into a single
2-cusped tootb, wbile tbose on left side are normal.

t I bave seen no Pyrenean specimens of Sorex alpinus. For record of
its occurrence see Trutat, Bull. Soc. Hist. Nat., Toulouse, xii, p. 100, 1878
("massif de la IMaladetta '"). Tbe animal is probably less common in tbe
Pyrenees tban in tbe Alps, as I was unable to find it in several localities
resembling tbose wbere it regularly occurs in Switzerland.

SOREX 63

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

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

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

Switzerland : St. Cergues, Vaud (Jura), 4 (U.S.N.M. and Mottaz) ;
Briinig, 5 (U.S.N.M.); Vitznau, 4 (B.M. and U.S.N.M.); Andermatt, 1
(U.S.N.M.) ; St. Gothard, Uri, 6 (B.M. and U.S.N.M.) ; Untervatz, Grisons,
1 (U.S.N.M.) ; Murgthal, St. Gallen, 1 (U.S.N.M.) ; Santis, St. Gallen, 1
(Mottaz) ; no exact locality, 3.

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

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

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

Longchamps (p).
1, 1 al. St. Gothard, Uri. Purchased (Brandt). 46. 2. 13. 18.

46. 2. 13. 14.
1. St. Gothard, Uri. Purchased (Stock- 46. 6. 2. 32.

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

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

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

sylvania, 2000 ft.
Hungary.
i, 2 9. Hatszeg, Hunyad, Tran- C. G. Danford (c). 3. 11. 8. 12-14.

sylvania, 1500-2000 ft.

SoREX ALPINUS HERCYNICUS Miller.

1909. Sorc.x: alpinus heixynicus Miller, Ann. and Mag. Nat. Hist., Sth ser.,

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

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

Tjipe localitij. — Miiuseklippe, Bode Valley, Harz Mountains,
Germany.

(Teograpliical distrihntion. — Harz Mountains and Riesen-
gebirge.

Diagnosis. — Skull and teeth not so large as in typical
SorcK alpinus (condylobasal length of skull, 19 to 19 "6 mm.;
upper tooth-row, 8"2 to 8*6 mm. ; lower tooth-row, 7-8 to 8'0 mm.).

Measurements. — ■External measurements of type : head and
body, 71, tail, 67 ; hind foot, 15*4. Average and extremes of
six specimens from the Harz Mountains : head and body, 73 • 4

64

INSECTIVORA

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O O O O O O O O O CT O C3 C5 03 0> C5 03 O C5 03
0'llMC<IC^<M(MOq<MCMiHOqi-l T-I.Hi-lTHr-tr-(,Ht-l

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O+O^CH-OOO+CH-OO+OOO* 000-CH-'-0*-o'0*XD

&

inrHCMOOi-H^_r/-i'-,r-)^c-i lOCOOC-l^t-COOl

o lO o in CO CO & S S S ?o ^ 0-1 fM CO on CI 0-1 c<i c-1

-iH -H -41 -* -^H -* S S S S S S C-1 01 CM CM C-1 C-1 C-1 0-1

S33?13?3wSS§SS ;:1 ;=! S ^; ;il ;:J ^1 ;:j

Locality.

^ . ... .2

S. alpinus alpinus.

France : Chamonix
Switzerland : St. Ccrgues, Vai

Vitznan
Briiuig

Andermatt

S. alpinus hercynicus.

Germany : Wolfshau, Silosia

Eulengrund .

Bahrcnberg, Harz
Miiuseklippe ,

65

(72-77); tail, 66-6 (6c-68) ; hind foot, 15-1 (14-S-I5-8)
Average and extremes of five specimens from the Riesen-ebir-e •

ui i'lf.R^r\P'i ^''~"\' ^^^^ ^'^'^ ^•^^■'^^) ' '"^d fo^^t,"
1* J (14 a-lo-U). ior cranial measurements see Table, p. 64.

Spechnens examined. -Eleven (all in U.S.N.M.), from the foUowin-
localities m Germany: Bahienberg, Harz Mts., 2; Bodethal, Harz
w iVi' ^^^^^'^'^l'PP'r.HarzMts.,3; Eulengrund, Riesengebirge, Silesia 3
Wolfshau, Riesengebirge, Silesia, 2. 6 ^^ 'e«. '^uebid, o ,

BemarJc^— In external measurements the Alpine shrew of
the Harz MountaiiLs and Riesengebirge agrees with the Swiss
animal ; but the differences in length of skull and of tooth-rows
seem enough to warrant the recognition of the two forms as
distinct.

Genus NEOMYS Kaup.

1829. Keomus Kanp, Entw.-Gesch u Natiirl. Syst. Europ. Thierwelt, i,
p. 117 (boicv daiibcntomi Erxleben).

1829. LeiccorhyncMcsK^ni,, Entw.-Gesch. u. Natiirl. Syst. Europ. Thier-
Th!, fi'' f • I" ('^•^'•^'^' ^^«^«i"^ Geoffroy and ,S'. Icucodon Hermann •
p lotMatHsol).*""' '" Thomas, The Zoologist, 4th ser., n',

1829. Hydrogalc K^n^, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt i

p. 119 {Sorcx remifer Geof!roy). ^ J-nierweu;, i,

1832. Crossopus Wagler, Isis, p. 275 [Sorex fodiens Schreber)
1835. i7y^ro.or.r Duvernoy, M6m Soc. du Mus. d'Hist. Nat., Strasbourg,

II, p. 19 (&ore.i:/ofZjcws Pallas, see p. 17). °

1835. ^«y./»sor.,r Duvernoy, M^m. Soc. du Mus. d'Hist. Nat., Strasbourg

animal).^ P^r'" " ^ ' ^^""^^ "*" ^°'''' '"''"'''"' tetragonurits
1838. Pinalia Gray, Proc Zool. Soc, London (1837), p. 12G, June 14, 1838

(MS. synonym of Crossop^ts).
1848. Galeviijs Pomel, Archiv Sci. Phys. et Nat., Geneve, ix p 249

November 1848 part, included £V«cfe,/4o,-,,, Duvern^y, cX'ssopus

Wagler, and Pachyiua do Selvs-Lougchamps) ^'o.sopus

1854. Myosidis Pomel Catal. M^.th. Vert. Foss. Bassin de la Loire p 14

/S.)°'' "" ''°' distinguished specifically from Nelmys

1857. Crossopus Blasius, Saugethiere Deutschlands, p. 119
1898. Neomys Thomas, Zoologist, 4th ser., ii, p. 100, March 15, 1898.

Ti/pc species.—Sorex daubentonii Erxlehen = S. fodiens Schreber
(geographical distribntion.—I'ali^.iivctic region north of the
Mediterranean from Spain and England to the Caucasus and
Asia Minor, north to northern Scandinavia.

Characters.— Uppev unicuspid teeth 4-1, their form more pre-
Uensory than in Sorex; dental formula : / ;'=i\ t- tj, jjih --=?, vi ?== = 30 •
posterior lobe of anterior upper incisor 'less 'than' lillf 'as hi-h
as hrst ; anterior lower incisor with one ill-defined lobe on cuttin-
edge near middle of tooth ; third lower molar and second lowe?
unicuspid as in Sorev ; points of all the teeth pigmented; fur
unusua ly soft and den.se, entirely concealing ears ; feet with a
noticeable fringe of elongated hairs at edges of soles and toes

F

66

INSECTIVORA

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

Semarlcs. — In its external form, slight reduction in the
number of teeth, and in the noticeably prehensory adaptation
of the upper incisors and unicuspids, the members of the genus
Neomys are distinctly more specialized than the species of Soreoc.
They retain, however, the primitive form of the third lower
molar and lower premolar characteristic of Sorcx, these teeth
showing no tendency to the more highly modified structure
found in Crocidura. The members of the genus Neomi/s are the
only Europenri shrews specially adapted to aquatic life. They
are at once recognizable Iw their dense, velvety fur, and large,
distinctly fringed hind feet. In the commonest and most widely
distributed species the tail is provided with a keel of stiffened
hairs, a character not found in any other Eurojiean shrew.

KEY TO THE EUROPEAN FORMS OF NEOMYS.

Tail with median keel of stiffened hairs extending

entire length of under surface N. fodicns, p. 66.

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

Peninsula) S.f.fodienx, p. 69.

Underparts usually brownish (England and Scot-
land) y.f- Mcoloi; p. 73.

Tail without keel.

Length of tail 47 to 53 mm. ; hind foot 16 to

17 mm. (Alps and Pyrenees) N. milleri, p. 78.

Length of tail 55 to 61 mm.; hind foot 16-8 to

18 mm. (Northern and central Spain) K. anomalus, p. 81.

NEOMYS FODIENS Schreber.

(Synonymy under subspecies.)

Gco[irapMral (listribiition. — From Norway and England to the
Pyrenees, northern Italy and western Siberia. ISTorthern and
eastern limits of range not known.

Diagmi^is. — Tail with keel always present ; feet Aery con-
spicuously fringed ; hind foot usually more than 1 7 mm. ; lachrj'-
mal foramen (.)pening over ^^osterior half of m^ ; anterior upper
incisor robust.

E.iternal cliar(tcfert<. — Fur very dense, its depth at middle of
back about 6 inm. in summer, 9 mm. in winter, its texture soft
and velvety, this ji'^i'ti^-'idarly noticeable on underparts : a
few longer hairs usually present on flanks and rump. Eyes
small and inconspicuous ; ears completely hidden by the fur.
Muzzle less slender than in Sorea- araneiis, the median ridge on
upper lip with distinct wart-like outgrowth which tits between
points of ujjper incisors when mouth is closed. Feet larger than

NEOMYS

67

more widely spaced ardbette;- defined Tf ■'" T''7' ^"'

pencil evident, utally ab™ 3 to°f ° ''™'''"f'='- 1' ■"^l" ^
median li„e «t tail bcLt frL extrameTaie'l r°"'\ "^'""^

and on head mssin<- ilf-Lr ''f ""'"j''""" everywhere evident,

size''!:ht^Iw t"'' ^'^'" ^™^" ^'^^^ «^ ^-- — '"'^ in larger
size leldtnely larger, more inflated brain- "

case (depth decidedly more than half breadth
and lower, more flattened rostrum and
interorbital region. The brain-case is sub-
circular m outhne, though distinctly flattened
antero-externally and postero-exteLallv-in

rSnl-'' ""^"^"'^ °^ '"^^ interJrbital
region by a conspicuous, well deflned anc^le
lu ateral view the proflle of brain-case^ is
S . '''''''''^f posteriorly, low anteriorly
o s ly abrupt curve into the nearly flat dJrsal

V ."'l> iT*"'""'^'*"' '"^Sion and rostrum.
Anteorbital foramen relatively larger and more
Midelv open than in the European species of
hnlTnf ^^"\7"^^^ f^'-'-^n^en over posterior
half of u\ i^Jesopterygoid space decidedly more than half ..
-Ida as long; hamular processes shorter than in^W arteTs

KlQ. 14.

yeomys fodiens.
-Nat. size.

68 INSECTIVORA

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

Teeth. — Though in general not strikingly different from those
of Sorex araneus the teeth show a tendency towards greater
elevation of the cusps as well as a more pronounced hooking
forward of the anterior upper incisor, peculiarities which impart
to the entire dentition, though j^articularly to the teeth lying
in front of the large premolars, an unmistakably prehensive
character. Anterior upper incisor with main hook more projecting
than in Sorex araneus ; posterior talon low though rather long,
its outline when viewed from the side somewhat triangular, its
height less than half that of main cusp and much less than that
of first unicuspid. Lower incisor more slender than that of
Sorex araneus, its cutting edge with only a single low, ill-defined
lobe near middle. Upper unicuspids large and strong, their
crowns when viewed from below decidedly longer than wide,
truncate posteriorly, narrowed an-
terioi'ly and terminating in a slight
though evident point. Jn lateral
view the cusps appear to be more
slender and set further forward
than in Sorex araneus, and the
posterior part of crown is more
produced backward, the cingulum
often rising to a distinct postero-
internal angle or rudimentary cusp,

particularly in first tooth. Spaces
Neomus fodiens. Anterior teeth t , ,. n . . , ,

in profile, x 5. between tips ot unicuspids greater

than in the European species of
Sorex. Each cusp is nearly terete, but with a small postero-
internal concavity continuous with the concave crushing area
which occupies posterointernal fourth of crown. The con-
cave area of cusja and crown is bounded externally by a
low but distinct ridge extending to middle of posterior border
of crown. First and second unicuspids sub-equal ; third about
half as large as first or second though quite similar to
them in form ; fourth not half as large as third, in tooth-
row or crowded somewhat inward, always distinctly visible
from outer side, in contact posteriorly with large premolar.
Lower unicuspids relatively longer and lower than in Sorex
araneus, their form when viewed from the side much as in
Sorex alpinus, except that the first has a very rudimentary
posterior cusp. First somewhat more than half as large as
second, its general outline much the same as in corres-
ponding upper teeth. Second essentially as in Sorex araneus.
Larger cheek-teeth as in Sorex araneus, but main cusps higher
and hypocones better developed. Pigmentation of teeth about

NEOMYS 69

as in Sorex nraneus araneits, the hypocones usually if not always
white.*

Measurements. — In external measurements this species is
unusally variable, the head and body ranging from 72 to 96 mm.,
tail from 47 to 77 mm., and hind foot from 16 to 20 mm. These
differences appear to be for the most part strictly individual.
The skull is more constant : condylobasal length, 1 9 '6 to 22 • 2 mm.;
upper tooth-row, 9*6 to 11 mm. Here the variation is to a
certain extent geographic, as the skull of the British race
averages smaller than that of the Continental form.

SemnrJcs. — Neomys fodiens is the common and widely dis-
tributed water-shrew of Europe. It may always be recognized
by its large size, fringed feet and keeled tail. While the keel
varies greatlj' in depth and distinctness, some trace of it is
always present on basal half of tail, the region in which the
median hairs are never modified in the round-tailed members
of the genus.

Neomys fodiens fodiens Schreber.

1776. Sorex aquaticus P. L. S. Miiller, Natursyst. Suppl. u. Regist.-Band,

p. 36 (Prance ; based on Buffon, viii, pi. 11). Not Sorex aquaticus
Linnseus, 175S.

1777. Sorex fodiens Schreber, Saugthiere, iii, p. 571 (Berlin, Germany).
1777. [Sorex] dauhentonii Erxlebeu, Syst. Regni Amm., i, p. 124 (Burgundy,

France ; based primarily on the Musaraigne d'eau of Daubenton,
Hist, de I'Acad. Roy. des Sci., Paris, 1756, p. 42).
1780. Sorex carinatus Hermann in Zimmermann, Geogr. Gesch., ii, p. 382
(Strassburg, Germany).

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

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

1793. [Sort\r] fluviatilis Bechstein, Gemeinn. Naturgesch. Deutschlands,

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

synomym of Sorex eremita, perhaps from Bechstein MS.).
1800. S[orcx] flodiens'] alhus Bechstein, Thomas Pennant's Allgem. Uebers.

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

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

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

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

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

but not named bv Geoffroy, M^m. Mus. d'Hist. Nat., Paris, I,

p. 309, 1815.

* In an adult male from Geneva, Switzerland (No. 104G Mottaz), the
right upper incisor and first unicuspid and right lower incisor are white
throughout. All other teeth normal.

70

INSECTIVOKA

1822. Sorcx inacrourus Lehmanu, Observ. Zoologies in fauoam Hamburg-

ensem, i, p. 5 (Sachseuwald, near Friedrichsruh, Schles\Yig-Holstein,

Germany).
1826. Sorcx aniphibins Brehm, Ornis, ii, p. 44 (Renthenclorf, Thiiringen,

Germany).
1826. Siyrex nutans Brehm, Ornis, ir, p. 44 (Renthenclorf, Thiiringen,

Germany).
1826. Sorex stagnatilis Brehm, Ornis, ri, p. 47 (Renthenclorf, Thiiringen,

Germany) .
1830. Sorex rivalis Brehm, Isis, p. 1128 (Renthenclorf, Thiiringen, Germany).
1832. Sorex musculus Wagler, Isis, p. 54 (Bavaria, Germany).
1882. Sorex psilurus Wagler, Isis, p. 54 (Bavaria, Germany).

1834. S[ore,r] nigripcs JMelchior, Den Danske Stats og Norges Pattedyr,

]}. 68 (Sielland, Denmark).

1835. Sorex fodu'iis Duvernoy, ilem. Soc. du ilus. d'Hist. Nat., Strasbourg,

II, p. 17. Part : animal, not skull.
1835. Sonw hcrmanni Duvernoy, !\Iem. Soc. du ilus. d'Hist. Nat., Stras-
bourg, II, p. 23 (Strassburg, Germany). Part : skull, not animal.

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

1838 (North Bothnia, Sweden).

1839. \ Sorcx fodiens] var. Icucotis de Selys-Longchamps, Etudes de Micro-

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

Blanc, Haute-Savoie, France).
1839. ? [Sorcx fodiens] var. albiventris de Selys-Longchamps, Ktudes de

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

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

nudum.
1857. Crossopus fodiens Blasius, Saugethiere Deutschlands, p. 120 (part).
1868. Sorex timbriatiis and Crossop)us fimbriaius Fitzinger, Sitzungsber.

kais. Akad. Wissensch. Wien, ]NIath.-Naturwiss. Clas>"e, lvii, pt. 1,

p. 610 (Synonyms of " Cvossopns daubcntonii nigripcs," wrongly

attributed to Giebel, Saugethiere, p. 899). Not Sorex jinibriatus

Wagler, 1832.
1868. Crossopiiis ciliatiis, griscogularis Fitzinger, Sitzungsber. kais. .\kad.

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

(Chartres, Eure-et-Loire, France).
1870. Sorer intermedins Cornalia, Catal. Descrit. IMamm. Ital., p. 27 (HiUs

of Brianza, Como, Italy). Part: tail only (see Sordelli, Atti

Soc. Ital. Sci. Nat. e del 3Ius. Civ. Stor. Nat., Milano, sxxviii,

p. 364, 1899).
1895. Neoniys fodiens Thomas, The Zoologist, 4th ser., ii, p. 100, March,

1908.
1899. Sorcx alpiniis var. longobardus Sordelli, Atti Soc. Ital. Sci. Nat. e

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

of intermedius).
1901. Ncomys fodiens minor Miller, Proc. Biol. Soc, Washington, xiv,

p. 45, April 25, 1901 (Moutrejeau, Haute-Garonne, France). Type

in U.S. National Museum.
1905. Crossopus ou Sorcx ignotiis Fatio, Arch. Sci. Phys. et Nat., Genfeve,

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

skull, not mandible. Type in Geneva Museum.
1905. Ncoviys fodiens naias Barrett-Hamilton, Ann. and Mag. Nat. Hist.,

7th ser., XV, p. 507, May, 1905 (Hatszeg, Hunyad, Hungary). Type

in British Museum.

NEOJIYS 71

1906. Ncomys fodiens nanus Lydekker, Zoological Record, xlii (1905),
Mamm., p. 34, August, 190C (Accidental renaming of naias).

1910. Neornys fodiens, N. fodiens naias and N. fodiens minor Trouessart,
Paune l\Iamm. d'Europe, pp. 56, 57.

Type localiti/. — Berlin, Germany.

Oeographical distribution. — Continental Europe from Norway
to Italy and the Pyrenees.

Diagnosis. — Underparts buify whitish, occasionally tinged with
a stronger shade of yellow or suffused with salmon-colour, and
sometimes though rarely washed with wood-brown, especially on
chest and along median line of belly. Condylobasal length of
skull usually more than 21 mm.

Measurements. — Average and extremes of ten specimens from
Brunswick, Germany : head and body, 86 • 6 (83-92) ; tail, 65 " 5
(58-77)] hind foot, IS '5 (17-19). Average of three specimens
from Montrejeau, Haute-Garonne, France : * head and body, 83
(82-85); tail, 52 (-50-60); hind foot, 17-1 (16-18). Average
and extremes of seven specimens from Luchon, Haute-Garonne,
France: head and body, 75*7 (72-80) ; tail, 53-1 (47-58) ; hind
foot, 16 '4 (16-17). Average and extremes of five specimens
from Porte, Pyrenees-Orientales, France : head and body, 93
(83-96) ; tail, 6''l (57-65) ; hind foot, 18-2(18-19). Average and
extremes of nine specimens from Meiringen, Bern, Switzerland :
head and body, 82-7 (80-86); tail, 66"- 2 (62-71); hind foot,
19 "3 (18—20). Average and extremes of ten specimens from
Hatszeg, Hunj^ad, Transylvania : f head and body, 77 • 7 (72-82) ;
tail, 60 (56-66) ; hind foot, 18-5 (18-19). For cranial measure-
ments see Table, p. 75.

Specimens examined. — One hundred and thirty-four, from the following
localities: —

Norway: Vefsen, Nordland, 1; Kvikne, Hedemarken, 1; Brekke-
bygden, Trondhjem, 1.

Sweden : Bothnia, 1 ; Upland, 1 ; Medstugan, Jemtland, 1 ; Upsala, 1
(U.S.N.M.) ; no exact locality, 1.

Belgium: Waremme, Liege, 3 (U.S.N.M.).

France : Guines, Pas-de-Calais, 2 ; Abbeville, Soiime, 2 (B.!M. and Paris ;
the latter type of remifer Geoffrey) ; near Paris, 1 (Paris; type of lineatus
Gooffroy) ; Dinan, C6tes-du-Nord, 1 ; Cadillac-sur-Garonne, Gironde, 2
(U.S.N.M.); Montrejeau, Haute-Garonne, 3 (U.S.N.M.); Porte, Pyrenees-
Orientales, 5 ; I'Hospitalet, Ariege, 2; Luchon, Haute-Garonne, 7 ; liareges,
Hautes-Pyrenees, 3 ; Barcelounette, Basses-Alpes, 1 ; Charaonix, Haute-
Savoie, 1 (U.S.N.il.) ; Cranves-Sales, Haute-Savoie, 1 : Scientrier, Haute-
Savoie, 1 (Mottaz) ; fitupes, Doubs, 6 (Mottaz) ; no exact locality, 1.

Germany : Brunswick, 10 (U.S.N.M.) ; Saxony, no exact locality, 1
(U.S.N.:\r.); Bahrenberg, Harz Mts., 1 (U.S.N.M.): Frankfort, Wies-
baden, 1 ; Niesky, Silesia, 1 : Strassburg, 1.

Austria-Hungary : Haida, Arva, Bohemia, 3 ; Hatszeg, Hunyad,
Transylvania, 11.

Switzerland : Geneva, 4 (Mottaz) ; Chesieres, Vaud, 4 (Mottaz) ; Les
Plans, Vaud, 3 (U.S.N.M.) ; Meiringen, Bern, 9 (U.S.N.:M.) ; GrindelwrJd,

* " Neomys fodiens minor ^' Miller.

t " Ncomys fodiens naias" Barrett-Hamilton.

72

INSECTIVORA

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

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

Remarks. — There is much individual variation in colour as
well as in external measurements and proportions,* though the
size of the skull and teeth, as may be seen from the Tables,
remains very constant. The degree of development of the fringes
on feet and keel on tail is also variable, the depth of the keel in
some individuals being about equal to diameter of tail, while in
others it is too slight to be measured. This is partly, if not
entirely, due to season, as the deepest keels are found in winter
specimens. With regard to the variations in external measure-
ments shown by the averages and extremes in a preceding
paragraph, it is probable that they are much exaggerated by
different methods in taking the measurements. This is well
illustrated by the two series of Pyrenean specimens measured
by A. Robert at an interval of six years. AVhile much
material has been examined it is not wholly satisfactory, con-
sisting chietly of small lots taken by many diflferent collectors.
Eventually it may be necessary to recognize certain Continental
forms such as minor, naias, and the Scandinavian linneana, as
distinct from true fodiens ; but for the present, in view of the
uncertainties concerning external measurements and the striking
similarity of skulls from the entire range of the animal, there
seems to be no other alternative than to regard all the Con-
tinental water-shrews with keeled tail as belonging to a single

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

way. Waldo (c & p).

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

2200 ft. (A'. F. Ticc- (p).

hurst.)

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

hjem (A'. F. Ticehurst.) (p).

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

berg.)

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

550 m. (G.Kolthoff.)

2 9. Gaines, Pas-de-Caiais, 0. Thomas (c \- p). 94. G. 6. 1-2.

France.

6, 2 ?. Porte, Pyr6n6es-Urien- 0. Thomas (p). 8. 9. 1. 39-41.

tales, 1600-1700 m.

France. (A. Robert.)

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

73

9.
6,2.39.

2 6.

2 9.

6 6,i9.
2.

4:9.

9.

3 6.

i.

(5,2 9.
1 al.

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

L'Hospitalet, Ariege.

Luclion.Haute-Garonue,
G00-900m. (A.Robert.)

Bareges, Hautes-Pyr6-
n6es, 1300-1500 m.

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

Cranves-Sales, Haute-
Savoie.

Abbeville, Somme.

Frankfort, Wiesbaden,
Germany.

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

Strassburg, Alsace.
(C. Mottaz.)

Haida, Bohemia,
Austria.

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

Hatszeg, Transylvania,
1500-2000 ft. "

Miirren, Bern, Switzer-
land.

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

Faido, Ticino.

Locarno, Ticino.

Switzerland.

Busalla, Liguria, Italy.

Vallombrosa, Florence.

0. Thomas (p).

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

G. S. MiUer (c).

0. Thomas (p).

A. Robert (p).

V. BaiUon (c).

Dr. Dieffenbach (p).

Lord Lilford (p).

O. Thomas (p).

Lord Lilford (p).

C' G. Danford (c).

C. G. Danford (c).

\V. Gurtner (p).

0. Thomas (p).

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

8. 9. 1. 42.

8. 8. 4. 139.
G. 4. 1. 12. 14-

18.
8. 8. 4. 13G-

137.
8. 8. 10. 13.

5. 4. 9. 2.

54. A.

47. 1. 8. 46.

99. 1. 9. 8.

8. 8. 10. 12.

11. 1. 1. 148-

149.
3. 2. 2. 1.

3. 11. 8. 2-11.
92. 10. 5. 1-2.

4. 4, 5. 48-51.

5. 8. 2. 20.
5. 8. 2. 8-10.
46. 6. 15. 61.
5. 8. 3. 16-18.
1. 8. 2. 1.

Neomys fodiens bicolor Shaw.

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

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

England).
1837. AmpMsorcx pennantiC^T&y , Proc. Zool. Soc, London, p. 125 (England).
1840. Crossopus soiverbyi iBonaparte, Iconogr. Faun. Ital., i, fasc. 29, in

text under C. fodiens.
1857. Crossopus fodietis Blasius, Saugethiere Deutschlands, p. 120 (part).
1905. Nemnys fodiens ciliatus Barrett-Hamilton, Ann. and Mag. Nat. ffist.,

7th ser., xv, p. 508, May, 1905.
1910. Neotnys fodiens ciliatus Trouessart, Faune Slamm. d'Europe, p. 56.

Ty2)e locality. — Oxford, England.

Geofjraplncal distribution. — Great Britain.

Diagnosis. — Similar to Neomijs fodiens fodiens, but underparts
usually washed with wood-brown. Condylobasal length of skull
usually less than 21 mm.

Measurements. — External measurements of two adult females
from Scotland (Grantown-on-Spey and Cortachy, Forfar) : head

74

INSECTIVOEA

and body, 83 and 71 ; tail, 57-5 and 53 '5 ; hind foot, IS and 17.
Adult male from Wellington, Somerset : head and body, 75 ;
tail, 48 ; hind foot, 17. Adult male from Yalding, Kent: head
and body, 82 : tail, 52 ; hind foot, 17. For cranial measurements
see Table, p. 77.

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

Scotland : Gordonstown, Elgin, 1 ; Dunphail, Elgin, 1 ; Grantown-on-
Spey, Elgin, 1 (Wilson) ; Cortachy, Elgin, 1 (Wilson) ; Cromlis, Stirling-
shire, 1 ; Knibtuck, Stirlingshire, 1 ; Aberdeen, 1 ; Penkill, Ayrshire, 1.

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

England : ilill-on-Tyne, Northumberland, 1 ; Leeds, Yorkshire, 1 ;
Wellington, Somerset, 1; Great Grimsby, Lincolnshire, 2; Shaftesbury,
Dorset, 1; Halesworth, Suffolk, 1; Camliridgeshire, 2; Thornhaugh,
Northampton, 1 ; Podingtou, Wellingborough, Northamptonshire, 1 ;
Henley-on-Thames, Oxfordshire, 2; Drinkstone Park, Bury St. Edmunds,
Suffolk, 1 ; Woolpit, Bury St. Edmunds, Suffolk, 3 ; Halesworth, Suffolk, 1 ;
Cambridgeshire, 2 ; Epping, Essex, 1 ; Banstead, Surrey, 3 ; Esher, Surrey,
1 ; Garratt Park Lake, Surrey, 1 ; Godalming, Surrey, 1 ; Tillingbourne
Stream, Surrey, 1 ; New Forest, Hampshire, 4 ; IBasingstoke, Hampshire, 2 ;
Hampshire, no exact locality, 2; Yalding, Kent, 1.

6. Dunphail, Elgin.

1 al. 9 Aberdeen, Scotland.

1. Penkill, Ayrshire.

5. Fishguard, Pembroke-

shire, Wales.

1. Wales.

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

1 al. Wellingborough, ISTorth-

amptonshire.
1. Halesworth, Suffolk.

9. Cambridgeshire.

1. Cambridgeshire.

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

2 al. Henley, Oxfordshire.

1 al. Bury St. Edmunds,

Suffolk.
Sal. Bury St. Edmunds,

Suffolk.
i al. Esher, Surrey.
9 al. Crodalming, Surrey.

S, 3 ?. New Forest, Hampshire.

2 ?. Basingstoke, Hampshire.

2 al. Hampshire.

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

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

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

Rev. W. R. Tate

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

J. Baker (p).

W. Royal Dawson

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

Duncan Parker (p).

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

(c & p).
Miller Collection.

Miller Collection.

E. Bartlett (p).

7. 7. 7. 2Si8.

10. 11. 28. 1.

79. 'J. 25. 8.

11. 1. 3. 376.

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

84. 4. 21. 1.

87. C. 7. 1.

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

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

83. 8. S. 1.

84. 5. 16. 1-3.

11. 1. 3. 378.
94. 8. 5. 2.

7. 7. 7. 2845-

2849.
7. 7. 7. 4470-

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

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INSECTIVORA

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78

IXSECTIYORA

NEOMYS MILLERI Mottaz.

1907. Ncomys milleri Mottaz, !Mtm. Soc. Zool. de France, xx, p. 22,

September 20, 1907.
1910. Neomys milleri Trouessart, Fauna Mamm. d'Europe, p. 58.

Tyjw localitij. — Chesieres, Alpes A^audoises, Switzerland.
Altitude, 1,230 m.

Geographical (listrihitlon. — Pyrenees and Aljjs ; Hungary.
Details of distribution not j^et known.

Diagnosis. — Slightly smaller than Neomys fodiens : tail with
keel absent, oi* represented by a slight lengthening of hairs on
under side of terminal third only ; fringes on sides of feet not
conspicuously developed; hind foot usually less than 17 mm.;
tail usually less than 55 mm. ; lachrymal foramen opening over
point of contact between m^ and vf ; anterior upper incisor
slendei".

External characters. — Similar to Neomys fodiens, except that
the feet are less fringed and the tail lacks the definite keel on
under side. When unworn the hairs on ventral surface of tail
are slightly longer than those on upper side, and on terminal
third this elongation is occasionally sufficient to produce a
rudimentary keel ; but there is never any trace of a well-
defined ridge extending to base of tail. Caudal annulations
slightly less distinct than in Neomi/s fodiens. Mamma^ : a 1-1,
i 4_4 = 10.

Colour. — The colour is similar to that of typical specimens of
Neomys fodiens fodiens. Among the skins examined none shows
any noticeable wash of lirown or suffusion of buff on underparts.

Skull and Teeth. — The skull closely resembles that of Neomys
fodiens, but may be distinguished by its smaller general size and
relatively lower, more slender rostrum ; lachrymal foramen over
point of contact of m^ and nr. Teeth smaller throughout, a
difference especially noticeable in the anterior upper incisor.
Upper unicuspids with crowns longer and narrower than in
Neomys fodiens, the long posterior portion especially noticeable
in lateral view by comparison with height of cusp ; cingula
not so well developed as in the related species, and seldom, if
ever, forming a postero-external cusp.

Measurements. — External measurements of type: head and
body, 7G ; tail, 59 ; hind foot, 16 ; hind foot, including claws,
17. Average and extremes of ten specimens from the tyjje
locality: head and body, 77-4 (71-87); tail, 53-7 (50-59);
hind foot, 15 "8 (15-16 '2). Average and extremes of six
.specimens from Untervatz, Grisons, Switzerland : head and body,
79-3 (75-82); tail, 46-8(45-51); hind foot, 14-16 (14-15-4).
Two adult males from Locarno, Ticino, Switzerland : head and
body, 86 ; tail, 50 and 53 ; hind foot, 16 and 17. Average and
extremes of three adult females from 1' Hospitalet, Ariege,

NEOMYS 79

iooc 10 t (10 4-16). Adult male from Barec^es Hanfp«
Pyrenees: head and body, 76; tail, 56; hind frtf 16-4 For'
cranical measurements see Table, p. 80.

Specimens c.ra»M»ie(Z.— Thirtv-thrpp frnm fv,^ j^^n •

Switzerland: Near Geneva i V ^ *^^ f°H?'^i?S ^"''^^''^^s :-
Chesieres, Alpes Vaudoises 10 m M T^T'Jf' y^^*^' ^ (U.S.N.M.) ;
(U.S.N.M.); Untervl^SSois Su's N M f f 'f ^ = Meiring^n, Bern/i
(B.M. and V.S.N.M.) ; Sorengo Ti ino fm S N An .\""^*^"' ^t- Gallen, 3

Italy: Porlezza, Como. 1 (Mottaz) ^^■'^•^•^^'^•) ' L°°amo, Ticino, 2.

Fr™ ■^"^S-"^''-'/,^''^"'''^'-'' Northern Hungary 1
HaSe^^Tren^efr'*^'''' ^"'^^' '^ ^ ^-^°-' Haufe-Garonne, 1 ; Bareges,

complete absence T t;^ trie keetl^ the'Tr'T, '^-^
elongation of the hairs in'^meTan.^giorwi ^^^'

ThTsir r r;r- ^^ ^^- p--^^- s Uiroftj
S^sf^faji^hr^^^^^

Sa'S^'a^ttMr^™ l^^^r^^^ a^l^t:;; of
Their bnC. 1 " ^ specialized than iV^eo,««s /bf?/e„«

of the c 1^;^^^^^^^^^^^^^^^ t-; thoTe

locaHties where ^^^i^:^^^^^' ^^^ '^^^^ ^^^^^her in

''• "^Fratt^'^*' ^"'°^' G-S-^IillerCc). 8.8.4.138,140.

?. Chesjeres Alpes Vau- C. Mottaz (p) r, q r a

doises,1200 m. Swit- ^^ °- ^- "• ^•

zerland.

^ (^- -^^ Zollikofer.)

'^Sn\^"''^°' ^"^*- 0- Thomas (c&P). 5.8.2.6-7.
1. Zubere,-., Hungary. Budapest Museum 94 3 1 ^G

(e).

Ha^eiy^ILSf Srerif ituu^d^Trrtwi'^'^^P'^^^*' ^^''^ ^^ ^^-^oCS.
were taken alternateh in the sameTr°urs'^t n^H^'^T *°^^ ^^'^

streams. In the original descrint^on nif^ ■ ^\^ """^^^ °^ mountain

observes (p. 23) that according to^i°"llr.!^ \^ov,eyer, Mr. Mottaz

is the more inclined trwander awav fiom f hi ' *^^- ^.'^"'^d-tailed animal
The habits of .V. «.w,„a? fsin SI CenTral ^n"'"'''^'^'" vicinity of water,
aquatic as those of N. fodiens ^^'^ ^^P^'"' *° ''^ »« «t"ctlv

so

INSECTIVOEA

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PACHYURA

81

NEOMYS ANOMALUS Cabrera.

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

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

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

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

Oeographical distribution. — Northern and central Spain.

Diagnosis. — Similar to Neomys milleri, but with longer tail
(usually over 55 mm.), and larger hind foot (16 "8 to 18 mm.).
MammiB : a 1 — 1, i 4 — 4 = 10.

Measurements. — Type specimen, male (from Cabrera) : head
and body, 73 ; tail, 60 ; hind foot, 17 "5. Average and extremes
of nine sj^ecimens from Silos, Burgos, Spain : head and body,
83-2 (76-88); tail, 60 (56-61); hind foot, 17-4 (17-18).
Adult male and adult female from Barracas, Castellon, Spain :
head and body, 99 and 100; tail, 53 and 52; hind foot, 17.
For cranial measurements see Table, p. 82.

Specimens examined. — Tweuty-two, from the following localities in
Spaia: Silos, Burgos, 9; La Granja, Segovia, 2; Barracas, Castellon, 2 ;
L^rida, 9.

Remarlcs. — The Spanish water-shrew is well differentiated
from Neomys milleri by its longer tail and larger hind foot. As
yet no member of the keel-tailed group has been found in the
Iberian Peninsula, though it is probable that N. fodiens occurs in
the region north of the Ebro, and perhaps also in the Asturias.

2 (5, 7 9. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 22-30.

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

6,9. Barracas, Castellon. O. Thomas (p). 8.2.9.41-42.
(N. Gonzalez.)

Genus PACHYURA de Selys-Longchamps.

1889. Pachyura de S^lys-Longchamps, Etudes de Micromamm., p. 32

(Sorex ctruscus Savi). Sub-genus of Crocidura.
1857. Pachyura Blasius, Saugethiere Deutschlands, p. 147 (Sub-genus of

Crocidura).
1897. Pkrodus Schulze, Mamm. Europ., in Helios, Abhandl. u. Vortraga

Gesammtb. Naturwiss., xiv, p. 90 (Crocidura suaveolens Blasius

= Sorex ctruscus Savi).

Type species. — Sorex etruscus Savi.

Geographical distribution. — ^Africa and warmer portions of
Asia ; in Europe confined to the Mediterranean region.

Diagnosis. — Like Crocidura (p. 86), but with upper uni-
cuspids 4-4, the dental formula : i '"*, c ^i, pm ^3^, m -^"^ = 30.

Remarlcs. — Although perhaps not forming a natural group,
the 30-toothed species of Crocidura may for convenience be

G

82

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PACHYURA

83

treated as ineinbers of a distinct genus. The only European
species is immediately recognizable by its excessively small size.

PACHYURA ETRUSCA Savi.

1822. Sorex ctrnscus Savi, Nuovo Giorn. de' Letterati, Pisa, i, p. 60 (Pisa,

Italy).
1835. [Sorex] pachyunis Kiister, Isis, p. 77 (Cagliari, Sardinia).
1841. Pachyura etrusca Bonaparte, Iconogr. Faun. ItaL, i, Indice disfcrib.
1857. Crocidura suaveolens Blasius, Siiugethiere Deutschlands, p. 147.
1910. Pachyura etrusca Trouessart, Faune Mamm. d'Europe, p. 43.

Tfipe locality. — Pisa, Italy.

Geographical distribution. — Italy and adjoining portions of the
Mediterranean region.* Limits of range not known.

Diagnosis. — Much smaller than Crocidura russula (head and
body, 3.5-40 mm. ; hind foot, 7 ' 6-8 mm.), but with tail (25-
30 mm.) relatively longer, its ratio to head and body about
70 ; skull excessively small, its cond3'lobasal length only about
12 '8 mm., the dorsal profile nearly straight from front of nasal
to back of parietals, usually a little concave in interorbital
region, the brain-case narrow but even more flattened than in
Crocidura leucodon and C. mimula ; teeth not conspicuously
difiei'ent from those of the smaller European species of Crocidura,
except for the presence of the fourth upper unicuspid, and their
much smaller size (maxillary tooth-row about 6 mm.).

External characters. — ^In general external form Pachi/ura
I'trusca does not differ notably from Crocidura rus.mla except in
the relatively longer tail. The weight of a full grown individual
is, however, probably not more than one-fourth or one-third that
of an adult C. russula, and the head and feet are equally small
in proportion, a character by which Pachyura etrusca mav be
easily distinguished from young individuals of the larger animal.
Mamma;, i 3 — 3 = 6.

Colour. — Upper parts a uniform slaty brown, perhaps best
described as drab-grey washed with light bister, the indi^■idual
hairs slate-grey at base. Underparts faintly contrasted, rather
pale drab-grey with silvery reflections in certain lights. Feet
dull light slaty grey not noticeably contrasted with back. Tail
like back above, usually becoming somewhat darker toward tip,
lighter and more like belly below, but with no evident contrast
between the colours of the two surfaces.

Skull. — Apart from the small size the skull differs from that
of its European allies in .several important characters. Most
noticeable among these is the extreme flatness of the dorsal
profile, which is essentialh' straight from nares Lo occipital,
usually a little concave in interorbital region, while in the
European foims of Crocidura it is usually a little convex
throughout, and never evidently concave in interorbital region.

* Spain. See footnote, p. 8G.— O. T.

84 INSECTIVORA

The l)rain-case is excessively flattened, relatively more so than in
Crocifhira miviula, so that there is less contrast between its depth
and that of rostrum. This flattening is not accompanied by any
increase in breadth of brain-case, the ratio of
^^^T^ which to condylobasal length is about 46,

J\^^;^a^ slightly less than in Crocidura mhimla. Ante-

^^^ orbital foramen relatively large, its position

^ « normal. Lachrymal foramen over posterior

f\ i\ ^^^^^ "^^ '"'■ P'^.late relatively short and wide.

/ s M^-\ Mesopterygoid space relatively longer than in

( I ) W%] ^^^® European forms of Crocidura, its anterior

\gi>/ Xjdy border on line with jjosterior edge of third

Fig. 16. niolar instead of distinctly behind it, its lateral

Nat. size. borders nearly f)arallel, though slightly con-

verging posteriorly : hamular excessively deli-
cate, bowed outward and upward, its length relatively greater
than in C. russula, and about equal to greatest breadth of fossa.
Mandible not peculiar except for its small size (length about
7 mm. or less), and very delicate structure.

Tei'th. — While in general, and aside from the presence of
the fourth unicuspid, the teeth do not differ materially from
those of the smaller European Crocidurse except in size, they
show several jjeculiarities in form. The anterior upper incisor
projects more strongly forward, and has the anterior cusp rather
shorter than in the species of Crocidura, while the posterior cup
is more distinctly separated from the cingulum, these two
characters together imparting to the tooth a form somewhat
suggesting that assumed in Sorex. First unicusjaid relatively
larger and higher than in Crocidura rus.mJa, its cingulum less
curved, and its width distinctly exceeding that of palate instead
of barely ecjualling it. Contrast in both height and crown area
of first and second unicuspids greater than in C. runi<n1a, the
area of second about one-third that of first instead of evidently
more than one-third. Third unicuspid slightly larger than second.
Fourth about half as large as third, crowded inward from the
tooth-row, but visible from the outside through space separating
third unicuspid from large premolar, the width of this space
rather more than half diameter of crown of fourth unicuspid.
Large premolar as in C. Jeucodon, except that cutting blade is
shorter and even higher, and posterior border of crown is less
concave. Upper molars with hypocones less distinct than in
C. ruamla, but otherwise not showing any tangible peculiarities.
Mandibular teeth essentially as in Crocidura russula and C. leu-
codon, but posterior section of third molar even more reduced.

Measurements. — Average and extremes of six specimens from
Florence, Italy: head and body, 38-5 (36-42); tail, 27-1
(26-28) ; hind foot, 7-7 (7 • 6-7 • 8). Adult male and female
from near Turin, Italy : head and body, 42 and 40 ; tail, 28 and
27 ; hind foot, 7 • 8 and 7 • 4. Adult male and female from near

PACHYDRA

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INSECTIVOEA

Genoa, Italj- : head and body, 41 and 42 : tail, 26 and 28 ; hind
foot, 7 • 6 and 8 • 0. Adult female from Hassari, Sardinia : head
and body, 41 ; tail, 29 ; hind foot, 7*8. For cranial measure-
ments see Table, p. 85.

Specimens examined. — Twenty-nine, from the following localities : —

Italy : Near Turin, 2 (Genoa) ; Perti, Finalborgo, 2 (Genoa) ; near
Genoa, 5 (U.S.N.M. and Genoa) ; Pisa, 5 (B.il. and Mottaz) ; Florence, G
(Li .S.N.M. and JMottaz) ; Tuscany, 2 ; Rome, 1 (Genoa) ; no exact locality, 2.

Sicily : No exact locality, 2.

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

Greece : Lamia, 1.

lal.

Pisa, Italy.

Marquis G. Doria (p).

G9, 3. 4. 1.

2al.

Tuscany.

(No hist

Dry).

lal.

Italy.

Zoological Society's
Collection.

55. 12. 26. 295.

Skeleton.

Italy.

Purchased.

58. 10. 21. 13.

6, ? al.

Sicily.

Purchased (Parzu-
daki).

62. 2. 26. 22-23

lal.

Sardinia.

Hon. N. G. Roths-
child (1').

11. 10. 1. 1.

1 al.*

Lamia, Greece.

E. MacDonell (p).

8. 7. 22. 1.

Genus CROCIDURA Wagler.

1832. Crocidiira Wagler, Isis, p. 275 (Soivx leucudun Hermann).

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

1869. Leucodan Fatio, Faune Vert. Suisse, i, p. 132 (substitute for

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

Gesammtb. Naturwiss., xiv, p. 90 (Sorcx Icucodon Hermann and

iS'. arnncus Schreber = .S'. russulus Hermann).

Ti/pe species. — Sorex leucodon Hermann.

GeograjjJiical disirihufion. — Africa and warmer portions of
Europe and Asia, including the Malay Archijjelago ; in Europe,
north to northern Holland and central (Germany, west to the
Atlantic coast and the Channel Islands. Ab.sent from Great
Britain and Ireland.

Characters. — Upper uuicusjjid teeth o-o (dental formula :
' ^ '» ^ il'' i"'* ^' ™ %Er — '^^ '■> posterior lobe of upper incisor
noticeably less than half as high as main cusp ; anterior lower
incisor without lobes on cutting edge ; third lower molar with hypo-
conid and entoconid coalesced, so that form of tooth is strikingly
different from that of other molars, its crown 4-cusped instead
of 5-cusped ; second lower unicuspid without I'udimentary second
cusj) and commissure ; teeth white throughout ; skull lieavier and
more robust than in Sore.v and Neomijs, with conspicuously deeper
rostrum and less contrast between width of brain-case and that
of anterior portion ; rudimentary zygomatic process of maxillary
obsolete; external form rather heavy; ear rising noticeably

* <^, 2 9. Marismas, Lower Guadal- A. Chapman (c & p). 11.12.19.1-3.
quivir, Spain.
Received December, 1911. — 0. T.

CROCIDURA

87

above, t'ur, the meatus closed by two valves as in Neomys ; tail
somewhat thickened, its surface covered with short hairs, among
M^iich are sprinkled numerous longer ones ; habits terrestrial.

Beiiutrkn. — Though not highly modified in external pecu-
liarities the genus Crocidura is the least primitive group of
shrews occurring in Europe. This is indicated by the reduced
number of upper unicuspids, but is more clearly shown by the
highly modified form of the third lower molar and the completely
unicuspid character of the lower premolar. Among the European
members of the family Soricidse the species of Crocidura may
at once be recognized by their moderate size, entirely white teeth,
large ears, and by the presence of long, loosely spreading hairs
scattered over the surface of the tail. Badly prepared specimens,
in which the true characters are obscured, may often be detected
among skins of Sorcx by the noticeable silvery reflections on
hairs of back. Eight species are now known to occur in western
Europe, while the number of forms thus far described from other
portions of the range of the genus is not far from one hundred.

KEY TO THE EUROPEAN FORMS OF CROCIDURA.

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

Hind foot about 14 mm. (Sicilv) C. caudata, p. 110.

Hind foot 11 to 12-5 mm.

Upper tooth-row about 8'6 mm. (Corsica) C. ci/rnensis, p. 111.

Upper tooth-row about 8 mm. (Balearic Islands)... C. balearica, p. 112.
Tail about half as long as head and body, the ratio
normally varying from 40 to 60.
Condylobasal length of skull 16 to 17 '6 mm.

Second upper unicuspid with crown area about

equal to that of third (Crete) C. canem, p. 109.

Second u^sper unicuspid with crown area distinctly

less than that of third C. mimiila, p. 94.

Colour tending to be pale and greyish (Basque

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

Colour tending to be dark and brownish.

Brain-case not decidedly flattened (Charente,

France) C. »«. kiUisma, p. 98,

Brain-case decidedly flattened (entire range

of species, western portion excepted)... C. 711. mimula, p. 95.
Condylobasal length of skull 17-6 to '20 '4 mm.
(usuallv more than 18 mm.).
Brain-case about half as high as wide or less.
Animal conspicuously bicolor ; condylobasal
length of skull usually more than 19 mm.

(Central Europe, south into Italy) C. h'ucodon, p. 88.

Animal obscurely bicolor ; condylobasal length

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

Brain-case more than half ashigb as wide C. ritsstila, p. 99.

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

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

Condylobasal length of skull usually 18 to 19 mm.

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

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

88

INSECTIVORA

CROCIDURA LEUCODON Hermann.

1780. Soirx leucodon Hermanu in Zimniermann, Geogr. Gesch., ii,

p. 382 Cvicinity of Strassburg, Germany).

1781. Sorcx leucodon Hermann in Schreber, Saugtbiere, pi. clix.

1782. Sor[ex] leucodon Hermann, Tabula Affinitatum Animalium, p. 79

(footnote).
1792. Sorcx alhipes Kerr, Anim. Kingd., p. 208 (based on Pennant's account

of S. Ieucodo7i Hermann).
1832. Croc[idura] leucodon Wagler, Isis, p. 275.

1857. Crocidura leucodon Blasius, Siiugetbiere Deutscblands, p. 140.
1869. Leucodon niicrourus Fatio, Faune Vert. Suisse, i, p. 173 (Substitute

for leucodon).
1897. [Crocidurci] leucodus Schulze, Mammalia Europsea, p. 18 (Substitute

for leucodon).
1910. Crocidura russula leucodon Trouessart, Faune Mamm. d'Europe, p. 44.

TyjK' locallfj/. — Vicinity of Strassburg, Germany.

Oeograpliical distribution. — Central Europe, from Belgium to
Hungary ; south into Italy. Not known from the Iberian
Peninsula.

Diagnosis. — Size large (among the European species) ; hind
foot, 1 1 to 13 mm. ; condylobasal length of skull, 18-4 to 20 mm. ;
upper tooth-row, 8 "8 to 9*0 mm. ; tail short, its actual length
28 to 38 mm., its ratio to head and body usually ranging from
38 to 43 ; skull with brain-case noticeably depressed, its height
less than half its width ; large upper premolar with antero-
external cusp rather large, its height usually greater than that
of first unicuspid ; colour of underparts whitish, strongly con-
trasted with rather slaty brown of back, the line of demarcation
along sides well defined.

External characters. — Fur shorter and more dense than in
Sorcc araneus, the length of hairs at middle of back about
3 • 5 mm. in summer, 5 mm. in winter, its texture not .specially
modified ; a few slightly elongated hairs (8 mm.) on flanks and
across rump. Eyes small and inconspicuous ; ears small but
rising conspicuously above fur, the two well developed valves a
conspicuous feature of the ear in freshly killed specimens. Feet
not peculiar in form ; less slender than in Sorcx araneus, finely
pubescent on dorsal surface and on lateral portions of posterior
half of sole ; fingers proportioned as in S. araneus, but graduation
less, especially in hind foot ; pads 6-6, those on palm distinct
though somewhat crowded, the surface of palms and soles finely
rugose between the pads. Tail less slender than in Sorcx and
Neomi/s, nearly terete or with under side somewhat flattened,
its length equal to about half that of head and body, its hairs of
two kinds : (a) finely appressed hairs less than 1 mm. in length,
nearly concealing the annulations and forming a very slight
pencil ; and (h) loosely spreading hairs about 5 mm. long, rather
thickly sprinkled among the others ; annulation ill-defined,
about 35 to the centimeter at middle. Mamma?, i 3 — 3 = 6.

CROCIDURA g9

Colour.-Uppev parts varying from a slaty drab to dull
usse , the hairs with a slight znetallic gloss and with silve.y
reflections which in certain lights produc'e an evident effect of
fane speck ing Underparts and inner surface of limbs buf^
white, dulled to a varying degree by the slaty under colour th^
chin and throat often suffused with cream-buff S'e of
demarcation sharply defined, extending just below ear and eye
and alongside of muzzle to middle of pad' The two extremes of
colour probably represent ill-defined dichromatic phases Tlost

?U duirXSh 'ofr^'' "", '? ^^^"^ degree ^termediate
^eet dull whitish often irregularly clouded with drab Tail
sharply bicolor, whitish below, concolor with back above tbe
longer hairs silvery grey. ^^^ auo>e, uie

Skull -The skull is slightly larger than that of Sorex araneus
and noticeably more heavily built, particularly that portion

feCd To ^it ;r""'' i^ ^' ;^'^^^^ '^ both VoadenL Ls
S S- ? } i ^T'^'-'^^ ^"*^"^^ ^^P^^« less conspicuously

fiom behind forward, whether skull is viewed from above or

legion than in the European species of Sorex and Neomvs its
main sutures closing early in life. Brain-case ^ '

slightly longer than broad, its posterior out- ^ --__

Ime rounded but broken by the slightly (L^=:^^X
projecting points of the condyles, its antero- ^^^T""^'''^'^
external border straight, sharply angled in
front ; sagittal crest low but evident in adult
skulls, meeting the complete lambdoid crests
posteriorly. Depth of brain-case at middle
slightly but constantly less than half greatest
breadth (see tig. 20, page 1 00). Dorsal profile
with a slight concavity at front of brain-case
and slight convexity over middle of rostrum
-Nares squarely truncate posteriorly, the ^"'- "•

lateral wall abruptly angled near middle. "^""Nri^r""'-
Anteorbital foramen relatively smaller than in
Sorex aranens, and region between it and edge of alveolus dis-
tinctly wider. Lachrymal foramen over meta^tyle of J PlaJe
forming outer wall of anteorbital canal nearly thi-ee times as wMe

ZoU^T ^'""''T- ^"^"^'•^^" '^'^^-^^'^ ^^gi"" over posterior
molars broader and more prominent than in th'e European si.ecies
of t^orex, but rudimentary zygomatic process of maxillary reduced

floor ofT" """;• /^^-P*-yg-d fo-a as in SoreLranT;
floor of bra n-case between tympanic bones narrower and with
distinct median ridge.

snecieT'if"^!?'"^'^'^'! noticeably heavier than in the European
species of .STor^'.r and Neamy,, the difierence in general asnect

.TttT' A^''" ^^'^^"^^^' brown colouring matter'n jli
ot cusps. Anterior upper incisor with main cusp lon< and
slender, abruptly hooked downward; basal lobe low and

90

INSECTIVOEA

Fig. is.

Crocidura leucodon. Anterior
teeth in profile, x 5.

triangular, its height less than half that of main cusp and only
a little more than half that of first unicusjiid tooth. General
form of first incisor not unlike that of
Neomys fodiens, but with even more
contrast between height of the two
cu.sps. Anterior lower incisor simple,
rather robust, the shaft slightly taj^er-
ing, the point slightly bent upwai'd, the
cutting edge without lobes. Upper
unicuspids robust, strongly contrasted
in size. First about double the height
of second and third, the points of which
are nearly in line with that of basal
lobe of anterior incisor. Its crown is
.somewhat longer than broad, with main
axis slightly oblifiue to that of tooth-row. In lateral view it is
nearly triangular, with anterior-border about half as long as upper
and lower. Cutting edge essentially
as in Neomys, but better develo2)ed
and reaching posterior border of
tooth somewhat outside of middle.
Crushing surface well developed,
occujaying about one-third area of
crown. Second and third unicus-
pids approximately equal, in both
height and crown area, to basal
lobe of anterior incisor, their crowns
distinctly narrower as well as much
shorter than that of first
unicuspid, their cutting
ridge and crushing sur-
face less well diflferen-
tiated than in first.
Second smaller than
third, its crown about
as broad as long. Third
sepai'ated from large pre-
molar b}^ a slight space,
its crown longer than
broad, its cusp about
efiual in height to para-
cone of large premolar.
Lower unicusjiids essen-
tially alike in form, the
second differing from that
of Sorex and Nrontys in
the complete absence of the rudimentary second cusp. First low,
narrow and long ; second high, its crown about as broad as long.
Upper cheek-teeth with crowns narrower than in Sor-ex and

FlC ID
CrocitJuj'd leucodon.

CROCIDURA 91

Neomys, the emargination of posterior border deeper, particularly
in jm^, and hypocones more distinct. Large upper premolar
with antero-external cusp (paracone) well developed, its height
usually about equal to that of third unicuspid or somewhat
more, the distance from its point to well-defined angle in
cingulum over anterior root of tooth distinctly more than half
length of anterior border of main cusp ; posterior cutting blade
high, the angle formed between its edge and the moderately
projecting point of main cusp ill-defined and obtuse. Third
upper molar actually as well as relatively smaller than in Sorcx
araneus, but containing the same elements. Lower cheek-teeth
essentially as in Sorex aranetis except for the pecuharities of m^
characteristic of the genus.

Measurements. — Average and extremes of nine specimens
from Etupes, Doubs, France: head and body, 78-3 (70-87)-
tail, 35-3 (32-39); hind foot, 12-3 (12-13). Average and
extremes of nine specimens from Untervatz, Orisons, Switzerland :
head and body, 81 (77-85); tail, 31-6 (29-34); hind foot,
12-5 (11-8-13). Average and extremes of four specimens from
Lugano, Ticino, Switzerland: head and body, 80 (77-81); tail,
32 • 2 (31-36) ; hind foot, 1 2 • 5 (1 1 • 8-13). Average and extremes
of three specimens from Florence, Italy (in alcohol, bodies
contracted): head and body, 63 '3 (63-64); tail, 34-3 (32-36) ;
hind foot, 12-3 (11-6-13). " For cranial measurements see Table,
p. 92.

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

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

France: Dinau, Brittany, 1; Paris, 1 (U.S.N.M.); Manouville,
Meurthe-et-Moselle, 1 ; Etupes, Doubs, 14 (B.M. and Mottaz).

Germany : Brunswick, 1 (U.S.N.M.) ; Nussberg, Brunswick, 1 ; Frank-
furt, Hessen-Nassau, 1 ; Taucha, Saxony, 1 (U.S.N.M.) ; Marxheim,
Bavaria, 4 ; Ummerstadt, Thiiringen, 1 ; no exact locality, 1. '

Austria-Hungary : Hatszeg, Hunyad, Transylvania, 1.

Switzerland : Geneva, 3 (Mottaz) ; Briinig, Bern, 1 (U.S N.M.) ; Mei-
rmgen, Bern, 4 (U.S.N.M.); Untervatz, Grisons, 15 (B.M. and U.S.N.M.);
Grisons, no exact localitv, 1 ; Lugano, Ticino, 4 (U.S.N M ) • Ziiber-
wangen, St. Gallen, 1. "

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

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

92

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■2 ?, 1. Esneux, Liege, Belgium Lord Lilford (p). 'Jo. 1. 1. 1-2.

[H. Griinvold.)
i. Dinan, Brittany, Prance. G. Barrett-Hamilton 11. 1. 2. 158.

i. Manonville,Meurthe-et- Lord Lilford (p). 8. 9. 8. 1.

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

(C. Mottaz.)
2 9. Marxheim, PSavaria, Lord Lilford (p). 8.9.8.2-8.

Germany.
1. Frankfort, Hesseu- Dr. Dieffenbacli (p). 47.1.8.50.

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

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

9. Hatszeg, Transylvania, G. G. Danford (c). 3. 2. 2. 10.

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

Switzerland. 10. 8. 16. 9-13.

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

(E. H. Zollikofer.)
9. Lugano, Ticino. ' O. Thomas (p). 10. 8. 16. 15.

(E. II. ZoUil-ofcr.)
?. Ziiberwangen,St.Gallen. 0. Thomas (p). 10. 8. IG. 14.

(E. H. Zollikofer.)
6. Rome. (C. Coli.) G. Barrett-Hamilton 11. 1 2. 103.

(P).

CROCIDURA MIMULA Miller.

(Synonymy under subspecies.)

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

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

Colour. — Ti^pper pai'ts varying from a dull russet tinged with
sepia to a dark hair-brown with a tinge of drab, the liaii-s with
metallic gloss and silver}' reflections ; underparts varying from
dull ochraceous-buff" to greyish cream-buff", never sharply defined
from colour of back and sides. Feet bufly whitish, usually with
some dark clouding. Tail greyish or brownish, very obscurely
bicolor.

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

CROCIDURA

95

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

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

Remarhs. — Ciocidnra mimiihi differs from the other continental
European members of the genus, in its small size, a character in
which it is approached by the small races of C. russulu, though
not sufficiently to cause any confusion. In addition to the
typical form two geographical races have been described, one from
south-western France, the other from northern Spain, the
status of neither of which is clearl}' understood.

Crocidura mimula mimula Miller.

1839. ?? [Crocidura aranea] var. minor de S61ys-Longchamps, Etudes de

Micromamm., p. 35 (Silesia).
1901. Crocidura mimula JMiller, Proc. Biol. Soc, Washington, xiv, p. 95,

June 27, 1901 (Ziiberwangeu, St. Gallen, Switzerland). Type in

U.S. National Museum.

1901. Crocidura antipse Matschie, Sitz.-Ber. Gesellsch. Naturforsch.

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

1902. [Crocidura"} minuta Lydekker, Zool. Record, xxxviii (1901), Mamm.,

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

Tijpe localitji. — Ziiberwangen, St. Gallen, Switzerland.

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

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

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

96

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98 INSECTIVORA

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

Si^ecimens exaynined. — Thirty-seven, from the following localities : —
Phance : Abbeville, Somme, 1 ; Agay, Var, 2.
Germany : Marxheim, Bavaria, 2.

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

RouMANiA : Gageni, Prahova, 1 ; Bustenari, Prahova, 1.
Bulgaria : Sofia, 1 (Andersen) ; Varna, 1 (Andersen).
Switzerland: Ziiberwangen, St. Gallon, 3 (U.S.N.M. and Mottaz) ;
Untervatz, Grisons, 2 ; Faido, Ticino, 1 ; Santa Margherita, Ticino, 3
(Mottaz) ; Davesco, Ticino, 1 (U.S.N.M.) ; Lugano, Ticino, 2 (B.M. and
Mottaz) ; Locarno, Ticino, 1.

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

Greece : Corfu, 2.

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

Prance.
2 (5. Agay, Var. G. S. Miller (c). 8. 8. 4. 162-168.

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

Germany. ( Wolter-

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

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

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

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

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

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

land.
6. Locarno, Ticino. O. Thomas (c & p). 5. 8. 2. 1.

6,9. Untervatz, Grisons. O. Thomas (p). 4.4.5.21-22.

{E. H. Zollikofcr.)
9. Lugano, Ticino. 0. Thomas (p). 4.4.5.57.

[E. H. Zollikofcr.)
9. Viareggio, Lucca, 5 m. 0. Thomas (c & p). 6. 8. 2. 21.

Italy.
6 Rome. (Coli.) G.Barrett-Hamilton 11.1.2.97.

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

(C. Mottaz.)

Crocidura mimula iculisma Mottaz.

1908. Crocidura nmmila icvMsma Mottaz, BuU. Soc. Zool. de Geneve, i,

p. 119, April 30, 1908. Type in Mottaz Collection.
1910. Crocidura mimula iculisma Trouessart, Paune Mamm. d'Europe, p. 47.

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

Geographical clistribution. — Known from the type locality
onl}'.

Diagnosis. — Size as in C. tiiimnla mimula or slightly smaller
(hind foot, 10; condyloba.sal length of skull, 16); brain-case
deep, nearly as in C. russida.

CROCIDURA 99

Measuremenh. — Type (from Mottaz) : head and body, 60 ■ 5 ;
tail, 38*5; hind foot, 10 •2. For cranial measurements see
Table, p. 97.

Specimens examined. — Three, all from the type locality (Mottaz).

BemarJcs. — While this race appears to be distinct from true
miniuJa, the material seen is insufficient to form the basis of any
final opinion as to its status.

Cbocidura mimula cantabra Cabrera.

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

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

Ti/jje localiiy. — Bastjue Provinces, Spain, exact locality not
known.

Geographical distribution. — Basque Provinces, Spain.

Diagnosis. — Colour paler and more grey than in the other
races ; size small.

Measurements. — Type (from Cabrera) : head and body, 55 ;
tail, 24 ; hind foot, 10 ; ear, 6*5 ; upper tooth-row, 7*2. (Cranial
dimensions not known.)

Bemarks. — I have not seen this animal, but from the original
description, as well as from information received from Mr. Cabrera,
it appears to be paler than the typical form, to which it bears
much the same relation as C. russula pulehra to true russula.

CROCIDURA RUSSULA Hermann.

(Synonymy under subspecies.)

Geographical (lislrihution. — Central and southern Europe,
from the Mediterranean coast to Holland and central Germany.
Not found in the British Islands.

Diagnosis — Size rather large (among the European forms) :
hind foot, 11 to 14 mm. ; condylobasal length of skull, 18 to
20 '4 mm.; upper tooth-row, 8-2 to 9 mm.; tail rather short,
its actual length 33 to 45 mm., its ratio to head and body
varying from 45 to 55 ; skull with brain-case not noticeably
depiessed, its height always at least half greatest width and
usually more ; large upper premolar with autero-external cusp
small, its height usualh' less than that of third unicuspid ;
colour of underparts not strongl}' contrasted with that of back,
the line of demarcation along side vaguely defined.

E.vtcrnal chararteis. — In external characters, aside from the
relatively longer tail, Crocidura russula agrees with C. Icucodon.
Depth of fur at middle of back about 5 mm. in summer, 8 mm.
in winter.

H 2

100 INSECTIVORA

Colour. — Upper parts varying from a dark hair-brown,
tinged with bister to a light drab with or without a shade of
wood-brown, the darker colour more frequent in winter pelage,
the light apparently pecuhar to summer. The pelage has the
usual metallic gloss, and the individual hairs show strong silvery
reflections in certain lights, particularly in the long full winter
coat. Underparts usually a dull buffy grey or ecru-drab, but
sometimes almost whitish, rarely tinged with a bright yellowish
brown,* never strongly constrasted with back, the line of
demarcation along sides always vague. Feet dull buffy grey or
light drab Tail obscurely bicolor, like back above, like belly
below.

Shtll and teeth. — The skull resembles that of Crocidura
leucodon, except that the brain-case is noticeably less flattened,
its depth at middle always exceeding one half greatest width, a
character readily appreciable to the eye when skulls of the two
animals are viewed from beliind. Teeth essentially as in the
related species, but large upper premolar with antero-external

Fig. 20. Fig. 21.

Posterior view of skull of Crocidura rus$itla. Anterior

Crocidura leucodon (upper teeth in profile. X ."i.

figure), and C. russu!a (lower
figure). X li.

cusp (paracone) low, its height often much less than that of
third unicuspid, the distance from its point to ill-detined angle
in cingulum over anterior root of tooth about half length of
anterior border of main cusp ; posterior cutting blade not so
high as in Crocidura leucodon, the angle formed between its
edge and conspicuously projecting point of main cusp well
defined and less obtuse than in the related species.

Measurements. — In the diflerent races the head and body
ranges from 04 to 95 mm., tail from 33 to 46 mm., hind foot
from 10 '8 to 14 mm., condylobasal length of skull from
18 to 20*4 mm. The unusual apparent variability in length
of head and body is probably in great part due to diflerences in
method of taking the measurement and to differences in the
condition of the specimens measured.

* Such specimens evidently formed the basis of Savi's Sorex tltoracicns
and Dehne's S. chrysothorax.

CEOCIDCRA 101

Crocidura russula russula Hermann.

1777. Sorex araneus Schreber, Siiugthiere, iii, p. 573 (not of Linnaeus, 1758).
1780. Sorex russulics Hermann in Zimmermann, Geogr. Gesch., ii, p. 382

(vicinity of Strassburg, Germany).
1780. ?? Sorex constrictns Hermann in Zimmerinann, Geogr. Gesch., ir,

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

1792. ■?•? Sorex unicolor Kerr, Anim. Kingd., p. 208 (Strassburg, Germany.

Based on Pennant's account of Sorex constrictus).

1793. Sorex musaraneus Cuvier, Tabl. Til6m. de I'Hist. Nat. des Anim.,

p. 109 (France).

1800. ?? Sorex Icncurus Shaw, Gen. Zool., I, pt. 2, p. 538 (Strassburg,

Germany. Based on Schreber, pi. clixc, S. constrictns Hermann).

1801. S[prex^ alraneiis'] cincreus Bechstein, Gemeinn. Naturgesch. Deutsch-

lands, I, 2nd ed., p. 867, misprinted 863 (Thiiringen, Germany).
ISOl. S[orcx\ a[raneus'] candidus Bechstein, Gemeinn. Naturgesch. Deutsch-

lands, I, 2nd ed., p. 867, misprinted 863 (Thiiringen, Germany).
1832. Sorex fimbriat us Wagler, Isis, p. 54 (Bavaria, Germany).
1832. Croc[;idu7-d] moschafa Wagler, Isis, p. 275 (Substitute for Sorex

fimbriatus).
1832. C\^rocidiira] major Wagler, Isis, p. 1218 (Bavaria, Germany).
1832. Crocidura rufa Wagler, Isis, p. 1218 (banks of the Rhine, Germany).
1832. Crocidura poliogaster Wagler, Isis, p. 1218 (banks of the Bhine,

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

(near Pisa, Italy).
1839. ? Sorex inodorus de S61ys-Longchamps, Etudes de Micromamm.,

p. 34 (Savi cited as authority, but name apparently published here

for the first time as synonym of aranea (= russula)).
1839. ? [Crocidura ara7iea] var. ininor de Selys-Longchamps, Etudes de

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

Micromamm., p. 36. (No locality given.)
1839. ? Crocidura hydruntina Costa, Fauna del Ragno di Napoli, Mamm.,

p. 6 (Otrauto, Calabria, Italy).
1855. Sorex chnjsothorax Dehne, Allg. deutsche Xaturhist. Zeitung, Neue

Folge, I, p. 241 (Wilsdurf, near Dresden, Germany).
1857. Crocidura araneus Blasius, Saugethiere, Deutschlands, p. 144.
1895. Crocidura russula Thomas, The Zoologist, 3rd ser., xix, p. 63,

February, 1895.
1910. Crocidura russula Trouessart, Panne Mamm. d'Europe, p. 43.

Tyjje locality. — Vicinity of Strassburg, Germany.

Geographical distribution. — Central Europe, from Holland and
central Germany to the valley of the Garonne and the coast of
south-eastern France (Var) ; Italy ; Sardinia 1 * ; Guernsey and
Alderney, Channel Islands.

Diagruh<<is. — Size rather large (hind foot, 11 "7 to 14, condy-
lobasal lengtli of skull, 19 to 20-4), and colour usually dark,
seldom, if ever, becoming a light drab except in rather worn
summer pelage.

Measurements. — Average and extremes of four specimens from
Oosterbeek, Gueldei'land, Holland : head and body, 78 (76-81) ;
tail, 41 (37-45); hind foot, 12-8 (12 -2-13 -9). "Average and

* I have seen five Sardinian specimens in the Genoa Museum
resembling the typical form of Crocidura russula and differing widely
from the Corsicau C. cyrnensis (see pp. 111-112).

102

INSECTIVORA

extremes of nine specimens from Esneux, Liege, Belgium : head
and body, 77 (72-85); tail, 35-6 (33-38); hind foot, 12-4
(11' 7-13). Average of ten specimens fi-om Pas-de-Calais, France :
head and body, 74-5 (71-80) ; tail, 40-9 (38-46) ; hind foot, 13-1
(12 •5-13' 5). Average and extremes of six specimens from St.
Cergues, Vaud, Switzerland : head and body, 87 (83-95) : tail,
38-1 (35-41); hind foot, 12-8 (12-4-13-2). For cranial
measurements see Table, p. 104.

Spemyiens examined. — One hundred and two, from the following
localities : —

Holland : Oosterbeek, Guelderland, 4.

Belgium : Esneux, Liege, 9 ; Waremme, Liege, 8 (U.S.N.IM.) ; no exact
locality, 1.

France : Boulogue-sur-Mer, Pas-de-Calais, 8 ; Guines, Pas-de-Calais, 2 ;
Abbeville, Somme, 2 (B.M. and Mottaz) ; Cxuernsey, Channel Islands, 4;
Alderney, Channel Islands, 1 ; St. Briac, Brittany, 1 ; Lignieres,
Charente, 1 (Mottaz) ; Nancy, iIeurthe-et-]Moselle, 1 (Merriam) ; Etupes,
Doubs, 9 (Mottaz) ; Montauban, Haute-Savoie, 4 ; Valescure, Var, 2 ; Ax-
les-Thermes, Ariege, 1 ; Luchon, Haute-Garonne, 2.

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

Switzerland : Geneva, 10 (LT.S.N.INI. and jNIottaz) ; St. Cergnies, Vaud, 9
(U.S.N.M. and Mottaz); Cxrosjoan, Vaud, 1 (^Mottaz) ; Chesieres, Vaud, 1
(Mottaz) ; Lucerne, 1 ; Vitznau, Lake of Lucerne, 3 ; Thurgau, Roggwil, 1 ;
St. Gallon, 3 (U.S.N.M.) ; Ziiberwangen, St. Gallen, 5 (B.M. and U.S.N.M.) ;
Degersheim, St. Gallen, 5 (B.M. and U.S.N.M.) ; Engelberg, Unterwalden, 1.

Italy: Ceresole d'Alba, Turin, 2 (Turin).
3 <?, 1 ?. Oosterbeek, Guelderland, O. Thomas (c & p). 98. 2. 1. 9-12.
50 m. Holland.
6, ?. Oosterbeek, CTuelderland. Miller Collection. 7. 7. 7. 3851-

(0. Thomas.)
4 <5, 49,1. Esneux, Liege, Belgium.

Belgium. (H. Grunvohl.)
2(5, ?,1. Guernsey, Channel Is-
lands. (R.H. Bunting.)

Alderney.

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

Guines, Pas-de-Calais.

Abbeville, Somme.

St. Briac, Brittany.

Montauban, Haute-
Savoie.

IMontauban, Haute-
Savoie, 900 m.

(.4. Robert.)

Valescure, Var.

Ax-les-Thermes, Ariege.

Luchon, Haute-Garonne,
600 m. [A. Robert.)

Ingelheim, Rheinhessen,
Germany.

Strassburg, Alsace.

(C. Mottaz.)

Vitznau, Lake of
Lucerne, 500 m.
Switzerland.

Engelberg, 3300 ft.
Switzerland.

9.
5(5,3 9.

2 6.
1.
1.

2rf.

<5, 9.

9.

2 9.

3 9.

1 al.

Miller Collection.

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

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

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

0. Thomas (p).

G. S. Miller (c).
V. Builles (c & r).
O. Thomas (r).

C. Hilgert (c).

0. Thomas (p).

0. Thomas (c & p).

7. 7. 7.

3852.

95. 1. 1.

7. 1. 1. 30.

8. 9. 2. 18-21.

-11.

9. 3. 28.
98. 1. 9.

1.
4-11.

94. 6. 6. 8-9.
56. A.

94. 10. 3. 1.
97. 1. 9. 2-3.

6. 4. 2. 2-3.

8.8. 4.164-165.
8. 3. 27. 1.
6. 4. 1. 19-20.

8. 11. 2. 9-10.

8. 8. 10. 43.

5. 8. 3. 9-11.

Dr. J. Anderson (p). 99. 7. 17. ] .

CROCIDDRA 103

Crocidura russula pulchra Cabrera.

1907. Crocidura russula ptilchra Cabrera, Ann. and Mag. Nat. Hist.,
7th ser., xx, p. 213, September 1, 1907. Type in Cabrera col-
lection.

1907. Crocidura russula pulchra Cabrera, Bol. Real Soc. Espafi. Hist.
Nat., Madrid, vii, p. 223, October, 1907. (For date see Cabrera,
Ann. and Mag. Nat. Hist., 8th ser., i, p. 189, February, 1908.)

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

Type locality. — Valencia, Spain.

Geographical distribution. — Central and southern Spain ; low-
lands of France south of the Gironde.

Diagnosis. — Smaller than Crocidura russula russula (hind foot,
10" 8 to 13, condylobasal length of skull, 18 to 19 '4), and jjaler
in colour, the back a light drab brown tinged with sepia or dull
russet.

Measurements. — External measurements of type, male (from
Cabrera): head and body, 71; tail, 41*5; hind foot, 12.
Average and extremes of seven specimens from 'Silos, Burgos,
Spain: head and body, 72-7 (67-78); tail, 36 (34-37); hind
foot, 11 "9 (10 •8-12 -4). Average and extremes of .seven
specimens from Ciranada, Spain : head and body, 69 * 6 (68—74) ;
tail, 38-2 (36-40); hind foot, 12-2 (12-12-8). Average and
extremes of ten specimens from Cadillac-sur-Garonne, Gironde,
France: head and body, 75-2 (70-81); tail, 37 (33-40); hind
foot, 11*6 (11-12). For cranial measurements see Table, p. 106.

Specimens examined. — Sixty-five, from the following localities ; —

Feance : Cadillac-sur-Garonne, Gironde, 19 (U.S.N.M.) ; Moutr6jeau,
Haute-Garonne, 3 (U.S.N.M.) ; St. Genies, near Nlmes, Gard, 2.*

Spain : Silos, Burgos, 18 ; Dehesa de Valencia, Valencia, 1 ; Alcoy,
Alicante, 8 ; Elche, Alicante, 2 ; Venta del Baul, Granada, 2 ; Granada, 8 ;
Barracas, Castellon, 1.

Portugal: Sierra de Gerez, 1 (in alcohol; perhaps referable to cintrai).

9. St. Genies, Gard, 102 m. 0. Thomas (p). 8. 8. 10. 42.

France. (C. Mottaz.)

4 6. Silos, Burgos, 980 m. G. S. MiUer (c). 8. 8. 4. 38-41.

Spain.

9. Dehesa de Valencia. O. Thomas (p). 8. 2. 9. 46.

(N. Gonzalez.)

2 ,i, 1 9. Alcoy, Alicante. 0. Thomas (p). 8. 2. 9. 43-45.

(N. Gonzalez.)

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

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

<5al. Sierra deGerez,Portugal. Dr. H. Gadow 87.3.28.1.

(c & p).

* Intermediate between pulchra and true russtila.

104

INSF.CTIVOKA

5

J

O

Teeth slightly worn.

,, not worn.

,, slightly worn.

,, not worn.

,, slightly worn.

,, not worn.

,, slightly worn.

„ moderately worn.
,, slightly worn.

,, much worn.

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Number.

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98. 2. 1. 12

95. 1. 1. 3

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C. russula russ

Holland : Oosterbeek

Belgium : Esneux .
Warcmme

France : near Boulogi
Calais .

CROCIDURA

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INSECTIVOEA

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8. 8. 4. 39
8. 8. 4. 40
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8.8.4.42

172114
8. 7. 7. 44

1

C. russula russula {continued).

Switzerland: Ziiberwangen, St. \
Gallen . . ./
Degersheim

Lugano, Ticino .

Italy : Ceresole d'Alba .

Pisa

C. russula pulchra.

Spain : Silos, Burgos

CUOCIDURA

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108 INSECTIVORA

Crocidura russula ciNTRyE Miller.

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

XX, p. 390, November, 1907. Type in British Museum.
1910. Crocidura russula cintrx Trouessart, Faune Mamm. d'Europe, p. 45.

Type locality. — Cintra, near Lisbon, Portugal.

Geogrnpliical distrihution. — At present known onlj' from the
type locality.

Diagnosis. — Size as in C. russula pulchra (hind foot, 11 '4 to
12 "7, condylobasal length of skull, 18 to 19-2), but colour fully
as dark as in true russula, the back between the mars-brown and
russet of Ptidgway, the hairs with a peculiar, strong, coppery
lustre rarely indicated in the typical race.

Measurements. — External measurements of type : head and
body, 64; tail, 3.3 ; hind foot, 11*4. Average and extremes of
ten specimens from the type locality : head and body, 67 • 6
(64-72); tail, 37-7 (33-42); hind foot, 11 "9 (11 •4-12-7). For
cranial measurements see Table, p. 107.

Specimens examined. — Eleven, all from the type locality.

Remarks. — In its small size the Cintra shrew agrees with the
Spanish race, but the colour is conspicuously darker. Taken as a
whole the series, in winter pelage, is about as dark as in French
and Belgian russula ; but the noticeable coppery lustre is highly
characteristic of the Portuguese form.

6 6, 5 9. Cintra, Estremadura, 0. Thomas (c & p). 98. 2. 2. 10-20.
300-350 m. Portugal. (T7/|;eof subspecies 98. 2. 2. 11.)

CROCIDURA SICULA Miller.

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

1901. Crocidura sicula Miller, Proc. Biol. Soc, Washington, xiv, p. 41,
April 25, 1901 (Palermo, Sicily). Type in U.S. National Museum.

1910. Crocidura sicula Trouessart, Paune Mamm. d'Europe, p. 47.

Type locality. — Palermo, Sicily.

Geographical distribution. — Sicily.

Diagnosis. — Size and colour essentially as in Crocidura russula
pulchra (hind foot, 12 to 13 mm., condylobasal length of skull,
17 'G to 19 mm.); brain-case nearly as much flattened as in C.
leucodon, its depth usually a little less than half greatest width ;
crown ai'ea of molars slightly reduced.

Colour. — Upper parts a light bluish drab tinged with sepia ;
underparts faintly constrasted pale smoke-grey, with or without
a bufly cast. Feet dull whitish grey. Tail obscurely bicolor,
brownish above, whitish grey below.

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

CROCIDCKA 109

the skull resembles that of the small races of Crocidura russula.
Teeth as in C. russula, but upper molars with crown area
somewhat reduced, a character readily appreciable on comparison,
and large upper premolar with autero-external cusp slightly
enlarged, its form and relative size approaching the conditions
found in C. leucodon.

Measurements. — External measurements of type (male) and
a second specimen from the type locality : head and body, 68
and 75; tail, 32 and 35; hind foot, 12 and 12. Average and
extremes of six specimens from San Giuglielmo, Castelbuono,
Sicily : head and body, 76-3 (72-80) ; tail, 35-3 (32-41) ; hind
foot, 12'8 (12-13). For cranial measurements see Table, p. 113.

Specimens examined. — Fourteen, all from Sicily. Exact localities :
Palermo, 5 (B.M. and U.S.N.M.) ; Marsala, 2; San Giuglielmo, Castel-
buono, 6 ; Ficuzza, 1.

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

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

3 cJ, 1 ?. San Giuglielmo, Castel- 0. Thomas (p). 8. 9. 1. 6-9.
buono. (^4. Bobcrt.)

CROCIDURA CANE.ffi: Miller.

1909. Crocidura canese Miller, Ann. and Mag. Nat. Hist., 8th ser., iii,

p. 418, May, 1909. Type in British Museum.

1910. Crocidura canex Trouessart, Fauna Mamm. d'Europe, p. 48.

Type locality. — Crete.

Geographical distribution. — Island of Crete.

Diagnosis. — Size and general appearance as in the smaller
forms of Crocidura russula, and skull with similarly deep cranium ;
but second upper premolar as large as third, and entire anterior
portion of upj^er tooth-row unusually long relatively to cheek-
teeth.

Colour. — The colour does not difter appreciably from that of
dark individuals of C. 7-ttssula.

Skull. — The skull is essentially .similar to that of the smaller
forms of Crocidura russula. Brain-case slightly more than half as
high as wide. Anterior portion of palate between unicuspids
and anterior incisors more nearly parallel-sided, and more
elongate than in any of the related species.

Tectli. — The teeth differ from those of all the other known
European members of the genus in the approximately equal size
of the two small upper unicuspids and in the longer, relatively
narrower crown of the first imicuspid. In the related species the
first unicuspid is so wide posteriorly that it makes an abrupt and
noticeable break in the outline of outer side of tooth-i'ow. In
C. caneie this tooth, though larger than usual, is not sufficiently
wide to project beyond the general line of the outer margins of
the unicuspid teeth. Second unicuspid fully as large as third,
which is of normal size. The unicuspid row is thus distinctly

110 INSECTIVOKA

increased in length, so that the distance from front of large
premolar to front of incisor equals that from front of large
premolar to mesostyle of second molar, while in the related
species^it equals that from front of large premolar to metastyle
of first molar. Large premolars and molars, particularly those
of mandible, more robust than usual though not peculiar in form.
Measurements. — External measurements of type (male) : head
and body, 65 ; tail, 42 ; hind foot, 11 '8. External measurements
of adult male from Canea : head and body, 71; tail, 47;
hind foot, 12" 6 ; ear, 9*5. For cranial measurements see Table,
p. 113.

Specimens examined. — Two, both from Crete.

6 al. Crete. Purchased (LinnEea, 84. 3. 14. 2.

Frankfort).

[Type of species.)
6. Canea, Crete. A. Trevor Battye (p). 8. 10. 24. 1.

(C. H. B. Grant.)

CROCIDURA CAUDATA Miller.

1901. Civcidura caxidata Miller, Proc. Biol. See. Washington, xiv, p. 42,

April 25, 1901. Type in U.S. National Museum.
1910. Crocidura caudata Trouessart, Faune Mamm. d'Europe, p. 49.

Ti/pe locality. — Palermo, Sicily.

Geographical flistrihution. — Sicily.

Diagnosis. — Size about as in large specimens of C. 7u.9si(ln
(hind foot, 14 mm.). Tail very long, its ratio todaead and bodj-
about 80, and so thickened that its diameter at middle is 3 mm.
(in other European sjjecies the diameter of tail scarely if at all
exceeds 2 mm.).

External characters. — Except for the unusual length of the
tail Crocidura caudata does not difter in external characters from
C. russula. The tail is so long that when laid forward over back
it extends to between ears. It is distinctly 4-sided, broader
below than above, its greatest diameter at middle 3 mm.*

Colour. — After six months' immersion in alcohol the colour of
the type specimen was essentially as in Crocidura sictda. After
eight years more in the same fluid the back appears to have
assumed a somewhat more brownish cast.

Sliull and teeth. — The only known skull is so injured that the
details of its form cannot be seen. -The rostral portion does not
difter appreciably from that of C. russtda. Teeth essentially as
in C. russula, but lirst upper unicuspid larger, third unicuspid
more crowded against large premolar, and cutting edge of large
pi-emolar higher, its antero-external cusp, however, of the same
form as in C. russula.

* In the type the tail is flattened laterally for about 13 mm. from tip,
evidently as the result of an accident.

CROCIDDRA 111

Measurements. — External measurements of type : head and
body, 63; tail, 52; hind foot, 14. For cranial measurements
see Table, y>. 113.

Specimen examined. — -The type.

Hemarhs. — The tail is actually as well as relatively longer in
this species than in any other European member of the genus.

CROCIDURA CYRNENSIS Miller.

1907. Crocidura cyrnensis Miller, Ann. and Mag. Nat. Hist., 7th ser., sx,

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

Type locality. — Bastia, Corsica.

Geographical distribution. — Corsica.

Diagnosis. — Smaller than Crocidura caudata (hind foot, 12 to
12- 4 mm.) but with tail relatively almost as long, its ratio to
head and body about 70.

External characters. — Similar to C. caudata except for the
smaller size ; tail apparently less thickened than in the Sicilian
animal, its diameter at middle only about 2 mm.

Colour. — Back and sides drab washed with a brown inter-
mediate between wood-brown and raw-umber, this especially
noticeable on posterior half of back, but scarcely extending to
sides, which are a nearly clear drab ; underparts a light buffy
drab-grey, inconspicuously contrasted with sides ; tail dull dark
drab, essentially unicolor ; feet (both fore and hind) like tail on
outer half, rather sharply contrasted pale buffy grey on inner
half.

Skull and teeth. — AVhile its general size and form are essentially
as in Crocidura russula, the skull of the Corsican shrew is
distinguishable by its broader, more deepened rostrum. In the
type the mandible is peculiar in the unusual depth of ramus,
though in a second specimen this character is less marked.
Teeth essentially as in C. russula.

Measurements. — External measurements of type (adult male) :
head and body, 67 ; tail, 48 ; hind foot, 12-4. Very old female
from the type locality : head and body, 62 ; tail, 46 ; hind foot,
12*4 ; ear, 8 '2. External measurements of well made skin from
La Foce de Vizzavona : head and body, 72 ; tail, 51 ; hind foot,
12. For cranial measurements see Table, p. 113.

Specimens examined. — Three, all from Corsica.

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

112 INSECTIVORA

Ovile Seardu and two from Zinnigas) in the Genoa museum
appear to be strictly of the russula type, though without com-
parison of the skulls it is impossible to say whether they are most
nearly related to true russula or to sicula. Their average and
extreme measurements (from spirit specimens) are as follows :
head and body, 65-2 (60-69); tail, 36-8 (33-6-39): hind
foot, 12 (11 -8-12 -2).

1 La Foce de Vizzavona, Col. J. W. Yerbury (c & p). 93. 9. 15. 3.
Corsica.
6 al. Bastia, Corsica. Mrs. Southwell (c & p). 6. 3. 14. 1.

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

CROCIDURA BALEARICA Miller.

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

1907. Crocidura halearica Miller, Ann. and Mag. Nat. Hist., 7th ser., xs,

p. 391, November, 1907. Type in British Museum.
1910. Crocidura halearica Trouessart, Faune Mamm. d'Europe, p. 49.

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

Geographical distribution. — Balearic Islands, Spain.

Diagnosis. — In general similar to Crocidura cyrnensis, but
tooth-row distinctly shorter than in the Corsican form.

Colour. — The three skins are more noticeably brownish above
than in the one skin of ci/rnensis at present known, though the
actual elements of the colour are the same. Feet with the same
colour pattern.

Shdl and teeth. — In cranial and dental characters the Balearic
.shrew agrees with Crocidura cyrnensis, except that the mandible
is less robust (essentially as in the small races of C. russula) and
the tooth-row is distinctly shorter. Brain-case somewhat more
flattened than in the small races of C. russula.

Measurements. — External measurements of type (female) :
head and body, 62 ; tail, 45 ; hind foot, 12-5. External measure-
ments of two other specimens from the type locality (male and
female) : head and body, 71 and 72 ; tail, — and 45 ; hind foot,
12 "5 and 12. For cranial measurements see Table, p. 113.

Sjxcimcns examined. — Three, all from the type locality.

c?, 2 9 San Cristobal, Minorca; 0. Thomas and R.I. Pocock 0. 7. 1. 40-42.
Balearic Islands. (c & p).

(0. 7. 1. 42. Tij2)e of species.)

CROCIDURA

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114 insectivora

Family ERINACEID^.

1821. Erinaceidx Gray, London ^Med. Eepos., xv, p. 300, April 1, 1821.

Geogrcqjhical disiribntion. — Tropical and temperate Africa,
Europe and continental Asia ; in Europe west to Ireland, north
to central Sweden and south-eastern Norway.

Characters. — Skull deep and heavy, not specially tapering
anteriorly, most of the sutures persistent ; zygomatic arch com-
plete and heavy ; tioor of brain-case completely ossified ; tympanic
bone annular, not attached to skull ; auditory process of basi-
sphenoid large, sometimes forming an evident half-bulla ; glenoid
surface directed downward (normal) ; a large external pterj'goid
plate ; teeth anterior to molars neither well differentiated by
form into incisors, canines and premolars nor strictly " unicuspid,"
the anterior upper incisor higher than the others but not specially
modified in form ; anterior lower incisor short, oblique ; crowns
of upper molars rather high, sub-quadrate in outline (except the
reduced third), the cusjas sub-equal, subterete, near margin of
crown, the styles and commissures reduced or absent, never
forming an important functional part of the tooth ; form short
and heavy ; eyes and ears well developed ; snout pointed, some-
what produced ; back normally covered with short, stifl" spines.

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

Genus ERINACEUS Linmmis.

1758. Erinaceus Linnreus, Syst. Nat., i, lOth ed., p. 52 {E. curopxus).

1857. Erinaceus Blasius, Saugethiere Deutschland.s, p. 152.

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

Ti/pe species. — Erinaceus europseus Linnaeus.

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

Characters. — Skull rather short and broad, the zygomatic
breadth distinctly more than half greatest length ; posterior
palatal region conspicuously fenestrate ; auditory process of basi-
sphenoid well developed, concave, sometimes forming a half-bulla ;
external pterygoid plate rather larger than internal pterygoid
plate, formed aliout equally of ectopterygoid and a broad horizontal
outgrowth from palatine; dental formula: i ^'j^, <-'}—, P«lr^,
m -lo = 36 ; canines not diflPerentiated by form from the contiguous
teeth ; third upper molar consisting of a large j'l'otocone and
minute j)aracone, all trace of crushing sui'face absent ; body
short and heavy, the back covered with stifT, sliarply pointed
bristles of uniform length : tail shorter than hind foot.

ERINACEUS 115

BemarJcs. — About twenty-tive species are currently referred
to this genus, four of them occurring in Europe.

KEY TO THE EUROPEAN F0R:MS OF ERINACEUS.

Third upper iucisor definitely 2-rooted ; elevated

portion of posterior lower premolar 2-cusped ; a

bare area among spines at middle of forehead

^ (Spain, Balearic Islands and southern France)... E. algirics, p. 130.

Underparts clouded with brown, at least in inter-

ramial and intercrural regions (Spain and

southern France) E. a. alg!n(s, p. 131.

Underparts entirely whitish (Balearic Islands).... A', a. vagaiis, p. 1.33.
Third upper incisor never definitely 2-rooted ;
elevated portion of posterior lower premolar
3-cusped ; no bare area among spines at middle
of forehead.
Greatest upper length of maxillary greater than
or at least equal to depth of rostrum at middle ;
anterior upper premolar sub-equal to canine,
its posterior border with small though evident
cusp (Eastern).
Hind foot 40 to 43 mm. ; condjiobasal length
of skull in adult male about 58 mm. (Eastern
Germany through Bohemia and Roumania

to Greece) E. roumanicus, p. 127.

Hind foot 35 to 38 mm. ; condylobasal length

of skuU in adult male about 55 mm. (Crete) E. nesiotes, p. 129.

Greatest upper length of maxillary less than

depth of rostrum at middle ; anterior upper

premolar decidedly smaller than canine, its

posterior border with cusp obsolete or absent

(Western) E. europseus, p. 115.

Size larger, the skull in old males exceeding
59 mm. in condylobasal length.
Average colour darker, the face never clear,
pale, huffy grey, but usually with notice-
able blackish markings (Central-western

Europe) E. c. europseus. p. 120.

Average colour lighter, the face usually clear,
pale, buffy grey without noticeable black-
ish markings (Iberian Peninsula) E. c. hupanlcus, p. 122.

Size smaller, the skull in old males not ex-
ceeding 59 mm. in condylobasal length.
Colour of head and shoulders lighter than

that of contiguous spiny area (Italy) E. e. italicus, p. 123.

Colour of head and shoulders darker than

that of contiguous spiny area (Sicily) E. e. coiisolei, p. 12G.

ERINACEUS EUROPSEUS Linnteus.

(Synonymy under subspecies.)

GeoijrnplikaJ distribution. — Europe from the Mediterranean
coast to Scotland and southern Scandinavia ; west to Ireland ;
eastern limits of range not known.

DiagnoniK. — Size large (head and body about 225 to 27-5 ; hind
foot 40 mm. or more ; condylobasal length of skull more than

116 INSECTIVORA

55 mm.) ; spines coarse and heavy, extending in an unbroken
line across forehead and decidedly overtopping ears ; skull with
heavy, deep rostrum, the distance from posterior extremity of
premaxillary to jjosterior extremity of maxillary less than rostral
depth at middle ; auditory process of basisphenoid short, wide-
funnel shaped, not forming a half-bulla ; third upper incisor with
one root, this sometimes partly divided longitudinally ; elevated
portion of posterior lower premolar with three cusps (tig. 25 a).

External characters. — General form short, thick and clumsy,
the legs short, the feet large, plantigrade. Legs, tail, underparts
and head, except ci'own, clothed with coarse fur, the finer more
woolly hairs of which are about 1 5 mm. long, the longer, straight
hairs about 40 mm. in length. The rest of the body is covered
b}^ a densely-set mass of sharply pointed bristles about 25 mm. in
length and 1 mm. in diameter.* The skin on wliich these bristles
are set is loosely attached to body, and jjrovided with a special
system of muscles by which the edges of the spiny area can be
drawn together ventrally over the animal's retracted head and legs,
forming a comjjlete protection for the entire body. Feet robust
with short digits and well developed claws, those on hind feet
longest. Fore foot broad and rounded with very short, thick
fingers, the third and fourth sub-equal and longest, second slightly
shorter, fifth extending to base of fourth, first well developed
but not reaching base of second ; balls of all five large and pad-
like ; palm entirely naked ; three large, semi-coniluent pads
at bases of median digits, a small tubercle (about 2 mm. in
diameter) at base of thumb, and two large pads at back of wrist,
the outer the larger ; skin between pads wrinkled ; hind foot
much like fore foot but longer ; second, third and fourth digits
sub-equal and longest, but their claws noticeably graduated from
second to fourth, fifth digit extending to base of fourth, first not
reaching base of second ; sole naked, the tubercles as on palm
but more crowded, the two posterior sub-equal. Tail short and
thick, its length much less than that of hiud foot. Ear simple,
rounded, shorter than bristles on crown, the meatus without
valves. Muzzle moderately produced, somewhat pointed, the
muzzle-pad well developed, naked, its surface marked with minute
furrows, its lateral edgesfinely scalloped, its lower border continued
as a pair of parallel ridges extending inward to palate. Eye
well develojaed but rather small. Mammie : p 1-1, ff. 2-2,
i 2-2 = 10.

Colour. — Furred portions of body varying from dull brown to
dirty whitish, the under fur usually a dusky hair-brown, the
longer hairs lighter and more buffy. Belly often irregularly
blotched and variegated with whitish and darker or lighter
brown. Feet usually darker than sides. Cheeks and eye-ring
often darker than rest of head. Sjjines buflty at base, then with

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

EEINACEUS

117

I

a slaty area of variable width, followed by a narrow but sharply
defined buffy annulation and an obscurely darker tip. The
general effect is a coarse grizzle, tlie exact tone of which varies
considerably in the different geographical races as well as in
individuals of the same race.

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

Fig. 22.
Erinafens eitropaeus.

conspicuously wider than interorbital region, the rostrum short
and deep (ilistance from anteorbital foramen to front of pre-
maxillary less than depth through anterior root of zygoma).
Occiput and interorbital region marked by noticeable" ridges.

118 INSECTIVOEA

Ventral profile straight, the dorsal profile essentially parallel to
it from lambda nearly to front of interorbital region, then sloping
forward at a slight angle (about 15°) ; occiput squarely or some-
what obliquely truncate. General outline of occijiital region as
viewed from behind truncate-triangular, slightly more than half as
high as wide, the base of the triangle formed by line joining tips
of widely projecting mastoid processes, the apex by the narrowly
rounded or bluntly j^ointed lambdal region. Paroccipital
processes nearly as large as mastoid processes and resembling
them in form, though more slender and directed more backward.
Basisphenoid with deep median pit between bases of half-funnel
formed auditory processes, the pit continuous anteriorly with
mesopterygoid fossa. Tympanic ring open jaostero-externally, its
greatest breadth (antero-internal) about 3 mm. Inner and outer
2)terygoid plates broadly triangular, approximately alike in size
and form, each containing more of the pterygoid than palatine
element. Hamular short, strongly curved. Mesopterygoid sjiace
slightly longer than broad. Palate terminating posteriorly in a
high transverse ridge and strongly projecting median spine, the
ridge nearly straight, its median portion well developed. In
front of I'idge the palatine bones are conspicuously and irregularly
fenestrate. Lambdoid crest high. Sagittal crest low but
evident, extending forward to back of interorbital region. Here
it divides into two low, diverging ridges which pass forward
toward lachrymal region. In some specimens they can be traced
as far as the high, well defined ridge which occupies edge of orbit
for a distance of about 7 mm. above lachrymal foramen. Ante-
orbital foramen small, separated from lachrymal foramen by a
space much greater than its own diameter, its anteiior border
over anterior root of large pi'emolar. Upper portion of maxillary
rather short, its length behind posterior point of premaxillary
less than depth of rostrum at middle. Posterior termination of
premaxillary variable in form : nearh' squai'e, broadly or narrowly
cuneate, rounded, or rounded with supplemental inner spicule.
ManiliV)le short and heavy, the greatest depth of I'amus about one-
third length of alveolar line. Coronoid process high, narrow,
sharply hooked backward at tip. Angular process about as wide
as coronoid process, but not so long, its apex slightly bent
inward.

Teeth. — General aspect of teeth as compared with that in
other European members of the order, short, heaA^y and blunt,
distinctly omnivorous rather than strictly insectivorous in type.
Anterior upper incisor about twice as high as the succeeding
small teeth, its shaft subterete, flattened posteriorly, directed
slightly forward and inward, the teeth separated at base by
space about ecjual to height of shaft, at tip by about half this
distance. The four succeeding teeth (two incisors, canine and
anterior premolar) are essentially alike in form, the crown
slightly longer than wide, its height slightly greater than length,

ERINACEUS

119

the blunt point of conical cusp somewhat in front of middle of
crown. Of these four unicuspid teeth the first is smallest, the
second and thii'd sul>equal and larger, the fourth intermediate.
On posterior side of crown of each unicuspid there is a faintly
developed ridge extending to apex of cusp, this ridge tending to
rise posteriorly, especially in fourth unicuspid, to form a verj'
rudimentary secondary cusplet. First and second incisors invari-
ably single-rooted. Third incisorsingle-rooted, but root occasionally
showing trace of longitudinal furrow on outer side. Canine
usually single-rooted, the root with or without longitudinal
furrow ; but in a small series of specimens every stage may be
observed from this condition to a completely two-rooted tooth,
each root with a distinct alveolus. First premolar single-rooted,
the root of ten showing traces of longitudinal division, and perhaps
rarely double. Anterior lower incisor essen-
tially like the corresponding upj^er tooth
but not so high. Its shaft is directed
obliquely forward in line with symphysis,
the teeth of opj^osite sides parallel, separated
throughout by a narrow space.
Three succeeding teeth unicuspid,
the crowns similar in outline to
those of upiaer unicusjiids, but an-
terior cusps obsolete and posterior
cusplets relatively better developed.
Second upper premolar scarcel)"
broader than the unicuspids, but
three-rooted and with a distinct
protocone, metacone and postero-
external commissure. Large upper
premolar with well developed pro-
tocone, hypocone, metacone and pos-
tero-external commissure, the cusps
much as in the molars except that
hypocone is relatively smaller and
metacone and its commissure larger
and more trenchant. Large lower premolar with a high anterior
three-cusped portion similar to first triangle of lower molars,
except that the metaconid is reduced to a slight thickening at
inner base of commissure of protoconid ; second triangle repre-
sented by a mere narrow ledge or thickened cingulum. First
upper molar sub-quadrate in outline, the crown slightly wider
posteriorly than anteriorly. Protocone with somewhat broader
base than the other cusps and with low ill-defined anterior
and posterior commissures. Paracone, metacone and hypocone
sub-equal, the metacone slightly larger than the others. All
three are subterete with faintl}- indicated commissures, that
extending outward and backward from metacone to rudimen-
tary metastyle the most distinct. Parastyle and mesostyle

Tig. 23.
Eriiiacetts europieus. Teeth.

120 INSECTIVORA

absent.* A small but evident metaconule. Second molar like
first but smaller, its crown area about equal to that of large
premolar, its greatest diameter anterior instead of posterior.
Cusps essentially as in first molar, except that paracone is larger
than metacone and hypocone, and metaconule is barely indicated.
Third molar reduced to a protocone nearly as large as in the
other teeth, and a rudimentary paracone, the two connected by
a cutting edge sloping obliquely outward, forward and upward.
The tooth is single-rooted. Lower molars with the usual cusps
and commissures, the cusps more terete and commissures less
trenchant than in other European insectivores. Third molar
consisting of the anterior triangle only, this somewhat smaller
than in the other two teeth.

Erinaceus EUROPyEUS EUROP^us Linujeus.

1758. [Erinaceus'} europxiis Linnteus, Systema Naturae, i, lOfch ed., p. 52

(Sweden).
1779. [Hystrix] erinaceus Blumenbach, Handbuch d. Naturgesch., p. 72

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

Nat., p. 67 (France).
1303. Erinaceus caninus Geofiroy, Catal. ISIammif. du 'Sins. Nat. d'Hist.

Nat., p. 68 (France).

1857. Erinaceus europxus Blasius, Saugethiere Deutscblands, p. 153 (part).

1858. E\_rinaccus'] canicejjs Hamilton Smitb, Jard. Nat. Libr., 2nd ed., xv

(Mammalia I), p. 148 (near Brussels, Belgium).
1897. [Erinaceus] echinus Scbulze, Abb. u. Vortr. Gesammtb. Naturw. iv,

No. 10, p. 19 (Substitute for europunis).
1900. Erinaceus europibus Barrett-Hamilton, Ann. and ilag. Nat. Hist.,

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

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

Scotland). Type in British Museum.
1910. Erinaceus europxus and E. europx-us occicloitaUs Trouessart, Faune

Mamm. d'Europe, p. 38.

Tjiije locality. — Upsala, Sweden.

Oeoijraphical distribution. — Western central Europe from
Scotland, southern Norway, and central Sweden to the Pyrenees
and Alps ; west to Ireland : eastern limits of range not known.

Diagnosis. — Size largest of the European hedgehogs (condylo-
basal length of fully adult skulls usually 61 to 63 aim., seldom
less than 60 mm.) ; colour dark, the sides and underparts seldom
if ever a light buffy grey : face with blackish area extending
from eye to muzzle.

Measurements. — Adult male and female from Upsala, Sweden :
head and body, 265 and 263 : tail, 34 and 37 : hind foot, 44
and 43. Adult male from Innerwick, Haddingtonshire, Scotland :
head and body, 218 ; tail, 17 : hind foot, 42. Adult male and

* The cingulum usually forms a small projection resembling a rudi-
mentary parastyle.

EKINACEUS 121

female from Oundle, Northampton, England : head and body,
■2id and 257 ; tail, 24 and 31 ; hind foot, 40 and 40. Adult
male and female from Maredsous, Namur, Belgium : head and
body, 270 and 251 ; tail, 39 and 33 ; hind foot, 44 and 44.
Adult male from Bouconne, Gers, France : head and body, 220 ;
tail, 20 ; hind foot, 41. Adult male atid female from St. Gallen,
Switzerland : head and body, 279 and 297 ; tail, 41 and 43 ; hind
foot, 44 and 47. For cranial measurements see Table, p. 124.

Specimens examined. — Seventy-three, from the following localities : —

Scotland: Dunphail, Elgin, 2; Gordonstown, Elgin, 2; Innerwick,
Haddingtonshire, 1 ; Glendoc, Inverness, 1 (Wilson) ; Lanarkshire, 1.

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

Wales : Cardiff, 2.

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

Norway : Asker, near Christiania, !.

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

Denmark : Copenhagen, 5 (Andersen).

Holland : No exact locality, !.

Belgium : Maredsous, Namur, 2.

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

CtERmany : Brunswick, 1 (U.S.N.M.) ; Heidelberg, 1 (U.S.N.M.) ; Strass,
near Burgheim, Bavaria, 3; Ingelheim, Rheinliessen, 1.

Switzerland : GJeneva, 2 (Mottaz) ; St. Gallen, 7 (B.M. and U.S.N.M.) ;
Uzwil, St. Gallen, 1 (U.S.N.M.) ; Heresau, St. Gallen, ! (U.S.N.M.) ; Wolf-
halden, Appenzell, ! (U.S.N.M.) ; Thurgau, 2.

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

6, 9. Crordonstown, Elginshire. W. E. Ogilvie-Grant II. !. 3. 379-

Scotland. (c & p). 380.

6. Innerwick, Haddington- W. Eagle Clarke (c & 0. 3. 13. !.

shire. p).

{Type of E. e. occidcntalis B.-Ham.)

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

6, 9. Oundle, Northampton, Hon. N. '0. Roths- !1. 1. 3. 381-

England. child (c & p). 382.

S. Graftonbury, Hereford- W. E. de Winton (c !1. !. 3. 883.

shire. & p).

* For discussion of this character see Lonuberg, Ann. and Mag. Nat.
Hist., 7th sor., v, pp. 542-544, June, 1900, and Barrett-Hamilton, I.e.,
pp. 245-246, August, 1900. The question cannot yet be regarded as
decided, since no adequate series of skulls has yet been brought together.

122

INSECTIVORA

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

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

9. Cardiff, Glamorganshire,

Wales.
6. Nenagh, Tipperary, Ire-
land. (W. SmitJuvick.)
S al. c?. Ennis, Clare.
2 6. Kilmanock, Wexford.

Asker, Christiania, Nor-

1.

way. (E).

6 juv. Upland, Sweden. {G. Lord Lilford (p'
KoltJwff.)

1 al. Holland." {Scha Coll.)

2 6. Maredsous, Namur, Bel-

gium.
6. Eoret de Bouconne, Gers,
250 m. France. (-4.
Rohnrt.)
9. Cranves - Sales, Haute-
Savoie. (.4. Robert.)
6, 9. Burgheim, Bavaria, Ger-
many. {Woltcrstorff.)
6. Ingelheim, Rheinhesseu.
2 ^, 9 juv. St. Gallen, 500 m.
Switzerland. (E. H.
ZolUkofer.)
6, 9. Thurgau, 400 m. (E. H.
ZolUkofer.)

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

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

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

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

(c & P).

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

Dr. J. Anderson (p). 93. 7. 81. 1-2.

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

R. Drane (c ir p). 11. 1. 3. 387.

G. Barrett-Hamilton 11. 1. 2. 95.

(P).

J. W. Scott (c & p). 93. 10. 30. 1-2.

Ct. Barrett-Hamilton 11. 1. 2. 93-94.

(c & p).

Christiania Tiluseum 93. 3. 1. 7.

11. 1. 1. 151.

Lidth de Jeude Coll. 67. 4. 12. 555.

Rev. G. Fouruier (c 1. 6. 2. 1-2.

A- p).

O. Thomas (p). 6. 4. 1. 10.

O. Thomas (p).

Lord Lilford (p).

C. Hilgert (c).
0. Thomas (p).

0. Thomas (p).

6. 4. 2. 1.

11. 1.1.92,150.

8. 11. 2. 5.
4. 4. 5. 27-29.

4. 4. 5. 30-31.

Erinaceus europ.ecs nisp.wicus Barrett-Hamilton.

19C0. Erinaceus curopieus liisjianicus Barrett-Hamilton, Ann. and Mag.

Nat. Hist., 7th ser., v, p. 363, April, 1900. Type in British Museum.
1910. Erinaceus curojJeeus hispanicus Trouessart, Faune Mamm. d'Europe,

p. 39.

Type loculUi/. — Seville, Spain.

Geograpltical (listribution. — Iberian Peninsula.

Diagnosis. — Size large, essentially as in E. europseus europseiis
(condylobasal length of skull in old individuals about 60 mm.) ;
general colour paler than in the typical race, the fur often a
uniform light buffy grey, seldom with any evident darker area
between eye and muzzle.

Colour. — There is much variation in colour, .some individuals
essentially agreeing with the palest examples of true europseus.
In its extreme phase, however, the colour is a whitish buff",
decidedly paler than the cream-buff of Ridgway, the muzzle and
region about eyes washed with ecru-drab ; feet tinged with drab ;
S2)ines the same whitish buff, about half of them with scarcely any

ERINACEUS 123

dark shading, the rest with a drab sub-terminal band, the general
effect of spiny area scarcel)- siDeckled, and nearly as pale as fur.
In the type specimen the fur is a dull cream-buff, and the feet
are washed with broccoli-brown ; spines drab brown with light
tips, essentially as in true europseus.

Measurements. — Type : hind foot (dry), 40. Adult male and
female from Burgos, Spain : head and body, 270 and 250 ; tail,
30 and 28 ; hind foot, 43 and 41. Adult male and female from
Pajares, Leon : head and body, 252 and 249 ; tail, 22 and 21 ;
hind foot, 44 and 42. For cranial measurements see Table,
p. 125.

Specimens examined.— Twenty, from the followiug localities in Spain :
Arrechavaleta, Vitoria, 1 ; Pajares, Leon, 7 ; Burgos, 5 ; Palacios de la
Sierra, Burgos, 1; Bejar, Salamanca, 2; Seville, 4.

Bemarhs. — The Spanish hedgehog is a moderately well
differentiated form. Extreme specimens are easily distinguish-
able from typical europseus ; but in general the difference between
the two races must be regarded as an average one.

i. Arrechavaleta, Vitoria, 0. Thomas (p). 8. 2. 9. 9.

Spain. {N. Gonzalez.)

(J, 2 ?, 9 juv. Pajares, Leon. (N. Gem- 0. Thomas (p). 8. 2. 9. 2-6.

zalez.)

6, 9, 9 juv. Burgos. G. S. MiUer (c). 8. 8. 4. 17-19.

2 6. Burgos. (N. Gonzalez.) 0. Thomas (p). 8. 2. 9. 10-11.

2 6 juv. Bejar, Salamanca. {N. 0. Thomas (p). 8. 2. 9. 7-8.

Gonzalez.)

i, 2 9, 1. SeviUe. {Dr. A. Ruiz.) Lord Lilford (p). 95. 3. 3. 1-4.

[Type of subspecies 95. 3. 3. 2.)

Erinaceus eueop.eus italicus Barrett-Hamilton.

1857. Erinaceus europxus Blasius, Saugethiere Deutschlands, p. 153 (part).
1900. Erinaceiis europxus italicus Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., v, p. 3G4, April, 1900. Type in British Museum.
1910. Erinaceus europasus italicus Trouessart, Faune Mamm. d'Europe,

p. 39.

Type localiti/. — Siena, Italy.

(• eographical distribution. — Italian Switzerland, Ital}' and
Sardinia ; Corsica ?

DuKjnosis. — Colour as in E. europseus europseus or slightly
paler ; size less than in the typical race, the largest skulls
probably not exceeding 59 mm.

Colour. — The colour is about as in the paler iudi^■iduals of
typical europseus, though the speckling of the spin}' area seems in
general to be finer, and the underparts usually lack all heavy
dark clouding.

Jleasureuients. — Adult female irom Curoggio, Ticino, Switzer-
land : head and body, 250; tail, 29 : liind foot, 41. Two adult

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126 INSECTIVORA

males from Siena, Italy : liead and body, 210 and 220 ; * tail, 30
and 28 ; ''' hind foot, 40 and 42.* Two adult females from the
same locality : head and body, 200 and 208 ; tail, — and 32 ;
hind foot, 43 and 38. For cranial measurements see Table,
p. 125.

Specimens examined. — Seventeen, from the following localities : —
Switzerland : Bigorio, Ticino, 1 (U.S.N.M.) ; Curoggio, Ticino, 1
(U.S.N.M.); Gentilino, Ticino, 1 (U.S.N.M.).

Italy : Empoli, Florence, 1 ; Siena, 4 ; Ostia, Rome, 1 ; Rome, 4.
Sardinia : Su Cramu, 1 ; Bare, 1 ; Marusei, 1 ; Trecorgia, 1.

Remarks. — Though not so pale as the Spanish race the Italian
hedgehog seems worthy of recognition as a form distinct from
true eurupseus. Its status is at present unsatisfactory, owing to
the lack of sufficient material ; but specimens from south of the
Alps seem never to attain the large size of Central European
adults. The Sardinian si^ecimens that I have seen are in general
paler than those from the mainland ; but here again the material
is insufficient. A hedgehog is known to occur in Cor.sica, but
no specimens have yet been compared with the Italian race.

9. Empoli, Florence, Italy. A. H. Savage Laudor 97.3.7.1.

(c & p).
2 ^, 2 9. Siena. (S. Brorji.) Dr. E. Hamilton (p). 98. 10. 2. 5-8.

(Type of subspecies 98. 10. 2. 5.)
1. Ostia, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 7.

3 6. Rome. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. 3-4. 96.

6. Su Cramu, Sardinia. (IT. O. Thomas (p). 0. 12. 3. 5.

WoUerstorff.)

6. Bare. (17. Woltersforff.) 0. Thomas (p). 0. 12. 3. 3.

9. Marusei. (W. Wolterstorff.) 0. Thomas (P). 0.12.8.6.

9 juv. Trecorgia. (ll'.iroZ^_Tsto>^.) 0. Thomas (p). 0.12.3.4.

Erinaceus europ.eus consolei Barrett-Hamilton.

1900. Erinaceus europxus consolei Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., v, p. 866, April, 1900. Type in British iluseum.
1910. Erinaceus europxus consolei Trouessart, Faune INIamm. d'Europe,

p. 40.

Type locality. — Palermo, Sicily.

Geographical distrihiUion. — Sicily.

Diagnosis. — Size apparently as in E. enrnp'ceus italicus (only
known specimen imperfect) ; colour differing from that of
italiciis in the uniform dusky brown head and shoulders ; quills
unusually robust, their dark and liglit markings strikingly con-
trasted.

Colour. — Whole head and sides of neck and shouldei'S a
uniform dark brown between hair-brow^i and sepia, sprinkled
with bufty grey hairs. The dark brown continues back along

* Type.

ERIXACEDS 127

edge of sj^iny area to tail, but throughout this region it is overlaid
by the uniform light cream-buff of underparts. Spines very dark
drab with light cream-buff tips, the light area shorter on spines
of mitldle of back than on those of sides, thus producing a slight
thougli evident darker median dorsal area. Feet so injured that
colour cannot be determined.

SJciiJI and teeth. — The imperfect skull shows no peculiarities.
Teeth as in specimens from the mainland.

Medsiirementit. — Type (sex not known) : head and body, 252 ;
tail, 50 ; hind foot, 40. For cranial measurements see Table,
p. 125.

Specimen examined. — The type.

Bemai-ks. — If not an abnormal specimen of E. europaeus italicns
the type of console! represents a very distinct local race.

1. Palermo, Sicily. J. I. S. Whitaker (p). 98. 10. G. 1.

{Tijpe of subspecies.)

ERINACEUS ROUMANICUS Barrett-Hamilton.

1900. Erinaceus europxus roiimanicus Barrett-Hamilton, Ann. and Mag.

Nat. Hist., 7th ser., v, p. 365, April, 1900. (Gageni, Roumania.)
Type in British Museum.

1901. Erinaceus danuhicus Matschie, Sitz.-Ber. Gesellsch. Naturforsch.

Freunde, Berlin, p. 229, December, 1901. (Prundu, Roumania.)
1910. Erinaceus europxus roiimanicus and E. curopxHs danuhicus Troues-
sart, Fauue Mamm. d'Europe, pp. 40-41.

Type locality. — Gageni, Pi-ahova, Roumania.

Geographical distribution. — From eastern Germany (Konigs-
berg) and northern Bohemia south through Hungarv, into
Greece.

Diagnosis. — In general like Erinacen.s europsens, but colour of
furred parts usually much darker, and chest often with a lai'ge,
conspicuously contrasted whitish area ; skull with upper length of
maxillary greater than depth of rostrum at middle ; teeth
essentially as in E. curopeeus, but first upper premolar with
postero-basal cusp usually more distinct.

Colour. — Furred area dark hair-brown interspersed with bufiy
and whitish hairs, the former most numerous on sides, the latter
forming a clear whitish pectoral area which sometimes spreads
over entire underparts. Feet and tail sepia. Face usually with
a seal-brown suffusion. Claws blackish horn-colour. Quills
with colours usually less contrasted than in E. europeeus, the
general effect of the spiny area darker and less speckled.

Shull and teeth. — The skull differs from that of E. eurrqjieus
in the relati^■ely longer, less deepened rostrum, a peculiarity
which seems chiefly to involve the maxillary bone. Upper length
of maxillary greater than depth of rostrum at middle. Posterior

1 28 INSECTIVORA

transverse palatal ridge usually lower and less developed,
particularly at middle, where there is often a slight angle.
Ridge at margin of orbit in lachrymal region not so long as in
E. curojjseus, and less distinctly marked off from general contour

Fig. 2-J.
Erinaceus rounuinicus. Nat. size.

of skull, its degree of development somewhat as in E. ahjirus.
Teeth not certainly distinguishable from those of Erinaceus
ruropseus, though anterior upper premolar is usually larger
relatively to canine, and its posterior cusplet tends to be better
developed.

Measurements. — External measurements of adult male and
female from vicinity of Konigsberg, Germany : head and body,
285 and 280; tail, 21 and 22; hind foot, 42 and 41. Type
(adult female) : head and body, 206 ; tail, 24 ; hind foot, 40-6.
Adult female from Corfu, Greece : head and body, 263 ; tail,
37 ; hind foot, 43 ; ear, 29. Adult male from Cephalonia,
Greece: head and body, 260; tail, 35; hind foot, 42; ear, 31.
For cranial measurements see Table, p. 132.

Specimens examined. — Eleven, from the following localities : —
Germany: Near Konigsberg, 3 (U.S.N.IM.).

KrsiNACEUS 129

Austria-Hunc4ary: Haida, Arva, Bohemia, 1; Vasoiir Eiscnbur^
Hungary, 1 (U.S.N.M.). °'

EoDMANiA : Gageni, Prahova, 2.
Greece: Corfu, 2; Cephalonia, 1 ; Tatoi, near Atheus, 1.

BemarJcs.— Though at first sight very similar to Erinacetis
europseus, this species is easily recognizable by its cranial characters.
In most specimens there is a strong contrast between the dark
posterior portion of underparts and whitish chest, throat and
shoulders, a pattei-n which appears to be rarely if ever well
developed in the related species.

9. Haida, Bohemia. Lord Lilford (p). 97. 8 14 1

i, 9. Gageni, Prahova, Roumania. Lord Lilford (p). 4. 4. 6. 15-16.

, „ ( "'• Dodson.) [Type of species 4. 4. 6. 16.)

.^. 1 otamos, Corfu, Greece. .J. I. S. Whitaker (p). 8 10 1 G

{C. Mottaz.) ■ ■ ■

■?. Argostoli, Cephalonia. J. I. S. Whitaker (p). S 10 1 7

(C. Motfaz.) ....

i. Tatoi, Athens. C. Mottaz (c). 8. 11. ,3. S.

ERINACEUS NESIOTES Bate.

1906. Erinaceim ciimjxns ncsioti's Bate, Proc. Zool. Soc. Loudon 1905 ii
p. 316, April 5, 1906. Type in British Museum. ' ' '

1910. Erinaceus europxvs ncsiotes Trouessart, Faune Mamm d'Eurone
p. 40. ^ '

Type locality. — Near Gonia, western Crete.

Gt'ograpMcal distribution.— Island of Crete.

-Dm»7«o.s;s.— Similar to Erinaccui^ roumanicm but smaller (hind
fo_(>t, 35 to 3S ; condylobasal length of skull in adult male, about
55 mm.) ; spines not so coarse as in the related species ; first
upper premolar distinctly larger than canine and of essentially
the same height.

(Jolour.—The colour is like that of E. roimnnicux. Under-
parts, sides and face dull whitish grey with faint dark clouding
between eye and muzzle, and in one of the three skins with a
dai-k wash on posterior half of under])arts.

Skull.— As in E. roumamnis, but not attaining as large size.

Mra.vu-ements.~Adu\t male"- and female from the type
locality: head and body, 208 and 204; tail, 29 and 19 • hind
foot, 40 and 38 (dry, 38 and 35). For cranial measurements see
lable, p. 132.

SjK'cimens f.vcnnined.—lhreL', all from Crete.

Bemarks.— The Cretan hedgehog is nearly related to Erinaccu>t
nmmuniruH, though well differentiated by its smaller size and by
the relatively large fourth unicuspid tooth.

'-, V. Gonia, Crete. Miss D. Bate (c). 5 10 o ii_i^

-,,,,. (5. 12. 2. 11 Type of speciesO ^

V. Mcsoghia. Miss D. Bate (c). 5. 12. 2. 13.

* Type.

130

IXSECTIVORA

ERINACEUS ALGIRUS Duvernoy and Lereboullet.

(Synon3-my under subspecies.)

Gcogrupliical distrihution. — Northern Africa ; also in southern
Spain, southern France, and the Balearic Islands.

Diagnosis. — Smaller than Erinaceiis europsens (hind foot less
than 40 mm.) : sjiines not so coarse ; middle of forehead with
bare area, among the spines ; skull with sagittal crest extending
forward to middle of frontal : a wide flattened area on each side
of bony palate behind transverse ridge ; third upper incisor with
two perfectly distinct roots ; elevated poi'tion of posteiior lower
premolar with only two cusps (fig. 2b h).

External characters. — Externally Erinaceus alijlnis is dis-
tinguishable from E. eurofseus by its smaller size, shorter, more
slender and apparently more densely-set bristles, and by the bare
area at middle of forehead. This bare area is about 7 mm. wide
and extends back about 10 to 15 mm. from front line of spines.
In dried skins it is sometimes partly hidden by shrinking. Fur
more dense and less coarse than in the larger animal. Claws on
front feet seldom attaining a length of 8 mm.

Colour. — The colour resembles in general that of the paler
races of E. euro^setis.

Slull. — The form of the skull is essentially as in E. i nropseus
except that rostrum is less elevated posteriorly, so that the dorsal
profile tends to become slightly concave. Sagittal crest when
fully developed extencUng forward to middle of frontal, while in
E. eiiropseiis it is usually confined to parietals, rarely encroaching
on posterior edge of frontal. Ridge at margin of orbit veiy short,
scarcely more than a process above lachrymal foramen. Bony
palate extending behind transverse ridge as a M-ell-defined Hat area
divided along median suture by a longitudinal ridge representing
the median spine of E. europaeus. Basisjahenoid pit narrower,
relatively deeper, and with more overhanging edges than in
E. enropsetis.

Teeth. — In general the teeth show no departure from those
of E. curopstus. The posterior lowei' pie-
molar, however, lacks all trace of the meta-
conid, so that the resemblance of the elevated
portion of the tooth to the first triangle .of
iH^ and 7/(.j is completely destroyed. In the
upper jaw the third incisor, canine, and
a h first premolar are two-iooted, apparenth'

i'lG. 25. without exception.

Large lower premolar of Measurements. — Head and body about

a.uiE.^aigiru!fh).'^i^l 200 to 250 ; tail, 25 to 40 : hind foot, 32 to
37 ; condylobasal length of skull, 54 to 59 mm.
Ilemarlcs. — This species is readily distinguishable from £v/«ace;f*"
europ»U8 by the bare spot among spines of forehead, the perfectly
two-cusped large lower premolar', and the t^\"0-rooted third upper

liRINACEUS 131

incisor. Although occurring wild in southern Spain, on the
Balearic Islands, and in south-eastern France, it seems not
improbable that the animal owes its presence in Europe to the
agency of man.

Ekixackus algirus AL(iiKus Duvernoy and LerebouUet.

1840. Erinaceus algirus Duvernoy and LerebouJlet, M6m Soc Mus

Q Hist. Nat. Strasbourg, iii, fasc. 2, p. 4.
1898. Erinaceus algirus de Winton, Proc. Zool. Soc. London, 1897, p. 955.

Type locality. — Oran, Algeria.

Geographical distribution.— 'i^orthern Africa ; also in .southern
Spain and south-eastern France.

J>ia(/Mos/s.— Condylobasal length of skull in individuals with
distinctly worn teeth 57 to 59 mm. ; underparts dusky through-
out or with at least an evident dark wash in interramial and
intercrural regions.

CoZor«»-.— Furred area buffy white to base of hairs, except on
muzzle, cheeks, interramial region, a narrow band aloncr sides
bordering quills and spreading posteriorly to cover tail hind
legs and intercrural region, all of which are a dark brown very
nearly the bister of Ridgway. Feet a lighter shade of the' same
brown. Occasionally the brown suffusion extends over most of
ventral surface. Quills dull horn-colour, each with a whitish
sub-terminal area about 7 mm. in length, the extreme tip usually
(ark. Throughout the spiny area the whitish strongly pre-
dominates, especially when animal is viewed from in front
Claws light yellowish horn-colour.

Measurement.s.~Adult male from Schaf-el-Kab, Morocco
(teeth much worn) : head and body, 206 ; tail, 2(i ; hind foot 32
For cranial measurements see Table, p. 1.32. '

Sp('ci77iens examined.— Nnmeious specimens from Northern Africa • also-
an adult from " Andalucia," Spain; a young, less than half grown, from
h.lche, Alicante, Spain ; and a still younger .specimen from Lecques, Var

Bcmarks.— Owing to the unsatisfactory nature of the Hpani.sh
and French material the status of the Continental European
hedgehogs of the Erinaceus algirus group is at present doubtful
Should they prove to be identical with tlie North African form
It would seem probable that they have been introduced within
historic times. The specimen from Elche, though undoubtediv a
wild-bred animal, is too young to be positively determined as' to
geographical race. The same is even raoi-e true of that from Var
The adult from " Andalucia " lacks detailed history. Externally
it resembles the African form as compared with E. ah/irns vaqans
but the skull is small,* es.sentially as in the Balearic" race. "

* Coudylolmsal length, 53-8; zygomatic breadth, 32-0; least inter-
orbital^breadth, 14-8 ; mandible, 41 -2 ; upper tooth-row, 27-0; lower tooth-

132

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KRINACEUS 133

1. Andalucia, Spain. Lord Lilford (p). 94. 6 11 5

_i juv. Elche, Alicante. G. S. Miller (c). 8. 8. 4. 1(3.

juv. al. Lecques, Var, France. Dr. P. Siepi (p). 98. 8. 25. 1.

Erinaceus ALGiRu.s VAGANs Thoma.s.

1901. Erinacens alr/irus imgans Thomas, Proc. Zool. Soc. London, p. 38.

Type in British Museum.
1910. Evinaccus algirus vagans Trouessart, Faune Mamm. d'Europe, p. 39.

Tyi^e locality. — Qsrn Cristobal, Minorca, Balearic Islands.

Geographical distribution. — Balearic Islands.

Diatjnosis. — Condylobasal length of skull in individuals with
worn teeth about 54 mm. ; furred area of body whitish through-
out, with no evident dark wash on face, in interramial or
intercrural regions, or on feet.

Measurements. — Type (adult male) : head and body, 250 ;
tail, 40 ; hind foot, 37. Adult male from Inca, Majorca : head
and body, 218 ; tail, 31 ; hind foot, 35. For cranial measure-
ments see Table, p. 132.

Specimens examine cl.~0\x6 from ^Majorca and five from Minorca
Balearic Islands. '

Bemarlcs. — The Balearic form of Erinaceus algirus is dis-
tinguishable from the African race by its paler colour and smaller
size. Its relationship to the animal occurring on the mainland
of southern Spain is not at present clear.

i. Inca, Majorca ; 300 m. O. Thomas & R. I. Pocock 0. 7. 1. 6.
Balearic Islands. (c & p).

4 6, 9. San Cristobal, ]\rinorca. 0. Thomas & R. I. Pocock 0. 7 1 35-39

(c & p).

(0. 7. 1. 36. Type of subspecies.)

134

CHIKOPTERA.

Order CHIROPTERA.

1779. Gldmptcra Blumeiibacli, Handbucb der Naturgescliichte, -p. 74.

Geofirajilrieal disirihutiou. — Practically cosmopolitan ; only
absent from the treeless arctic and antarctic I'egions, and from
the most remote islands of the Pacific and South Atlantic
oceans.

Characters. — Terrestial placental mammals with the anterior
limbs modified for true flight, the fingers greatly elongated (third
usually at least as long as head and body) and joined together
by a membrane which extends to sides of body and legs ;
shoulder girdle much more developed than pelvis, the sternum
usually keeled ; knee directed backwaixl.

Beniarhs. — ^The order Chiroptera, containing the only living
vertebrates, except birds, capable of true flight, is the most
sharply circumscribed of the main groups of mammals. Not
only are its living members invariably distinct from all other
recent forms, but the fossils also are, so far as known, equallj"
well differentiated. Therefore no intermediate stage has yet
been found connecting the bats with any other order. That
they are, however, not distantly related to the Insectirora, is
shown by numerous peculiarities of structure, among others the
relative!}' simple character of the brain. Two sub-orders are
recognized among the recent members of the order, the
MeijacMroptcra, not represented in Europe, with less highly
modified skeleton of fore limb, and more modified teeth, and the
Microchiropiera with more highly developed wing and in most
instances more primitive teeth.

Sub-Order MICROCHIROPIERA.

18'21. Inscdivora Gray, London Medical Repository, xv, x^. 299, April 1,

1821.
1872. Animalivora C-i-ill, Arrangement of the Families of Jlammals, p. IG,

November, 1872.
1875. Microcliiroptcra Dobson, Ann. and Mag. Nat. Hist., 4tli ser., xvi,

p. 346, November, 1872.
1878. Microcliiroptera Dobson, Catal. Ghiropt. Brit. ]Mus., p. 2.
1907. Microcliiroptcra Miller, Families and Genera of Bats, p. 78, June 29,

1907.

GeographicaJ di.vtribtit ion. -Thf: saiue as that of the order.
In Europe north to the limits of tree growth, west to Ireland
and the Azores.

MICROCHIROPTERA

135

Characters. — Anterior limb ^ery highly modified, the second
linger scarcely if at all independent of third, its ungual phalanx
never present, the humerus with trochiter and trochin large, the
former usually articulating with scapula ; mandible with angular
process well developed, long and narrow ; teeth usually not
modified for frugivorism (never in European species), the cheek-
teeth of upper and lower jaws very difierent from each other
(except when excessively reduced, as in the South American
Dcsmodontidse) ; margin of ear not forming a ring ; tragus usually
present. Mammaj in all European genera, jj 1— 1 = 2.

BemarJcs. — The sub-order Microchiroptera is essentially cos-
mopolitan in distribution. Though more highly modified than
the Megacliiroptcra in wing structure, the members of this group
foi' the most part I'etain the primitive tuberculo-sectorial type of
molar tooth, though certain South American frugivorous forms
show the stages through which the Megachiropterine molars
have probably passed. At jd resent 17 families and nearly 150
genera are recognized ; the species ai-e too imperfectly known to
permit any approximate estimate of their number. Three
families and ten genera are found in Europe.

KEY TO THE EUROPEAN FAMILIES AND SUB-FAMILIES
OF MICROCHIROPTERA.

Tragus absent ; muzzle with leaf-like outgrowths ;
premaxillaries represented by palatal branch
only, not fused with surromiding parts (often

lost in prepared specimens) PJdnolophidx, p. 136.

Tragus present ; muzzle without leaf-like out-
growths ; premaxillaries represented princi-
pally (entirely in European genera) by nasal
branch, very early and completely fused with
surrounding parts.
Fibula robust, its diameter about half that of
tibia ; tail projecting conspicuously beyond
hinder edge of narrow interfemoral mem-
brane Molo&sidx, p. 276.

Fibula very slender, its diameter much less than
half that of tibia ; tail scarcely or not pro-
jecting beyond hinder edge of broad inter-
femoral membrane Vespcrtilionidie, p. 165.

Presternum with median lobe much smaller
than body of bone ; coracoid curved out-
ward ; second phalanx of third linger not

specially elongated Vespei-tilioninie, p. 165.

Presternum with median lobe larger than
body of bone ; coracoid straight, directed
inward ; second phalanx of third finger
nearly three times as long as first Miniopteriiue. p. 26S.

i;56

CHIROPTEEA

KEY TO THE GENERA OF EUROPEAN BATS.

(A wholly artificial key based primarily on external characters.)

^luzzle with leaf-like outgrowths Rhinolophus, p. 137.

Muzzle without leaf -like outgrowths.

Tail projecting conspicuously beyond membrane ... Xi/ctinomus, p. 276.
Tail not projecting conspicuoush' beyond membrane.
Ears joined.

Ear longer than head Plccotus, p. 25G.

Ear shorter than head Barbastella, -p. 268.

Ears separate.

Second phalanx of tliird finger nearly throe

times as long as first Miii'wptenis, p. 268.

Second phalanx of third finger less than twice
as long as first.
Fifth finger alsout as long as metacarpal of

fourth or third Nydalus, p. 242.

Fifth finger much longer than metacarpal of
fourth or third.
Ear wider than high, its lower margin
forming a small pocket near angle of

mouth Vcispciiilio, p. 2.38.

Ear higher than wide, its lower margin
not forming pocket near angle of
mouth.

Upper cheek-teeth 6-6 Mijotis, p. 166.

Upper cheek-teeth less than 6-6.

Upper cheek-teeth .5-5 Pipistrcllus, j). 202.

Upper cheek-teeth 4-4 Ei^tcsic/is, p. 224.

Family RHINOLOPHID^.

1827. Bhmulophina Lesson, Man. de jMammalogic, p. 81 (part).
1857. Phyllostomata Blasius, Saugethiere Deutschlands, p. 26.
1866. Rhinolophidai Gray, Proe. Zool. Soc. London, p. 81 (part).
1878. Bhiiiolophidie, Dobsou, Catal. Chiropt. Brit. Mus., p. 100 (x)art).
1907. RJiinolopliidx Jliller, Families and Genera of Bats. p. 106, .June 29,
1907.

Geographical clistrlbution. — Tropical and temperate portions of
the Old World from Ireland east to the Philippine Islands,
Solomon Islands and north-eastern Australia. In Europe north
to northern England and the Baltic coast of Germany.

Characters. — Ear without tragus ; muzzle with conspicuous
leaf-like cutaneous outgrowths {i\g. 26) consisting of a horizontal
anterior horseshoe, a perpendicular median sella, and a posterior
erect lancet ; skull with premaxillaries rejaresented by palatal
branches only, the two bones partly cartilaginous and not fused
with surrounding parts (often lost in prepared specimens) :
.shoulder girdle highly abnormal, the seventh cervical and first
dorsal vertebrae, first and second ribs, and presternum fused into a
continuous ring ; secondary articulation of humerus with scapula
small l)ut distinct ; fibula thread-like ; foot normal, the hallux
with two phalanges, the other toes with three.

liciaarJi-s. — The I\lnnolophid;r are the most widely distributed

HHINOLOPHUS

137

of the Old World leaf-nused bats, and the only t'ainil}' known
to occur in Europe. Notwithstanding its extensive distribution
and its large number of species the group is represented by a
single genus.

Genus RHINOLOPHUS Lacepede.

1799. Hliinolophiis Lacepede, Tabl. des div. sousdiv. oidrcs et genres des

Maminif^res, p. 15 {femtm-cqtiinum).
1836. Rhinocrepis Gervais, Diet. Pittoresque d'Hist. Nat. iv, pt. 2, jj. 617

(attributed to Geoffroy and Cuvier, Mag. Encyclop6dique, 1795,

but the name does not occur in the paper alluded to).
1847. Aquias Gray, Proc. Zool. Soc. London, p. 15 (lucius and irifoliatus).
1857. lihinoloplius Blasius, Siiugethiere Deutschlands, p. 26.
1866. Phyllotis Gray, Proc. Zool. Soc. London, p. 81 {pliilijiprnsis) not

Phijllotis Waterhouse, 1837.
1866. CcelophyUuH Gray, Proc. Zool. Soc. Loudon, p. 427 {coslophyUus).
1878. Rhinolophus Dobson, Catal. Chiropt. Brit. Mus., p. 100.
1901. Euryalus Matschie, Sitz.-Ber. GeseUsch. naturforsch. Freuude,

Berlin, p. 225 [mchelyi).
1904. Euryalus Matschie and Andersen, Sitz.-Ber. Gesellsch. naturforsch.

Freunde, Berlin, p. 71 (euryalc group).
1907. Jlhinolophus Miller, Families and Genera of Bats, p. 108, -Tune 29,

1907.

Type species. — Vespertilio ferrum-equinum Schreber.

Geographical distribution. — Same as that of family (p. 136).

Cliaractcrs. — Dental formula: « .7-^^ , c i^ , j)m |~| , )h |^ = 32.
Upper incisor very small, but usually well formed and with
distinct rounded crown with slight cusp on inner side. Lower
incisors tritid, the outer larger than inner, the four teeth
forming a continuous row between canines. Upper canine
heavy, but without secondary cusps or conspicuous cingulum.
Lower canine rather weak. Anterior upper premolar (jnn^) and
middle lower premolar {pm.^) small, functionless, usually crowded
Huite out of tooth-row. Other teeth showing no special peculiari-
ties ; m^ and m- ^\'ithout hypocone, m^ with live cusps and
three commissures (in many species a rudimentary- fourth com-
missure), the crown area much more than half that of m^ or
m'-. Skull with large brain-case and much shortened, globularly
inflated rostrum, beyond which the maxillaries, bearing the large
canines, conspicuously project ; palate so deeply emarginated
both anteriorly and posteriorly that its median length is less
than least distance between tooth-rows. Tail well developed,
e.xtending to edge of wide interfemoral membrane. Calcar
slender. Ears large, .separate, without tragus. Muzzle with
conspicuous leaf-like cutaneous outgrowths, consisting of a hori-
zontal anterior horseshoe, a perpendicular median sella and an
erect posterior lancet (fig. 26).

Remarks. — Among European bats the members of the genus
lihinoloplms are at once recognizable by the presence of the nose-

138

CHIP.OPTERA

leaf and absence of tragus. The skull dift'ers from that of all
other members of the fauna" in the short, globularly inflated
rostrum and long, projecting maxillaries, between which lie the

Xoseleaf of RhiiKilopIn'ifen-mn-equinum (a), li. hrppaxiilerus (b), 11. /■iii-yole (o),
and R. blasii{A). Nat. size.

horizontal free premaxillaries (often lost in prei)ared specimens).
About 100 forms have been described,* eight of which occur in
Eui'ope.

KEY TO THE EUROPEAN FORMS OF RHINOLOPHUS.

Noseleaf with connecting process broadly
rounded above ; skull with nasal swellings
long, rising gradually above line of fore-
head.
Forearm over 50 mm. ; condylobasal length ^

of skull over 20 mm. ; sella pandurate ;
large upper premolar in contact with

canine (Greater Horseshoe) B. ferrum-eqitiniim, -p. 139.

Wing relatively long ; forearm 54-58 mm.,
longest finger 84 to 92 mm. (Southern

and central Continental Europe) i:. f. ferruvi-eijuiinnn, p. 142.

Wing relatively shorter ; forearm 52 to
55 mm., longest finger 83 to 88 mm.
(England) Ji. f. inaulanus, -p. 147.

* See Andersen, Ann. and Ma,g. Nat. Hist.. 7th ser., xvi, pp. G48-662.

RHINOLOPHUS 139

Forearm under 43 mm. ; condylobasal length
of skull under IG mm. ; sella cuneate ;
large ujJiper premolar not in contact

with canine (Lesser Horseshoe) 7?. hipiiomdews, p. 147.

Greatest length of skull 14 • -5 to 15-5 mm.

(Jlediterranean region) it. h. minimiu^, ]}. 151.

Greatest length of skull more than
15* 5 mm.
Forearm 80 -3 to 39 mm. (England and

Ireland) n. /,. m!inau>f, p. 154.

Forearm 89 to 41 -7 mm. (Central Fjurope) j;. h. hiiiimxidemit, p. 149.
Noseleaf with connecting process acutely
pointed above ; skull with nasal swellings
short, rising abruptly above level of fore-
head.
First phalanx of fourth finger more than
half as long as second ; sella bluntly
cuneate ; no marked contrast between
crown areas of anterior and posterior
lower premolars (Eastern Mediterranean

region) li. blasii, p. 162.

First phalanx of fourth finger less than half
as long as second; sella parallel-sided,
broadly rounded above ; a marked con-
trast between crown areas of anterior
and posterior lower premolars.
Size smaller, forearm 44-6 to 49 mm.,
upper tooth-row 6*2 to 6*6 mm.;
gradation between phalanges of fourth
. finger abrupt (ratio of first to second
about 38) ; ])oint of lancet gradually

narrowed, never linear R. curtjalc, p. 155.

Size larger, forearm 48 '6 to 51 '4, upper
tooth-row about 7 mm. ; gradation
between phalanges of fourth finger
less abrupt (ratio of first to second
about 44) ; point of lancet linear 7?. mclicliji, p. 159.

RHINOLOPHUS FERRUM-EQUINUM Schreber.

(S^^lonymy under subspecies.)

(reograpliicdl (Jistribntion. — From southern Japan and China,
through the Hiiiiahxyas, the Mediterranean sub-region (exclusive
of Egypt), and central Europe to southern England (Andersen).

Diar/noftis. — Hize largest of the European species (forearm
more than 50 mni., condylobasal length of skull about 21 mm.,
mandible, 15 to IG mm.); noseleaf with horseshoe less than
10 mm. wide, the sella pandurate, the connecting process low,
abruptly rt)unded ; fourth finger with first phalanx considerably
mor(! than half as long as second ; large upper prenujlar broadly
in contact with canine, the small ])remolar minute (sometimes
absent), completely external to tooth-row.

Extern<d characters. — Size large and form leather heavy
(among the European members of the genus). General outhne
of noseleaf a rather elongate ovate-pyriform, the width of horse-
shoe slightly greater than distance from flat area at base of sella

140

CHIROPTERA

to ti]i of lancet. Sella broadly rounded at tip, noticeably con-
stricted somewhat above middle, the resulting outline pandurate.
Connecting process rising slightlj" but evidently above level of
sella, its upper border straight anteriorly, its tip rather abruptly
rounded off. Eai' large, extending when laid forward slighth'
beyond extremity of muzzle, abruptly narrowed to a rather acute
recurved tip ; antitragal lobe less than half as high as conch,
its width about eclual to its height, its upper border nearly
horizontal. Wings broad, the membrane attached to ankle.
First phalanx of foui-th linger noticeably more than half as long
as second. Foot slender, nearly half as long as tibia.

Colour.- — General effect a light greyish or drabby brown
pi'oduced by varying combinations of the pale ecru-drab under
colour and the darker hair tips, the region between ears, across
shoulders and at sides of lumbar region usually jialer than back.
The colour of the tips is usually either a clear, rather light .sepia,
or a buffy brown resembling the wood-brown of Ridgway. While
intermediate shades occvir the extremes are more often met with,
evidently representing two dichromatic phases. Underparts
usually rather lighter than back but never approaching whitish.
Slciill. — General outline of skull long and narrow, the breadth
of brain-case much less than twice that across canines, and con-
tained about 2 A times in greatest length ; zygomata projecting
slightl}' beyond general outline ; interorbital region greatly con-
stricted and conspicuously hour-glass shaped ; occipital portion of
brain-case noticeably overhanging foramen
magnum and marked oti' from main j^ortion
Iw a slight transverse depression corre-
sponding to suture between parietals and
unusually large interparietal ; sagittal
crest well developed, extending forward to
nari'owest portion of interorbital region :
lambdoid crest low but evident ; auditory
bulla' small, covering less than half surface
of large cochleae, between which the floor
of brain-case is reduced to a longitudinal
bridge less than 1 mm. in diameter :
mesopterygoid fossa slightly widei-
anteriorly than posteriorly, about li
times as long as wide, its rounded
anterior margin at level of postero-
internal angle of ;/(- ; palate with an
evident emargination on each side ex-
tending between m^ and mesopterygoid space ; anterior palatal
emargination extending back to line joining protocones of
anterior molars ; premaxillary ligulate, somewhat wider pos-
teriorly than anteriorly, the innei' border entire, the outer border
with a deep almost circular emargination posteriorly, the two
bones closely applied to each other along inner margin and to

y\i^£^

Jihinolopliusferrum-eqiiimi
Nat. size.

liHINOLCIPHUS

Ul

bottom of palatal emai'gination posteriorly, but otherwise free:
rostral inflation evident but low, its posterior border running
gradually into that of interorbital region, its anterior margin
over middle of anterior molar ; maxillary triangular in outline
when viewed from the side, the heavy canine projecting forward
and downward from its anterior apex ; anteorbital foramen small,
over middle of second molar and directly beneath minute
lachrymal foramen, the plate forming outer wall of canal
thread-like, occasionally absent.

Teeth. — Except for the minute ]iremolars and upper incisors the
teeth are robust and heavy relatively to size of skull. Upper
incisor minute, low, the crown subterete, wider than root, lower
externally than internally, each tooth placed near middle of
oblique anterioi' border of premaxillary, the space between the
two nearly double diameter of crown. Lower incisors strongl}-
imbricated, forming a short, very convex row between canines,
their ci-owns longer than high, compressed (the outer tooth less
than the inner), deeply and et|ually trilid. Upper canine very
large, and noticeably the highest tooth in the maxillary series,
the root oblique, the shaft abruptly bent downward at level of
well developed cingulum, its length along cinguluin about three-
fourths height ; cross section of shaft triangular, the inner surface
flattened, slightly concave neai' cingulum, the posterior edge
trenchant, the antei-ior edge narrowly sub-ti'enchant, the outer
surface with well developed median longitudinal ridge and
noticeable posterior concavity. Lower canine not so large as
upper, the flattened surface of its shaft directed posteriorly to
o})pose front of upper canine in mastication. Anterior uppei'
premolar minute, resembling upper incisor in both size and form,
entirely external to tooth-row, occasionally absent. Posterior
uj)per premolar large, closely crowded against canine, its crown
area about equal to that of second molar, without secondary
cusps, its main cusp intermediate in height between canine and
metacone of first molar ; posterior border of crown slightly but
evidently emarginate. Anterior lower premolar less than half as
high as posterior premolar and with barely half its crown area,
the two teeth crowded closely together between canine and first
molar ; shaft of anterior tooth with slightly developed concave
area on inner side, that of posterior tooth with better defined
posterior concavity, its inner side convex ; middle lower premolar
resembling anterior upper premolar, its position equally external
to tooth-roM'. Upper molar with large though not unusually high
protocone, behind which in m^ and iii- there is a low-lying
lieel (best developed in »(*), but no indication of a true hypocone ;
paracone and metacone well developed, the latter slightly the
higher in m^ and m-, this reversed in m^ ; styles and commis-
sures in m^ and m- well developed, forming a normal VV-pJittern :
in »K^ the metastyle and fourth commissure are absent, and the
third commi.ssure is much reduced in length : crown area of in^

142 CHIROPTEEA

about two-tliirds that of //('- ; lower molars with protoconid
noticeably higher than the other cusps, the posterior triangle
slightly the wider of the two in m,^ and lu.^, a little the narrower
in m^ ; behind entoconid the cingvdum forms a minute though
evident accessory cusp.

Bemarks. — Among the leaf-nosed bats of Europe this species
is easily recognizable by its large size taken in connection with
the low, rounded upper margin of the connecting process of sella.
In B. meliehji, which approaches it in size, the connecting process
is sharply pointed above. Two geographical races ai'e known, one
occupying the Continental range of the species, tlie other confined
to Great Britain.

Rhinolophus ferrum-equinum februm-equinum Schreber.

1774. Vcspertilio fcrntm-cqiiimtm Sclireber, Siiugthiere, i, pi. lxii, upper

figures ; description, i, p. 174 under name : Die Hufeisennase

(part). Prance ; based primarily on Daubeuton.
177G. Vcspertilio equinus P. L. S. Miiller. Natursyst. Suppl. u, Kegist.-

Band, p. 20 (part), France.
1779. [T'esperiiZJo] perspiciUatus Blumenbacb, Handb. d. Naturgescb., p. 75

(part : included the leaf-nosed bats of Europe and Soutli America).
1785. [Vespertilio] ungula Boddaert, Elenchus Auimaliuni, i, p. 71

(Burguud}-).
1792. Vcsp\crfilio] fer[fnni\-equ[inuvi] viajor Kerr, Anim. Kingd., p. 99

(not 1'. molossus major Kerr, I.e., p. 97), Prance.
1798. Vespertilio hippocrcpis Sehrank, Pauna Boica, i, p. G4 (Renaming

of fcrrum-cquinum Schreber).
1803. Bhinolophiis major Geoffroy, Catal. Mamm. ;\Ius. Nat. d'Hist. Nat.,

Paris, p. 50 (Burgundy).
1S13. Bhinolophiis unihastatus (4cofii'oy, Ann. INIns. d'Hist. Nat., Paris,

xs, p. 257 (France).
1829. '! Ehinolophns nnifer Kaup, Entw.-Gcscb. u. Natiirl. Syst. Europ.

Tbierwelt, i, p. 104 (nomen nudum).
1857. Bhinolophus fcrrnm-cquinuni. Blasius, Siiugetliiere Deutschland.^.

p. 31.
18G3. \_Bhinolvphus frrrum-cqiiiniiiiij a var. qcniiaiuciis Kocli, Jalirb. des

Vereins fiir Naturlcunde im Herzogtlium Nassau, xviii, p. 522

(Wiesbaden, Hesson-Nassau, Germany).
18G3. [Blmiolophns fcrriaii-cqiiiniim] B var. italiciis Koch, Jalirl). des

Vereins fiir Naturkunde im Herzogthum Nassau, xviir, p. 523

(Italy).
1878. Bldnolopihiis fvrriiin-cqiiinum Dobson, Catal. Cliiropt. Brit. Mus.,

p. 119 (part).
18S5. Bhinolophus ■unihastatus^ var. hoinorodahnasicnsis Daday, Orvos-

Termeszettudomanyi Erteset(i, Kolozsvar, x, p. 274 (Homorod-

ALmas cave, Hungary).

1886. Bhinolophus nniliastatus var. Jiomorodalmasicnsis Daday, Verhandl.

u. Mittheilungen des Siebenbiirgiseben Vereins fiir Naturwissensch.
in Hermannstadt, xxxvi, p. 79.

1887. Bhinolophus fcrriim-cquinum var. homorodcnsis Daday, EIrtekezesek a

Term^szettudomanyok Kor^bi'd, Budapest, xvi, pt. 7, p. IH
(Renaming of homorodalniasiensis).

RHINOLOPHUS

143

1904. Bliinolophus ferrtiin-equinuvi obscurus Cabrera, ilem. Soc. Espafi.

Hist. Nat. Madrid, ii, p. 257 (Valencia, Spain).

1905. Bhinolophus ferrum-cquinum hjpicus Andersen, Proc. Zool. Soc.

London, 1905, ii, p. 113, October 17, 1905.

1905. Bhinolophus femtfn-equinum obscurus Andersen, Proc. Zool. Soc.
London, 1905, ii, p. 116, October 17, 1905.

1910. BMvolophus feirmn-equinum, B. fcmmi-equinum obscurus and B.
ferrum-equinum homowdensis Trouessart, Faune IManim. d'Europe,
pp. 4-5.

Type loralitji. — Burgundy, France.

Geographical distribution. — Central and southern Continental
Europe.

Diagnosis. — Wing relativel}^ long ; forearm, 54 to 58 mm. ;
third finger, 84 to 92 mm.

Measurements. — Average and extremes of six males from
8ilos, Burgos, Spain: head and body, 62 •! (58-64 '4); tail,
39-9 (37-6-43); tibia, 23-2 (21-8-24); foot, 11-7 (11-13);
forearm, 55-1 (54-8-55-4); third finger, 89-3 (88-92); fifth
finger, 72-5 (72-73); ear from meatus, 24-9 (24—26). Average
and extremes of five females from Silos, Burgos, Spain : head
and body, 61-8 (60-66); tail, 38-8 (37-42); tibia, 23-1
(22-6-24); foot, 11-7 (11-13) ; forearm, 56-2 (56-57) ; third
finger, 89-6 (88-91); fifth finger, 73-4 (71-76); ear from
meatus, 25-1 (24-8-26). Two males from Granada, Spain:
forearm, 54 and 54. Two females from the same locality :
forearm, 56 and 58. Two males from Elche, Alicante, Spain :
forearm, 54 and 55. Average and extremes of six adults
(3 males and 3 females) from St. Genies, Gard, France : tibia,
24-1 (23-4-24-6); foot, 12-1 (11-8-12-8); forearm, 55-4
(54-65) ; third finger, 87-3 (84-91) ; fifth finger, 71-5 (69-74).
Female from Marseilles, France : forearm, 57. Male and female
from near Genoa, Italy : forearm, 57 and 57. Two females from
Kimini, Ital}^ : forearm, 53 and 57. Female from Siena, Italy :
forearm, 54. Male from Rome, Italy : forearm, 56. Male from
the Parnassus region, Greece : forearm, 54. Adult female from
Tubingen, Wiirtemberg, Germany : head and body, 63 ; tail, 37 :
tibia, 24-4; foot, 11; forearm, 56-6; third finger, 88; fifth
finger, 72; ear from meatus, 24; width of ear, 16. Adult
female from Ofener Mts., Hungary : head and body, 61 ; tail, 40 ;
tibia, 24; foot, 11-6; forearm, 56; third finger, 87; fifth
finger, 73; ear from meatus, 23-4; width of ear, 16. Two
males from Herkulesbad, Hungary : forearm, 56 - 6 and 58. Four
females from Herkulesbad, Hungary: forearm, 56, 56-4, 56*6
and 57-4. For cranial measurements see Table, p. 144.

Specimens c.vamincd. — One hundred and forty-one, from the following
localities : —

Portugal: Cintra, 1.

Spain : Silos, Burgos, 22 ; Granada, 4 ; Elche, Alicante, 3 ; San
Cristobal, IMinorca, Balearic Islands, 4.

144

CHIROPTEKA

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ft w

RHINOLOPHUS

145

S .SO

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14G

CHIKOPTERA

Fkaxce : Troubate, Hautes-Pyrenees, 6 ; St. Genios, Gard, 4:3 (ISIottaz) ;
Marseilles, 1 (U.S.N.M.) ; Meounes, Var, 1.

Italy: Turin, 1; near Genoa, 8 (B.M., U.S.N.M. and Genoa); Grotta
de Isoverde, Liguria, 7 (B.M. and Genoa) ; Rimini, 2 (U.S.N.M.) ; Siena, 1
(U.S.N.M.); near Rome, 5 (U.S.N.]\I. and Genoa); no exact locality,!;
Sicily, 2.

Corsica : Commune di Barbaggio, 1.

Sardinia : Sassari, 1.

Greece : Parnassus region, 1 (U.S.N.M.) ; Island of Syra, Cyclades,

1 (U.S.N.M.) ; Mt. Goria Monastery, Crete, 3.

Germany : Tiibingen, Wiirtemberg, 1.

Switzerland : Geneva, G (U.S.N.M. and J\Iottaz) ; Boiidry, Neuchatel,

2 (Mottaz) ; Tremona, Ticino, 1 ; Mendrisio, Ticino, 2 ; Lugano, Ticino, 3.

ACSTRIA-HUNGARY : Herkulesbad, Hungary, G ; Ofener Mts., Hungary, 1.

BeiaarJcs. — Spanish specimens of BhinolopJiuti ferruui-equinum
have been regarded by both Cabrera and Andersen at represent-
ing a pecuhar race, R. f. ohscuius, distinguished by small size.
The material which I have examined, however, indicates that
the Iberian animal cannot be treated as distinct.

6.

Cintra, 500 mm. Portug

;al.

0. Thomas (c A- p).

98. 2. 2. 1.

G al.

Silos, Burgos, Spain.

N. iV- S. Gonzalez

8. 7. 7. 38-4.3.

2 0, 2 V.

Silos, Burgos.

G. S. MiUer (c).

8. 8. 4. 1-4.

6, 9.

Granada, 2250 ft.

G. S. Miller (c).

8. 8. 4. 8-9.

2 i, ?.

Elche, Alicante, 20 m.

G. S. Miller (c).

S. 8. 4. 5-7.

6, -2 V.

San Cristobal, Minorca,
learic Islands.

Ba-

0. Thomas and
R. I. Pocock
(c \- P).

0. 7. 1. 24-26.

6 al.

San Cristobal, ;\Iiuorca.

0. Thomas and
R. I. Pocock
(c & P).

0. 7. L G8.

G c^..

Troubate, Hautes-Pyrenees,

0. Thomas (p).

G. 4. 1. 1-6.

France. [A. Robert.)

1 al.

Meounes, Var.

Dr. K. .Jordan

(c & p).
Prof. Bonelli (p).

8. 3. 15. 1.

V al.

Turin, Italy.

9 al.

Isoverde, Genoa.

0. Thomas (CvV p).

88. 12. 7. 1.

Skeleton.

Italy. (I'riuce Bonaparte.)

Tomes Collection.

7. 1. 1. 727.

2 6 al.

Sicily.

rfal.

Barbaggio, Corsica.

Dr. C. I. Forsyth
Major (c & p).

6. 4. 14. 1.

6a\.

Sassari, Sardinia.

Marquis G. Doria

6. 12. 1. 11.

6, 2 9.

Mt. Goria Monastery, Crete.

Miss D. Bate (c).

5. 12. 2. 1-3.

9al.

Tiibingen, Wiirtemberg, Ger-

Dr. A. Giinther

66. 2. 1. 1.

many.

(P)-

9.

Tremona, Ticino, Switzerland.

0. Thomas (p).

2. 8. 4. 1.

[E. H. Zollikofer.)

2 9.

Mendrisio, Ticino. {E.
Zollikofer.)

H.

0. Thomas (p).

2. 8. 4. 2-3.

3.

Lugano, Ticino. {E. U. Z>
kofer.)

olll-

0. Thomas (p).

4. 4. 5. 1-3.

2 i, 4 9.

Herkulesbad, Hungary.

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

7. 9. 16. 1-6.

9al.

Ofener :Mts., Budapest.

Budapest iluseum

(E).

94. 7. 18. 5.

RHINOLOPHUS

147

JviiixoLOPHUS PERKUM-EQUINUM iNsuLANUs Barrett-Hamiltoii.

I'JJO. J titiiiolophns ferriivi-equimivi insulamts Barrett-Hamilton, Ann. and
Mag. Nat. Hist., 8tli ser., v, p. 292, March, 1910. Type in British
?tluseum.

lyiO. Rlnnoloplius ferriim-cqitiiium ijtsulaiiits Trouessarfc, Faune Mamm.
d'Europe, p. 273.

Ti/jtc locdlifij. -Clieddar, Somersetshire, Englaiul.

(T/'oiiraphiral (lii^frihuiion. — Central and southern England.

DlitfjDOsix. — Wing relatively short ; forearm, 52 to 55 ; third
finger, S.3 to 88.

Mfamireinents. — Type (adult male) : head and body, 67 ; tail,
•■i7 : tibia, 23-4 ; foot, 11 ; forearm, 54 ; third finger, 80 ; fifth
finger, 70; ear from meatus, 23 '6. Average and extremes of
seven males from the type locality : head and body, 65 (63-67) :
tail, 36-9 (35-40): tibia, 22-8 (22-24); foot, 11-2 (11-12):
forearm, 53 '5 (52-54) ; third finger, 85-5 (83-87) : fifth finger,
Gii'9 (69-71) ; ear from meatus, 24*2 (23-25). Tliree females
from the type locality : head and body, 64, G6 and 64 : tail, 35,
34 and 36; tibia, 23-4, 23 and 23; foot, 10-6, 12 and 12:
forearm, 55, 54*6 and 54: third finger, 87, 86 and 88: fifth
finger, 72, 72 and 72 ; ear from meatus, 25, 24 and 24. For
cranial measurements see Table, ]>. 144.

Specimens examined. — Twenty-four, from tlie following localities in
England : — Cheddar, Somersetshire, 10 ; Wells, Somersetshire, 10 ; Dorset-
shire, 3 ; Bonclmrch, Isle of Wight, 1.

5, 9. Cheddar, Somerset, J. A. Coward (c & p). 7. 1. 10. 1-2.

England.

(J al. Cheddar. • -T. A. Coward (c & p). 7. 1. 10. 3-10.

<, 2 9 al. Wells. Hon. N.C. Rothschild 5.1.23.1-3.

3 i. 6 juv., 9 St. Wells. Hon.N. C.Rothschild /I. 9. 3. 1-4.-

(p). \2. 9. 6. 1.

.^,9. Wells. Hon. N.C.Rothschild 11.1.3.3-4.

(P)-
2 9. Dorset. W. M. Hardy (c & p). 11. 1. 3. 1-2.

1. Bonchurch, Isle of Rev.C. A.Burv (c A" p). 11. 1. 3. 388.

Wight.

RHINOLOPHUS HIPPOSIDEROS Beclistein.

(Synouymx under subspecies.)

(icoijraphiral (Uxfrihittioii. — From Uilgit through the Mediter-
ranean sub-region and central Europe to Ireland ; north in
continental Europe to the Baltic, and in Great Britain to about
the southern border of Scotland.

DifKjnosix. — Size small, the forearm less than 43 mm. in
length, condylobasal length of skull about 14 to 15 mm., mandible
about 10 mm. : no.seleaf (fig. 26 b) with bluntly cuneate sella
and low , broadly rounded connecting process ; fourth finger with

L 2

148 CHIROPTERA

first phalanx slightly more than half as long as second ; large
upper premolar separated from canine by a noticeable space
occupied by the well developed small premolar, which lies per-
fectly in the tooth-row.

External characters. — A much smaller, more delicately formed
animal than Uliinoloplms ferni/in-equimim. General outline of
noseleaf nariower than in H. fcrriim-equinum, the width of horse-
shoe less than distance from flat area at base of sella to tip of
lancet ; sella narrowly rounded at tip, the sides straight, slightly
convergent above, the resulting outline bluntly cuneate ; connect-
ing process essentially as in the larger sjjecies but relatively
wider and lower, its upper extremity about on level with that of
sella ; lancet slender, scarcely or not contracted at middle, the
tip cuneate. Ear when laid forward extending about o mm.
beyond extremity of muzzle, the narrow tip abruptly curved
backward ; antitragal lobe more than half as high as conch, its
width less than its height, its upper margin noticeably obli(iue.
Wings and feet essentially as in B. ferruvi-equinavi.

Colour. — The colour does not difl'er appreciably from that of
Rliinoloplivs ferrum-equinum.

Skull. — The skull differs from that of Blii)wlophu.s ferruui-
equinum chiefly in its conspicuously smaller size (greatest length
about 16 mm. instead of about 24 mm.). There are also some
slight peculiarities in form, the principal of which is the greater
contrast between width of brain-case and anterior maxillary
region, the breadth of former being about twice that of latter.
As the breadth of brain-case is contained distinctly less than
2.1 times in greatest length it is probable that the diffei-ence
between the two animals is due rather to enlarged brain-case in
the smaller than to widened palate in the larger. Sagittal crest
slightly develojied ; lambdoid crest obsolete. Nasal region
relatively more inflated than in J?, ferrum-equinum, particularly
at side of nares, but dorsal outline not rising abruptly above
level of interorbital region. Mesopterygoid fossa so conspicu-
ously widened anteriorly that it occupies almost entire extent of
palate between posterior molars, leaving no space for lateral
palatal emarginations, a peculiarity by which the skull may be
distinguished from that of all other European members of the
genus.

Teeth. — Though in general resembling those of Mhiiiohq/hu.s
ferrum-equinum, ajmrt from their much smaller size, the teeth of
B. hipposideros show several notable peculiarities. Upper canine
relatively small, its apex in line with large cusps of molars, anrl
slightly below that of posterior premolar ; lower canine corre-
spondingly .short ; anterior upper jiremolar a well develojjed
functional tooth lying perfectly in tooth-row, with crown area
equal to nearly half that of canine, its shaft subterete though
flattened posteriorly, and only a little less than half as high as
main cusp of large jiremolar : anterior and posterior lower pre-

RHINOLOPIIUS

149

molars less contrasted in size than in the larger animal, and
separated from each other by a slight space in which lies
the minute middle premolar, less
crowded outward than in B. ferruvi-
eqwiniim. Large upper premolai
and upper molars essentially as in
R. ferrum-eqmnum except that in
m^ there is a distinct meta-
style and fourth commissure,
and area of tooth is nearly
equal to that of m"^. Lower
molars as in Hhinolophus
ferrKrn-t'quinum, except that
in M«3, almost exactly resem-
bles the other teeth, the second
triangle having undergone
practically no reduction.

Measurements. — Head and
body about 40 mm., tail about
.'jO mm., forearm, 34-5 to
4rl • 7 mm.,condylobasal length
of skull, 13 "8 to 16 mm.
(greatest length 1 4 • 5 to 1 6 • 2).
Details under sub.species.

Bemarks. — Three imperfectly differentiated forms of Bhino-
lophus hipposideros occur in Europe : a larger central race, a
smaller Mediterranean race, and an intermediate form peculiar
to Great Britain and Ireland. Without examination of much
more complete material than that now available it is impossible
to reach any wholly satisfactory conclusion as to the status and
interrelationships of these forms. The characters here given are
those published by Andersen in 1905.*

Fig. 2S.

ItliiiiolnpliHS Jiipposidems. Teetli X 10

Rhinolophus hipposideros hipposidkros Bechstein

1789. Vespertilio ferrum-equinum B., Die kleine Hufeiseunase, Bechstein.

Gemeinn. Naturgesch. Deutschlands, i, 1st ed., p. 186.
1792. Vcsi}[crtilio] fcrlnimyequlinum^ minor Kerr, Anim. Kingd., p. 99

(not 1". molossun minor Kerr, I.e., p. 97) (France).
1800. Vespertilio hijjposidcros Bechstein, Thomas Pennant's .\llgemeine

tJebersicht der vierfiissigen Thiere, ii, p. 6'29.
1803. lilmiolophus minor Geoffrey, Catal. ]\Iamm. !Mus. Nat. d'Hist. Nat.,

Paris, p. 57 (Neighbourhood of Paris).
181:5. Rhinolophus bihastatus Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xx,

p. 259 (Neighbourhood of Paris).
1816. ? I'hijllorhina mimtia Leach, Syst. Catal. Spec. Indig. Mamm. and

Birds Brit. Mus., p. 5 (uomcn nudum : " Small Leaf nose ").

* For further discussion of the subject see Mottaz, M6ni. Soc. Zool. de
France, Paris, xx, pp. 21-22, September, 1907 ; Andersen, Ann. and ^lag.
Nat. Hist., 7th ser., xx, pp. 384:^89, November, 1907.

150 CHIROPTERA

1S29. ? B]dnolo2)hubi bifer Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ.

Thierwelfc, i, p. 104 (nomeu nudum).
1S57. ? Rhinolophus hipposideros Blasius, Siiugetbiere Deutschlauds, p. 29.
1863. illhinolophus hipposideros] a var. typus Koch, Jahrb. des Vereins fiir

Naturkunde im Herzogthum Nassau, xviii, p. 5.30 (Wiesbaden).
1863. [Rhinolophus hijiposidcros] 0 var. alpinus Koch, Jahrb. des Vereius

fiir Naturkunde im Herzogthum Nassau, xviii, p. 530 (Alps).
1870. niiinoloj^ltus cggcnlioffncr Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Clasae, lxi, Abth. i, p. 151

(MS. synonym of bihastatas).
1878. Rhinolophus hipposideros Dobson, Catal. Chiropt. Brit. Mus., p. 117.

1885. Rhinolophus bihastatus var. kisnyiresicnsis Daday, Orvos-Terme'sz-

ettudoniAnyi Ertesitti, Kolozsvar, x, p. 274 (Kis-Nyires, Szolnok-
Dobaka, Hungary).

1886. Rhinolophus bihastcdus var. kisnyiresiensis Daday, Verhandl. u.

Mittheilungen des Siebenbiirgischen Vereins fiir Naturwisseusch.
in Hermannstadt, xxxvi, p. 80.

1887. Rhinolophus hipposideros var. troglophilus Daday, Ertekezesek a

Term6szettudomdnyok Korebol, Budapest, xvi, pt. 7, p. 8 (Re-
naming of kisnyircsiensis) .

1904. Rhuwlophus euryalc helvetica Bretscher, Vierteljahrsschrift der

Naturforsch. Gesellsch. in Ziirich, xlix, p. 256. See Mottaz,
Bull. Soc. Zool., Geneve, i., p. 172, 1908 (Baar, Zug, Switzerland).

1905. Rhinolophus liipposidenis ti/picns Andersen, Proc. Zool. Soc. London,

1905, II, p. 141, October 17, 1905.
1910. Rliinolophiis Idpposidcrns Trouessart, Paune ]Mamm. d'Europe, p. 9.

Tyije localitij. — France.

Oev'jrajjhical distribution. — Central Europe, north of the Alps,
ea.st through Ai'uienia and north-west Persia to the Himalayas.

Diagnosis. — Forearm* usually 39 to 41 '7 mm.; greatest
length of skuUf about 16 mm.

Mcasnrevients. — Two males from 8trass near Burgheim,
Bavaria, Germany: tibia, 17 and 18*4; foot, 7 '8 and 7*4:
forearm, 38-8 and 39-2 ; third finger, 61 and 60 ; fifth finger,
54 and 53. Female from the same locality: tibia, 17'4 ; foot,
7-4; forearm, 38-8; third finger, 60: fifth finger, 53. The
six following extremes of forearms of Eurojuean specimens are
given by Andersen (I.e. p. 142) : N. Bulgaria (1), 39 ; Roumania
(13), 39 to 41-2 ; Transsjdvania (2), 40 to 41 ; S. Carj^athians
(1), 39-3; Schlangenbad, Nassau, Germany (2), 40 to 40*1 :
Strassburg, Germany (3), 39 to 40*1. Forty-nine males from
the vicinity of Geneva, Switzerland (Mottaz collection), J ."7 -3
to 39. Thirty females from the same locality (Mottaz collection), J
38-5 to 40-6. Four males from the same locality (U.S.N". M.),
37 to 38-4 Six females from the same locality (U.S.N.M.),
37*4 to 40. Adult female from Dions, Gard, France (Mottaz
collection), 40. For cranial measurements see T\'ible, p. 152.

* Bases of metacarpals included.
t From back of occiput to front of canine.

% Measured by Chas. Mottaz (base of metacarpals included) and verified
by Gerrit S. Miller.

RHINOLOPHUS

1.11

Specimens examined. — One huudred and seventeen, from the following
localities : —

France: Dions, Gard, 1 (Mottaz) ; St. Cergues, Haute-Savoie, 1.

Germany : Strass, near Burgheim, Bavaria, 3 ; Mainz, 1 (Strassliurg) ;
Bitsch, Alsace, 1 (Strassburg).

Austria-Hungary : Hatszeg, Hiinyad, Hungary, 1.

Switzerland: Near Geneva, 9-i (B.M., U.S.N.M. and Mottaz); St.
Jloritz, 1 ; Thayngen, Schaffhausen, 2 (U.S.N.M.) ; Roggwil, Thurgau,
7 (B.:M. and U.S.N.M.) ; Canton Thurgau, 5 (B.M. and U.S.N.M.).

6. St. Cergues, Haute-Savoie, A.Robert (c & p). 5.4.9.1.

France.

■2 ?. Burgheim, Bavaria, 375 m. Lord Lilford (p). 11. 1. 1. 117-

Germanv. 118.

c5. Hatszeg, Hunyad, Tran- C.G.Danford (c). 8.11.8.1.

sylvania.

3 6. Grand Pr^, Geneva, Swit- C. jMottaz (c & r). 6. 2. C. 1-3.

1. St. Moritz, Grisous. Leon O. Galliard 75. 9. 20. 3.

(c X- P).
3 i, 2 ?, iram. Roggwil, Thurgau. (E. H. O. Thomas (p). 2. 8. 4. 4-9.
Zollikofer.)
9al. Ihurg&u. {E.H. Zollikofer.) 0. Thomas (p). 2.8.4.55.

RiiixoLOFiius HiPPOSiDERCs MINIMUS Heuglin.

1861. Rhinolophus minimus Heuglin, Nov. Act. Acad. Caes. Leop. -Carol.,
XXIX, Abliandl. viii, p. 6 (articles separately paged) (Ker^n,
Abyssinia).

18G'B.' [RhinoloiJhus hipiM.ndcro:^] y y&r. piallid-us Koch, Jahrb. des Vereins
fiir Naturkunde im Herzogthum Nassau, xviii, p. 531 (Mediter-
ranean region).

1904. Bhinolophus phasma Cabrera, Mem. Soc. Espan. Hist. Nat., Madrid,
II, p. 252 (Madrid, Spain).

1904. Tili[inolophus] h[ippiosidcnis] minimus .\ndersen, Ann. and ^lag. Nat.

Hist., 7th ser., xiv, p. 456, December, 1904.

1905. Rhinolophus hij^posidcrits minimus .\ndersen, Proc. Zool. Soc.

London, 1905, ii, p. 140, October 17, 1905.
1910. RJiinolophus hippiosidci'us minimus Trouessart, Faune Maram.
d'Europe, p. 10.

Ti/jje lornliiij. — Keren, Abyssinia.

Geofjraphical distribution. — Mediterranean region.

Diagnosis. — Forearm usually 34 ■ 7 to .'58 mm. ; greatest
length of skull, 14-5 to 15 '5 mm.

Measurements. — The following measurements of forearms are
given by Andersen (I.e. p. 141) : Keren (type), 86' 3 ; Cvprus (6),
34-7 to 37-7 ; Malta (8), 36 to 37 ; Ostia, Italy (2)," 35-7 t«
36-8; Corsica (1), 37-7; Balearic Islands (7), 36-2 to 37-6;
Seville, Spain (1), 37*7 ; Cintra, Portugal (1), 36 -2. Specimens
in U.S. National Museum : Ficuzza, Sicily ( ? ), 35 • 2 ; Rome ( ^ ),
36-4; Siena (3 $), 36-8 to 38; Marseilles, France ( ? ), 38;
Dions, Gard, France (cJ), 34 '8, (9), 36-6 Two females from
Ax-les-Thermes, Ariege, France, 36-6 and 38*4. Two males
from Silos, Burgos, Spain, 36 and 37. Female from the same
locality, 38. Five males from Ticino, Switzerland, 37 '0
(36-2-38). Mottaz collection: St. (denies, Gard, France (15,

152

CHIROPTERA

-5 '^

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CHIROPTERA

Loth sexes), 36 "4 to .SS-2) ; Sardinia (2 9 ), 37-4 to 37 -S. For
cranial measurements see Table, p. 153.

Specimens cxaminvd. — Eighty-one, from the following localities : —

Spain: Inca, Majorca, Balearic Islands. 1; San Cristobal, Minorca,
Balearic Islands, 3; Elche, Alicante, "1; Silos, Burgos, 5; Seville, 1.

Portugal: Ciutra, 1.

Fbance: As-les-Thermes, Ariegc, 6; St. Geuies, Gard, 17 (U.S.N.M.
and Mottaz); Marseilles, 1 (U.S.N.M.).

Switzerland : Locarno, Ticino, 4 (U.S.N.M.) ; Gordola, Ticino, 2
(U.S.N.M.) ; Minusio, Ticino, 6 (U.S.N.M.).

Italy : Liguria, :3 (U.S.N.M.) ; Siena, 3 (U.S.N.M.) ; Faenza, 1
(U.S.N.M.); Rome, 3; Osiia, Rome, 2; Ficuzza, Sicily, 6.

Sardinia : No exact locality, 2 (!Mottaz) ; Zimmigas, Siliqua, 2.

Corsica : Dintorni di Patrinionio, 3; no exact locality, 1.

^Ialta : 7.

?. Inca, Majorca, Balearic 0. Thomas and R. I. 0. 7. 1. 1-2.
Islands. Pocock (c & p).

i, 9. San Cristobal. Minorca. O. Thomas and R. I. 0. 7. 1. 27-28.

Pocock (c iV' p).

1 al. San Cristobal, i\Iinorca. 0. Thomas and R. I. 0. 7. 1. G9.

Pocock (c & p).

?. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 11.

c!. Elche, Alicante, Spain. G. S. Miller (c). S. 8. 4. 12.

5 al. Cintra, 500 m. Portugal. O. Thomas (c & p). 98. 2. 2. 57.
S,9. .4x-les-Thermes, Ariege, CV. S. Miller (c). 8.8.4.124-

2400 ft. France. 126.

6. Rome, 37 m. {C. Colt.) G.Barrett-Hamilton 11.1.2.42.

(P)-
2. Ostia, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 6-6.

6 9. Ficuzza, Sicily. (A. J!ol>crt.) O. Thomas (p). |g- g" f l^'^^^'

6, ? al. Zimmigas, Siliqua, Sardinia. Marquis G. Doria (p). 6. 12. 1. 20-21.

(7?. Meloni.)
3 6, 2 ?. Malta. {J. Micalleff.) Lord LiKord (p). 11. 1. 1. 119-

123.
2. :\[aUa. Lord Lilford (p). 95. 3. 2. 1-2.

Rhinolopiius hipposidekos MixuTus Montagu.

1808. Vespcrtilio niiniitus Montagu, Trans. Linn. Soc. London, ix, p. 163.
1905. Rhinolophiis liipposidcrus minutus Andersen, Proc. Zool. Soc.

London, 1905, ii, p. 142, October 17, 1905.
1910. lilnnokphits liipposidcruf! ■mhnitus Trouessart, Fauna Mamm.

d'Europe, p. 10.

Type locality. — Wiltshire, England.

Geographical iJitftrihution. — England and Ireland.

Diagnosis. — Forearm, 36 '3 to 39 mm.; greatest length of
.skull about 16 mm.

Measuronents. — Forearm in 30 Engli.sh and Irish specimens
measured by Andersen {i.e. p. 142), 37 • 6 (36-3 to 39). For
cranial measurements see Table, p. 152.

Specimens examined. — Sixteen, from the following localities in England :
— Great Grimsby, Lincolnshire, 1 ; Bowdou, Cheshire, 1 ; St. Asaph,
Denbighshire, 3 ; Conway, Carnarvonshire, 1 ; Hope End, Herefordshire, 1 ;
Wells, Somersetshire, 1 ; Devizes, Wiltshire, 1 ; Zeals, Wiltshire, 2 ; Devon-
shire, 2 ; Ragley House, Warwickshire, 3.

RHINOLOPHUS

1 55

3 6. St. Asaph, Denbighsliire, Charles 0]dham (c .^- i'). 11.1.3.5-7.
Wales.

1. Conway, Carnarvonshire. Sir W. Jardine (c Ar p). GO. 9. 17. 1.

9. Great Grimsby, Lincoln- G. Barrett-Hamilton (p). U. 1. 2. 'J8.
shire, England. (Caton
Uaigh.)

6. Bowdon, Cheshire. {T.A. G. Barrett-Hamilton (i-). 11.1.2.99.
Coioard.)

6 al. Hope End, Herefordshire. N. C. Hewitt (c .t p).

6 al. Wells, Somerset. S. Lewis (c .fc p). 5. 1. 24. 1.

1. Devizes, Wiltshire. J. E. Hartins (c & p). 87.2.21.1.

i, 9 al. Zeals, Wiltshire. F. Norgate (c & p). 4. 11. 6. 1-2.

2 9. Devonshire. Oxley Grabham (c >t p). 11. 1. 3. 8-9.

3. Ragley House, Warwick- Tomes Collection. 7. 1. 1. 289-
shire. 291.

RHINOLOPHUS EURYALE Basius.

1853. PMnoloplms euryale Blasius, Wiegmann's Archiv fiir Naturgesch.,

1853, I, p. 49 (Milan, Italy).
1857. Bhinolophus euryale Blasius, Siiugethiere Deutschlands, p. 35.
1878. Bhinolophus euryale Dobson, Catal. Chiropt. Brit. IMus., p. 116.
1904. E{uryalus] toscanus Andersen and Matschie, Sitz.-Ber. Gesollsch.

Naturforsch. Preunde, Berlin, p. 77 (Caverna di Parigiiano, lit.

Pisani, Italy).
1904. E[uryalus] atlanticus Andersen and Matschie, Sitz.-Ber. Gesellsch.

Naturforsch. Freunde, Berlin, p. 77 (St. Paterne, ludre-et-Loire,

France) .
1904. E[nryalus] cabrene Andersen and Matschie, Sitz.-Ber. Gesellsch.

Naturforsch. Freunde, Berlin, p. 78 (Alcah'i de Henares, Madrid,

Spain).
1910. Rhinoloph-us euryale, R. euryale atlanticus, R. euryale toscanus,

R. euryale cabrerai Trouessart, Faune I\Iamm. d'Europe, pp. 5-7.

Type Jocalitij. — Milan, Italy.

Geixjraphkal distribution. — Southern Europe from Portugal
to Greece, north to Hungary and central France.

Diagnosis. — Size medium, forearm, 44 "6 to 49, condylobasal
length of skull, 16 '4 to 18, mandible, 12 to 1-3, upper tooth-row
6 • 2 to 6 • 6 ; noseleaf with parallel-sided, bluntly rounded sella
and high, sharply pointed connecting process, the lancet gradually
narrowing to a bluntly cuneate tip ; fourth finger with first
lihalanx slightly more than one-third as long as second (ratio
about 38) ; large upper premolar separated from canine by a
narrow space occupied by the much reduced small premolar.

External cliaractera. — Size intermediate between that of the
Greater and Lesser Horseshoes. General outline of no.seleaf
(fig. 26 c) about as in lihinolophus ferrum-equintoii ; sella parallel
sided, rounded ofi" above, connecting pi'ocess .sharply linear-
pointc^d, rising conspicuously above .sella ; lancet with slight con-
striction above middle, beyond wliich the tip narrows gradually to
a bluntl\- cuneate point. Ear when laid forward extending about
5 mm. beyond extremity of nuizzle, its tip less attenuate and le.ss
noticeably curved backward than in lihinolophii.'^ //■rrum-eqiiinuui

156

C'HIROPTEi;A

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RHINOLOPHUS

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CHIROrXERA

and I\. Jiipposidenis ; antitragal lobe about lialf as High as conch,
its width about equal to its height. Wing pecuUar in the
shortening of the tirst jshalanx of fourth finger to a little more
than one-third that of second (average of 1 0 specimens from Gard,
Fi'ance : tirst phalanx, 6'G mm.; second phalanx, 17 '4 mm.;
ratio of first to second, 38 -f )• -Foot averaging slightly more
than half as long as tibia.

Colour. — While essentially as in Illiinoloijlins ferrmii-eqmnuni
and M. hipposideros, the colour usually differs slightly in the more
evident contrast of the light area between ears and the more
drabby general effect of underparts. Median region below
occasionally rather paler than usual, sometimes nearly as in
R. mehehj).

Skull. — In both size (greatest length about 19 mm.) and form
the skull is somewhat intermediate between that of mdywlophun
ferrum-equinum and B. hipposideros. It resembles or surpasses
the latter in the breadth of brain-case relatively to naTro^\"
maxillar}' region, but more nearly agrees with the former in the
nearly parallel-sided mesopterygoid space, the anterior boi'der of
which is separated from posterior molars by well defined palatal
eraarginations. The mesopterygoid .space is, however, shorter in
proportion to its width than in B. ferrum-equinum. Floor of
brain-case between cochlese less narrowed than in the preceding
species. Nasal region slightly less inflated than in R. hipposideros,
and inflated area relatively shorter, its posterior border rising
abruptly above interorbital level.

Tectli. — In all respects the teeth closely resemble those of
Blii iiolophu8 ferrum-equinum (apart from their smaller size), excejjt
that the upper canine is relatively less robust, the anterior upper
premolar is less reduced (its crown area about double that of
uj:)per incisor), and anterior lower premolar is less crowded
between canine and posterior premolar.

Measurements. — For cranial and external measurements see
Tables, pp. 156 and 160.

Specimens e.vamined. — About 130, from the following localities : —

Portugal : Cintra, 6.

Spain: Villalba, Lugo, 1; ^Madrid, 2; Silos, Burgos, 1.

Feance : St. Paterue, Indre-et-Loire, 3 (B.M . and U.S.N.M.) ; St. Genies,
Gard, about 50 (Mottaz) ; Gapeau Kiver, Var, 12.

Italy: Near Genoa, 33 (B.M., U.S.N.M., Genoa, and Mottaz); Monte
Pisanino, 2; Siena, 3 (U.S.N.M.) ; Rome, 2 ; Velletri, Home, 5 (U.S.N.M.) ;
Nicotera, Calabria, 1 ; Marsala, Sicily, 2.

Sardiijia: Mount Gennargentu, 3 (U.S.N.]M.).

Austria-Hungary: Ofener Mountains, 2 ; Orsova, 1.

Dalmatia : Zara, 1.

Greece: Missolungi, Acarnania, 2 (U.S.X.M.).

JteiiiarJcs. — Mhinolopihus euryale is so readily distinguishable
from all the other European membsrs of the genus, except
R. mehehji, as to renuire no special comparisons. From R. meheli/i
it is most easilv distinguished bv tlie form of the lancet and

RHINOLOPHUS

159

aiititragus, together witli its rather smaller size and iisnall\-
darker colour. With the material at hand I am unable to
recognize the local forms of this species described by Andersen
and Matschie, as the alleged differences appear tf) be within the
range of normal indi^'idual variation.

2 9. Cintra, 500m. Portugal. (). Thomas (c & p). US. 2. 2. 2-8.

2 i, 2 9 al. Cintra, 500 m. 0. Thomas (c & r). >JS. 2. 2. 53-5G.

9 al. Villalba, Lugo, N.W. Dr. V. L. Seoane (p). 'J4. 1. 1. 1.
Spain.

5, 9 al. Madrid. A. Cabrera (p). 5. 2. 3. 1-2.

1 al. St. Paterne, Indre-et- Roj^al Army Medical "J. 1. 4. 9.

Loire, France. College (p).

9 al. St. Paterne, Indre-et- (t. E. Dobson (p). 80. 12. 14. 3.

Loire.

'J al. Cxapeau River, Var. Dr. K. Jordan (C & p). 8. 3. 15. 2-10;

3 i, 9 al. Finalborgo, L i g u r i a , Marquis G. Doria (p). 6. 12. 1. 14-17.

Italy. (A. Gagi'iu.)

2 al. Monte Pisanino,Lignria. Lord Lilford (p). 73. 1. 8. (i.

•2 6. Rome. {C. CoIi.) G. Barrett-Hamilton 11. 1. 2. 40-41.

W-

c$ al. Nicotera, Calabria. Florence Museum (b). 85.7.6.1.

2 d. :\Iarsala, Sicily. {A. O. Thomas (p). G. 8. 4. 10-11.
Robert.)

6, 9 al. Ofeuer Mts., Budapest. Budapest Museum (e). 94. 7. IS. 2-3.
9. Orsova, Hungary. Hon. W. Rothschild 7. 9. 16. 7.

9. Zara, Dalmatia, 50 ni. Lord Lilford (p). 11. 1. 1. 128.

{Kolombafocic.)
1 al. S. Europe. Purchased (Parreys). 47. 5. 27. 44.

RHINOLOPHUS MEHELYI Matschie.

1901. RhinolopliHS mehehji ^Matschie, Sitz.-Ber. Gesellsch. Naturforscli.

Freunde, Berlin, p. 225 (Bucharest, Roumauiat.
1904. Rhinolophufi carpetcuiua Cabrera, ]\Iem. Soc. Espafi. Hist. Nat., ii,

p. 254 (Madrid, Spain).
1910. Rhinolopliua euryalc vwlieliji and R. cavpctanus Trouessart, Fauue

5Iamm. d'Europc, pp. 7-8.

Type locaHty. — Bucharest, Uoumauia.

(Teix/rapJucal flistribidioii. — Roumania, southern France
(Uard), Sardinia, central Spain. Details of distribution not
known.

Diagnosis. — Like Hhiiiolophns luryaJr but larger (forearm,
48 'G to 51 "4; upper tooth-row about 7 mm.); noseleaf with
lancet abruptly narrowed to a linear tip ; ear with antitragal
lobe relativel)^ broad and low ; fourth linger with iirst phalan.\
decidedly more than one-third as long as second (ratio about 44) :
colour usually paler than in tlie related animal.

J E.rirrtial characters. —i>\ight\y larger and more robu.st than
Rhinoloplnis ciiryalc, a difference especially noticeable in freshlv
killed individuals. Noseleaf as in I{. ciiryale, except tluit the
lancet is very abruptly narrowed above middle to a distinctlv
linear tip. Ear as in B. enryulr but bioader. the antitragal lobe

160

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162 CHIROPTERA

scarcely half as high as conch, its width sHghtly greater than
height. Wing difl'ering from that of the related animal in the
less degree of shortening of the first phalanx of fourth linger as
compared with second (average of ten specimens from Gard,
France, first phalanx, 8'1 ; second phalanx, 18'0; ratio of first
to second, 44 +). Foot as in B. euryale.

Colour. — Though not invariably distinguishable the colour is
usually paler than that of Bhinolophus euryale, a difference
especially noticeable in the region between ears, on sides of face
and neck, on chin and throat, and along median portion of chest
and belly, all of which are frequently a very pale almost whitish
drab-gre}'.*

Skull and teetJi. — Except for its slightly greater average size
(greatest length about 20 mm.) the skull agrees with that of
Bhinolophus euryale. Teeth more robust than those of the
related animal, but not peculiar in form.

Measurements. — For external and cranial measurements see
Tables, pp. 161, 163.

Specimens examined. — About fifty-five, from the following localities : —

Spain: Near Madrid, 1 (paratype of carpetcums).

France ; Near St. Genies, Gard, about 50, skins and in flesh (!Mottaz).

Sardinia : Sassari, 3 (B.M. and U.S.N.M.).

RouMANiA ; Bucharest, 1 (U.S.N.M.) ; Dobrudscha, 1 (Mottaz).

Remarks. — At first sight this species apjjears very similar to
Bhinolophus euryale, together with which it occurs ; but its
characters when once understood are readily appreciable. The
ranges of the two animals will probably be found to be essentially
coincident, though Bhinoloplius mehelyl may prove to be more
strictly confined to the Mediterranean region than the smaller
form.

9. Madrid. A. Cabrera (p.) 6. 2. 3. 1.

(Paratype of R. carpetayius Cabrera.)
2 i al. Sassari, Sardinia. Marquis G. Doria (p). 6. 12. 1. 18-19.

RHINOLOPHUS BLASII Peters.

1857. Bhiiiolopihus cliuosiis Blasius, Saugethiere Deutschlands, p. 33. Not
of RiippeU, 1824 (Italy, Sicily, Istria and Dalmatia).

1866. Bhinoloplius blasii Peters, Monatsber. k. Akad. Wissensch. Berlin,
p. 17 (Renaming of clivosus Blasius).

1878. Rhinolophus blasii Dobson, Catal. Chiropt. Brit. Mus., p. 117.

1910. Bhiiwlophus blasiusi Trouessart, Faune Manim. d'Europe, p. 9.

Type locality. — South-eastern Europe.

Geographical distrihution. — Eastern portion of the Mediter-
ranean region : Cyprus, Greece, Italy?

Diagnosis. — Size essentially as in Bliinolojjlnis euryale; nose-

* When seen by candle-light flying in caverns these bats are said to
appear entirely white.

RHINOLOPHUS

163

Observations.

Teeth not worn.

„ slightly worn.
,, moderately worn.
,, not worn.

„ slightly worn.
,, not worn.

,, slightly worn.
„ not worn.

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• 6. 12. 1. 18
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86536
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37508

Locality.

R. mehelyi.

lloumauia: Dobrudscha .

Bucharest
Sardinia : Sassari .

France : near Nimes, Gard

Spain : near ^ladrid

R. blasii.
Cyprus ....

Greece: Nauplia

M 2

1G+ CHIROPTERA

leaf with cuneate sella and very bigli, sharply poijited connecting
process ; fourth fingei' with iirst phalanx more than half as long
as second ; no marked contrast between crown areas of anterior
and posterior lower* premolars, a character unique among the
European members of the genus.

Colour. — The only skin of this species which I have examined
is in bad condition. It indicates that the colour is not essentially
different from that of Bhinolophus euryale.

Skull. — In general the skull resembles that of RhlnolophiiH
euryale, with which it agrees in size and in the form of the nasal
swellings as well as in that of mesopterygoid fossa and posterior
portion of palate. Constriction at front of interparietal more
pronounced than in any of the other European species, noticeably
marking off the occipital region from rest of brain-case.

Teeth. — Incisors, canines and molars as in Mhinoloplms euryale.
Small upper premolar slightly less reduced, perfectly in the
tooth-row, but showing no tendency to develoj) a cusp. Large
upper premolar with anterior and posterior margins of crown
essentially parallel, the posterior border nearly straight. Lower
premolars differing from those of all the other European membei's
of the genus in the approximately equal crown ai'eas of the two
larger teeth, the anterior subterete, the posterior with trapezi-
form section ; shaft of posterior tooth with diameter in axis of
tootli-i'ow much less than transverse diametei', the cusp when
viewed from the side appearing to rise from middle of crown with
noticeable flat area before and behind it.

Meanurements. — Average and extremes of four females from
Cyprus: head and body, 48-4 (44-51); tail, 24-7 (24-25);
tibia, 19-3 (19-20) : foot, 9-7 (9-4-10) ; forearm, 45 -^ (44-6-
47) ; thumb, 7-5 (7-8) ; third tinger, 70 3 (69-72) ; tifth finger,
57 "7 (56—60) ; ear from meatus, 19 "7 (19-20) : ear fi-om crown,
15-5 (15-16-4); width of ear, 14-7 (14-15). Adult from
Nauplia, Greece : tibia, 18 - N ; foot, 9 • 6 : forearm, 44 • 6 ; thumb,
8 ; third finger, 09 : fifth finger, 57. For cranial measurements
see Table, p. 163.

Specimens exaviincd. — Five, from the following localities : —
Cypbus : No exact locality, 4.
Geeece : Nauplia, 1 (U.S.N.:^[.).

Meuiurls. — This .species is so readily distinguished from the
other European members of the genus by the peculiarities of its
noseleaf and lower premolar as to reejuire no special comparisons.
Its range appears to be strictly confined to the eastern portion
of the Meditei-ranean i-egion, not extending west of Italy.

vespertilionin.e 165

Family VITSPERTILIONID^.

1821. Vespertilionichx Oray, London IMecl. Repos., xv, p. 299, April 1, 1821

(part).
1857. Vespertiiiones Blasius, Siiugethiere Deutschlands, p. .37.
1878. Vcspertilionidx Dobson, Catal. Chiropt. Brit. ]\Ius., p. 1G7 (except

the genera Natalus and Thyroptera).
1907. Vespertilionidx Miller, Families and Genera of Bats, p. 195, -Tune 29,

1907.

Geographical distribution. — Eastern and western hemispheres
to the limits of tree growth ; in the Atlantic to the Azores, and
in the Pacitic to the Galapagos and Hawaiian Islands from
America, and to Australia, New Zealand and Samoa from Asia.

Characters. — Ear with tragus ; muzzle without distinct leaf-
like outgrowths ; skull with premaxillaries represented b}' nasal
branches only, the two bones very earl\' fused with surrounding
parts ; median length of palate greater than least distance between
tooth-rows ; auditory bulla not emarginated on inner side ; shoulder
girdle normal, without fusion of its elements : secondary
articulation of humerus with scapula better developed than in
the Rhiiiolvphidse ; fibula very slender, not adding appreciably to
strength of leg ; foot normal, the toes slender ; tail not project-
ing conspicuously beyond membrane.

Memarks. — This family is the most widely distributed group
of bats as well as one of the richest in genera and species. Forty-
one genera are at present known, eight of which occur in
Europe.*

Sub-Family VESPERTILIONIN^.

1878. Vcspertilioius Dobson, Catal. Chiropt. Brit. IMus., p. 168 (except

genera Kcrivoula and Harpiocephalus) .
1907. Vespertilioninx Miller, Families and Genera of Bats, p. 197, June 29,

1907.

Oeof/rapliicdl distrihniion. — Same as that of the family
Vespertilionidse.

Characters. — Sternum slender, its entire length considerably
more than twice greatest width of presternum ; median lobe
very much smaller than body of pre.sternum : six ribs connected
with sternum ; seventh cervical vertebra not fused with first

* The American Nycteris cinerea has been recorded (under the name
Vespertilio pruinosiis) from South Ronaldshay, Orkney Islands, but the
occurrence seems open to question (see Wolley, The Zoologist, vii, p. 2343,
1849; Yiii, pp. 2G95-96, 2813-14, 1850; Barrett-Hamilton, Hist. Brit.
Mamm., i, pp. 222-224, March, 1911).

A specimen of another North American member of this genus
(N. borealis), bearing the label: " Villeveque (Maine & Loire), 8. 1. 89," is
hgured by Trouessart in Bull. Soc. Zool. de France, xxx, p. 152, 1905.
This is copied, with change of locality from Maryland to France, from a
figure published in Bull. U.S. Nat. Mus., No. 39, pt. N, 1899 and 1901.
Though intended merely as a guide in preparing specimens, Dr. Trouessart's
publication might be misinterpreted as a French record of the species.

166 CHIROPTERA

dorsal ; scapula with coracoid curved outward ; nostrils simple ;
lower incisors in all known genera, 3-3.

BemarJcs. — The sub-family Vespertilioninse contains all but
eight of the known genera of VespertUionidse, and all but one,
Miniopferus, of the eight found in Europe. It is the central,
least specialized portion of the family.

KEY TO THE EUROPEAN GENERA OP VESPEBTILIONID^.

Cheek teeth ~. Mijotis, p. 166.

Cheek teeth less than Jp^.
Upper premolars 1-1,

Rostrum noticeably concave on each side of middle
line ; nares extending about halfway to inter-
orbital constriction ; palatal emargination

broader than deep Vcspcrtilio, p. 238.

Rostrum evenly convex laterally ; nares not ex-
tending halfway to interorbital constriction;

palatal emargination deeper than broad Eptesicus, p. 224.

Upper premolars 2-2.

Lower premolars 3-3.

Auditory bulla large, its greatest diameter
more than twice width of basioccipital ;
ear much longer than head ; second
phalanx of third finger shorter than first Plecotus, p. 256.
Auditory bulla small, its greatest diameter
about equal to width of basioccipital ; ear
shorter than head ; second phalanx of
third finger nearly three times as long as

first Miniopterus, p. 268.

Lower premolars 2-2.

Fifth finger shortened, its length only a little
more than that of metacarpal of fourth

or third Nyctalus, p. 242.

Fifth finger normal, its length greater than
that of metacarpal and first phalanx of
fourth or third.
Upper surface of rostrum convex ; ears

separate Pipistrcllns, p. 202.

Upper surface of rostrum concave ; ears

joined Barbastclla, p. 263.

Genus MYOTIS Kaup.

1829. Myotis Kaup, Entw.-Gesch. u. Natixrl. Syst. Europ. Thierwelt, i,
p. 106 (ynyotis).

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

p. 108 (bcchsteinii).

1830. Lcuconoc Boie, Isis, p. 256 {datthcntonii) .

1839. Vespertilio Keyserling and Blasius, Wiegmaun's Archiv fiir Natur-
gesch., 1839, i, p. 306 (Not Vespertilio Linnfeus, 1758).

1841. Sclijsius Bonaparte, Iconogr. Faun. Ital. i., Introd. alia Classe
Mamm., p. 3 {m7jslaciniis).

1841. Capaccinius Bonaparte, Iconogr., Fauna Ital., i, Indice Distrib.,

p. 1 {capaccinii).

1842. Trilatitus Gray, Ann. and Mag. Nat. Hist., x, p. 258, December,

1842 (hasseltii, macellus = adversus and blcpotis).

MYOTIS 167

1849. Tralatitus Gervais, Diet. Univ. d'Hist. Nat., xiii, p. 213 (Modifica-
tion of Trilatitus).

1856. Brachijotus Koleuati, Allgem. deutsch. Naturhist. Zeitung, Dresden,
neue Folge, ii, p. 131 {nujstacimis, daubentonii, and dasycnemc).
Not Brachijotus Gould, 1837.

1856. Isotus Kolenati, Allgem. deutsch. Naturhist. Zeitung, Dresden, neue

Folge, II, p. 131 {nattciTri and cmargiiiatiis).

1857. Vespertilio Blasius, Siiugethiere Deutschlands, p. 78. Not Vespertilio

Linnaeus, 1758.

1866. Tralatitius Gray, Ann. and Mag. Nat. Hist., 3rd ser., xvii, p. 90,

February 1866 (Modification of Trilatitus).

1867. PtcrnoiAerus Peters, Monatsber. k. preuss. Akad. Wissensch. Berlin,

p. 706 (sub-genus of Vespertilio = Mijotis, type lobipes = muricola).
1870. Exochurus Fitziuger, Sitzungsber. kais. Akad. Wissensch. Wien,

Math.-Naturwiss. Classe, lxii, p. 75 (macrodactyius, horsfieldii =

adversus &nd macrotarsus).
1870. Aeoresies Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien,

Math.-Naturwiss. Classe, lxii, p. 427 (villosissimus, albescens, and

nigricans).
1870. Comastes Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien, Math.-
Naturwiss. Classe, LXII, p. 565 (capaccinii, incgapodius, dasycneme,

and limnophilus).
1878. Vespertilio Dobson, Catal. Chiropt. Brit. Mus., p. 284. Not Vespertilio

Linnaeus, 1758.
1897. Myotis Miller, Ann. aud Mag. Nat. Hist., 6th ser., xx, p. 382,

October, 1897.
1899. Euvespertilio Acloque, Faune de France, Mammif^res, p. 38 {emargin-

atus, mystacinus, nmrinus = myotis, nattereri, and bechsteinii) .
1907. Myotis Miller, Families aud Genera of Bats, p. 201, June 29, 1907.

Type species. — Vespertilio myotis Borkhausen.

Oeoiirapliical distribution. — Entire mainland of Eastern and
Western hemispheres to limits of tree growth ; also the Malay
Archipelago, New Guinea, Australia and Samoa, and in America
the Lesser Antilles.

Characters. — Dental formula : i ||, c J^, pm ^, m ;J-^ = 38.
General form slender and delicate, even in such large species as
M. myotis, the skull slender and lightly built ; muzzle narrow ;
ear narrow and rather long, without special peculiarities of form,
the tragus at least half as high as conch, straight or slightly
curved, tapering gradually to a narrow or acute point.

Bemarks. — The genus 3[!iotis is the most widely distributed
of the genera of bats. It is also probably the richest in species,
though these are at present so imperfectly known that no estimate
of their number can be made. Nine occur in Europe. These
present considerable differences in size, ranging from nearly the
smallest to nearly the largest members of the group ; they also
differ considerably among themselves in certain details of
structure, notably in the relative size of the hind foot ; but all
are recognizable, apart from their dental formula, by a certain
slenderness and delicacy of form, especially noticeable in the
muzzle, ear, tragus and skull.

168 CHIROPTERA

IvEY TO THE EUROPEAN SPECIES OF MYOTIS.

Size large (forearm 53 to 64 mm., condylobasal
length of skull 18-6 to 23-6 mm., upper tooth-
row 8'2 to 10"6 mm.) ; middle upper premolar
normally crowded inward from axis of tooth-
row ; third lower molar with second triangle
much smaller than first and noticeably different
from it in form.
Condylobasal length of skull 22 to 23 '6 mm.;
mandible 17 '8 to 19 mm. ; maxillary tooth-
row 9-8 to 10-6 mm. (Central and southern

Europe) M. mijotis, p. 192.

Condylobasal length of skull 18 '6 to 21-4: mm. ;
mandible 15 '2 to 17 "2 mm. ; maxillary tooth -

row 8-2 to 9'4 mm. (Mediterranean region) M. oxijgiuitlius, p. 199.
Size small or medium (forearm 34 to 47 mm., condy-
lobasal length of skull 12-4 to 16 '8 mm., upper
tooth-row 5 to 7 mm.) ; middle upper premolar
not crowded inward from axis of tooth-row ;
third lower molar with second triangle nearly
as large as first and essentially like it in
form.
Foot relatively large, obviously more than half
as long as tibia; calcar about twice as long
as free border of interfemoral membrane ;
skull broad, the width of brain-case more
than half greatest length ; crown area of
molars relatively small ; upper molars with
evident protoconule.
Forearm about 47 mm. ; condylobasal length
of skull about 16 mm. (Central and

southern Europe) M. dasijcnonc, ]^. 189.

Forearm less than 45 mm. ; condylobasal length
of skull never more than 15 mm.
Tibia and adjacent membrane densely furred ;
forearm about 42 mm. ; condylobasal
length of skull 14-0 to 14-8 mm. (Medi-
terranean region) 3/. ccqxiccinii, p. 187.

Tibia and adjacent membrane not furred ;
forearm about 35 mm. ; condvlobasal
length of skull 13-2 to 13 -"S mm.

(Distribution general) J\I. daubcntonii, p. 184.

Foot relatively small, about half as long as tibia ;
calcar about as long as free border of inter-
femoral membrane ; skull narrow, the width
of brain-case less than half greatest length ;
crown area of molars relatively large ; upper
molars without protoconule.
Ear not specially elongated, extending slightly
beyond nostril when laid forward.
Forearm about 40 mm. ; condylobasal length
of skull about 15 mm.; posterior border
of ear conch with deep, almost angular

emargination slightly above middle M. cmarginatus, p. 177.

. Forearm about 34 mm.; condylobasal length
of skull 12-6 to 13-2 mm.; posterior
border of ear conch with shallow, in-
conspicuous emargination M. mystacinus, p. 169.

Ear elongated, extending conspicuously beyond
nostril when laid forward.

MYOTIS 109

Combined length of tibia and foot less than

25 mm. ; condylobasal length of skull

14 • 0 to 14 • 6 mm. ; ear narrow, its width

about 10 mm. ; tragus conspicuously

more than half as high as conch M. nattcrcri, p. 174.

Combined length of tibia and foot about

30 mm. ; condylobasal length of skull

16 to 17 mm. ; ear broad, its width about

14 mm. ; tragus scarcely half as high as

conch M. bechsteinii, ]^. 179.

MYOTIS MYSTACINUS Kuhl.

1819. Vcs^xrtiiio mystacinus Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv

(= Neue Ann., i), pt. 2, p. 202.
1821. Vespertilio collaris Schinz, Das Thierreich von Cuvier, i, p. 177

(Mt. Blanc, Haute-Savoie, France).
1834. Tcs2iertilio humcmlis Baillon, M^m. Soc. Royale d'Emulatiou

d' Abbeville, 1833, p. 50 (Abbeville, Somme, France).
1837. Vespertilio schinzii Brehm, Ornis, Heft iii, p. 27 (Renthendorf,

Thiiringen, Germany).
1843. V[espcrtilio] schrankii Wagner, Wiegmann's Archiv fiir Naturgesch.,

IX, Bd. II, p. 25 (Munich, Germany? See Fitzinger, Sitzungsber.

kais, Akad. Wissensoh. Wien, Math.-Naturwiss. Classe, lxii, pt. 1,

p. 219, 1871).
1857. Vespertilio mystacinus Blasius, Saugethiere Deutschlands, p. 90.
1863. [Brachyottis mystacinus'] var. nigricans Koch, Jahrb. des Vereins fiir

Naturkunde im Herzogthum Nassau, xviii, p. 444 (Wiesbaden,

Nassau, Germany).
1863. [Brachyotiis mystacinus'] var. rufofuscus Koch, Jahrb. dcs Vereins

fiir Naturkunde im Herzogthum Nassau, xviii, p. 444 (Wiesbaden,

Nassau, Germany).
18G3. [Bruchyotus mystacinus] var. aureus Koch, Jahrb. des Vereins fiir

Naturkunde im Herzogthum Nassau, xviii, p. 445 (Breisgau,

Germany).
1869. [Vespertilio mystacinus] var. nigricans Fatio, Faune Vert. Suisse,

I, p. 92 (Switzerland). Not of Koch, 1863.
1869. IVespertilio] lugubris Fatio, Faune Vert. Suisse, i, p. 93 (Alternative

for nigricans Fatio).
1871. Vesjwrtilio mystacinus, nigro-fuscus Fitzinger, Sitzungsber. kais.

Akad. Wisseusch. Wien, jNIath.-Naturwiss. Classe, LXii, pt. 1,

p. 217 (Renaming of V. schinzii Brehm).
1878. Vespertilio mystacinus Dobsou, Catal. Chiropt. Brit. Mas., p. 314.
1900. Myotis mystacinus M6hely, Monogr. Chiropt. Hungarise, p. 200.
1910. Myotis mystacinus Trouessart, Faune Mamm. d'Europe, p. 33.

Geoffra^hical distribution. — Entire Continent of Europe north
to about the hmit.s of tree g!-owth ; west to Irehind ; ea.st into
Asia.

Diagnosis. — Smalle.st species of European Myotis (forearm
about 34 mm., longest finger about 60 mm., condylobasal length
of skull 12'() to 13 "2 mm.) : ear moderately long, extending 1 to
2 mm. beyond tip of muzzle when laid forward, its posterior
border with shallow incons})icuous emargination ; foot about half
as long as tibia ; wing membrane extending to ba.se of outer
toe ; last caudal vertebra free.

170

CHIROPTERA

External characters. — General form slender and delicate, the
legs and tail rather long, the membranes thin and semi-trans-
parent. Muzzle with rather noticeable glandular swellings.
Ear extending slightly beyond nostril when laid forward, its
general form rather slender, the tip narrowly rounded off, the
posterior border with shallow inconspicuous concavity extending
from just below tip to near middle of conch ; inner surface of
conch without well defined transverse striations. Antitragus
small (length about 2 mm.) but well defined Tragus a little
more than half as high as conch, its width slightly above level
of antei'ior base contained about 2^ times in length of anterior
border, the anterior border straight, the posterior border convex
below ; from widest region it narrows rather rapidly upward to
a rather blunt point ; posterior basal lobe well defined. Wing
rather narrow, with no special peculiarity of form, the third,
fourth and fifth metacarpals sub-equal, their distal extremities
falling short of elbow by about 3 mm.; membrane inserted at base
of outer toe. Foot about half as long as tibia ; calcar slender,
with barely indicated rudiment of keel and terminal lobe, its
length slightly greater than that of free border of interfemoral
membrane. Tail about as long as head and body, and twice as
long as tibia, its terminal vertebi'a free from membrane.

Fnr and colour. — The fur is soft and loose, the hairs on middle
of back about 10 mm. in length, tho.se of underparts a little more
than half as long ; it is closely confined to body, extending on
wings to extreme base of membrane only, slightly farther below
than above, and on interfemoral membrane over basal fifth or
fourth of both surfaces ; free border of uropatagium not fringed.
Colour of upper j^arts a clear light brown resembling the wood-
brown of Ridgway, but usually more yellow and always with a
distinct metallic gloss, the basal portion of the hairs blacki.sh-
slate, this colour sometimes appearing at the surface and
producing a general darkening effect ; underparts paler and more
buffy, usually not forming any decided contrast with back, but
occasionally almost whiti.sh, especially on
chest ; muzzle and cheeks dusky ; ears and
membrane blackish.

Skull. — The skull is slender and lightly
built, the breadth of brain-case conspicuously
greater than that of rostrum and slightly
but appreciably less than half greatest length.
Dorsal profile rising abruptly above low ros-
trum in interorbital region and forming a
strong convexity o%er anterior portion of
Fig. 29. brain-case ; occipital region distinctly pro-

MtiothmyHacinvs. ^^^^^^^ backward and rising slightly but evi-
dently above level of anterior portion of
brain-case, from which it is marked off by a shallow but notice-
al)le transverse constiiction foUowinjj anterior margin of inter-

171

parietal. Ventral profile rising slightly but e^sidently in region
of tloor of brain-case. Greatest depth of brain-case about three-
tjuarters mastoid width ; sagittal and lambdoid crests slightly
indicated in fully adult individuals. Interorbital region obscurely
short hour-glass shaped. Anteorbital foramen small, its posterior
border over posterior root of large premolar. Posterior palatal
region rather narrow, its width immediately behind molars about
equal to its greatest length, the median spine broad and short ;
mesopterygoid fossa slightly wider than long, the hamulars turned
inward. Auditory bulla moderate, its greatest diameter nearly
equal to distance between inner margins of bullje.

Teeth. — Upper incisors sub-equal, about half as high as canine,
each pair in contact or nearly so at base but diverging at tips, the
cingulum of inner tooth horizontal, that of outer tooth oblique ;
crown of outer incisor squarish in cross section, that of inner
somewhat elongated in axis of tooth-row ; near point of contact
each shaft bears a secondarj^ cusp, this usually though not always
better developed in outer than in inner tooth ; distance between
canine and outer incisor about equal to diameter of incisor, that
between pairs a little more than
twice as great. Lower incisors
slightly but evidently imbricated,
forming a continuous, broadly
V-shaped row between
canines ; crown of i^ sub-
terete, its area more than
half that of canine and
about equal to that oi pm^,
its blunt main cusp, the
highest in the incisor series,
situated at outer side, the
three smaller cusps (third
obsolete) on inner mai'gin ;
^1 and /., sub-equal, slightl}-
lower than i'g, their crowns
compressed, longer than
high, tritid, that of second
wider posteriorly than an-
teriorly, and usually with
a minute postero-internal
cusp. Upper canine rela-
tively large, slightly higher than main cusp of large premolar,
its shaft somewhat triangular in cross section, with broad
postero-internal concavity and anterior and postero-external
groove ; posterior cutting edge well developed ; cingulum com-
plete but not forming evident secondary cusps. Lower canine
lower and less acutely pointed than upper, its tip about on level
with highest cusps of molars ; cingulum usually forming a slight
secondary cusp anteriorly. First and second upper premolars

Flo. »l.
My Otis mystaciiius. Teeth X 10.

172 CHIROPTERA

alike in foim, crowded between canine and large premolar but
perfectly in the tooth-row, the first about the same size as the
ujjper incisors, the second with slightly more than half the
height and crown area of first, both with fully developed
cingulum and subterete, conical cusp. Large upper premolar
with crown area only a little less than that of first molar, the
main cusp large and with strongly trenchant posterior cutting-
edge, its height about eijual to that of largest molar cusps :
anterior and ^^osterior borders of crown slightly concave, inner
border narrowly rounded, usually with a distinct though small
cusp anteriorly, sometimes with another barely indicated pos-
teriorly ; a shght though evident concave crushing surface
between cingulum and inner base of main cusp. First and
second lower premolars essentially similar to the corresponding
upper teeth but slightly less reduced in size and with somewhat
higher cusps ; third lower premolar similar to first and second
Init with rectangular crown nearly twice as large as that of second
tooth, and main cusp as high as protoconid of first molar ; a
small cingulum cusp usually present at antero-internal angle.
Upper molars rather large relatively to size of skull, the crown
area of second tooth slightly greater than that of first ; anterior
and posterior borders slightly concave, inner border narrowly
rounded, especially in m'^, the antero-internal and postero-internal
outlines often flattened or even a little concave ; protocone large,
its base occupying entire inner border of tooth, its cusp a little
in front of middle, its anterior commissure simple, extending
uninterrujitedl}' outward to parastyle, its posterior commissure
terminating in a thickened rib-like rudiment of a hypocone,
between the outer base of which and inner base of metacone lies
a deep furrow : metacone larger than paracone ; styles and outer
commissures well developed ; third upper molar with crown area
equal to about three-quarters that of iu^, the protocone as in the
other teeth liut smaller, paracone larger than metacone, the first
outer commissure longer than in vi'^ and m'^, the second and third
about as in the other teeth but set at a different angle, the fourth,
together with metastyle absent. First and second lower molars
with second y slightly larger than first in cross section, this
condition reversed in m^ ; protoconid decidedly higher than
hypoconid in all three teeth ; inner cusps of about equal height
throughout (slighth? more than half as high as protoconid) ; a
distinct cingulum cusji behind entoconid.

Measurements. — Adidt female from Skane, Sweden : tibia,
15 ; foot, 7 • 6 ; forearm, 32 ; thumb, 6 • 2 ; third finger, 49 ; fifth
finger, 38 ; ear from meatus, 1 2 ; width of ear, 8. Adult male from
Madrano, Tyrol: head and body, 38; tail, 38; tibia, 16; foot,
8 ; forearm, 35 : thumb, 6-2 ; third finger, 56 ; fifth finger, 47 ;
ear from meatus, 13 : width of ear, 9. Adult female from the
Carpathian Mountains : head and body, 44 ; tail, 40 ; tibia, 15 • 4 ;
foot, 8 ; forearm, 34 ; thumb, 7 ; third finger, 54 ; fifth finger.

MYOTIS 173

45 ; ear from meatus, 14 ■ 2 ; width of ear, 9. Forearm, in other
specimens : Waremme, Liege, Belgium, 33 and 33 ; tStrass, near

Burgheim, Ba^■aria, 31 '6
Switzerland, 33 and 34 '4.
p. 182.

Dresden, 33 "2 ; Grotte de Vallorbe,
For cranial measurements see Table,

Specimens examined. — Thirty-five, from the following localities ; —

England : Newby Bridge, Lake Windermere, Cumberland, 1 ; Aberia,
Merionethshire, 2; Colwyn, Denbighshire, 1; Cheadle, Staffordshire, 1;
Manchester, Lancashire, 1 ; Ragley House, 1 ; Welford, Herefordshire, 2 ;
Macclesfield, Cheshire, 1 ; Pewsey, Wiltshire, 1 ; Dover, Kent, 2 ; Hastings,
Sussex, 1 ; Bath, Somerset, 1.

Sweden: Skullno, 1; Skane, 1 (U.S.N.M.).

Belgium: Waremme, Liege, 2 (U.S.N.M.).

Germany: Moritzburg, Saxony, 1 (U.S.N.M.) ; Dresden, 1; Strass,
near Burgheim, Bavaria, 1 ; Bavaria, no exact locality, 2 (U.S.N.M.).

ArsTRiA-HuNGARY : Haida, Arva, Bohemia, 1 ; Carpathian Mts., 1
(U.S.N.M.) ; Csallokoz-Somorja, Pressburg, 2 ; Tatra Mts., Hungary, 1.

Switzerland : Geneva, 1 (Mottaz) ; Grotte de Vallorbe, Vaud, 2
(Mottaz); Stein, Appenzell, 1 (U.S.N.M.); St. Gallon, 1 (U.S.N.M.); no
exact locality, 1 (Geneva: type of luguhris Fatio).

Remarks. — Mijotis mystacinus, the smallest European member
of the genus, is recognizable by its small size and relatively short
foot in combination with the insertion of wing membrane at base
of outer toie. From the small species of Pipistrellus it is at once
distinguished by the narrow muzzle and slender tragus.

2 9.

Aberia, Merionethshire,

G. H. Caton Haigh

11. 1. 3. 13-14

Wales.

(c & P).

9 St.

Colwyn, Denbighshire.

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

6. 2. i. 1.

1 al.

Newby Bridge, Cumber-
land, England.

T. Paul (c & p).

94. 9. 3. 1.

9.

Cheadle, Staffordshire.

E. Blagg (c & p).

11. 1. 3. 15.

6.

Manchester, Lancashire.

C. Oldham (p).

11. 1. 3. 10.

6.

Ragley House, Warwick-
shire.

Tomes Collection.

7. 1. 1. 498.

2 9.

Welford, Herefordshire.

Tomes Collection.

7.1.1.496-497

6.

Pewsev, Wiltshire.
(P. S.'Hembly.)

C. H. B. Grant (p).

11. 1. 3. 11.

9.

Dover, Kent. {B. Hesse.)

C. H. B. Grant (p).

11. 1. 3. 12.

1.

Hastings, Sussex.

MissI.Roods(c&p).

49. 1. 16. 1.

6 St.

Bath, Somerset.

G. Dalgleish (c & p).

4. 10. 13. 3.

1.

Skullno, Sweden.

Stockholm ^luseum

46. 1. 2. 22.

9.

Haida, Bohemia. (Ti"u/-
terstorff.)

Lord Lilford (p).

11. 1. 1.6.

1.

Csallokoz - Somorja,

Budapest Museum

94. 3. 1. 20-21

Pressburg, Hungary.

(E).

lal.

Tatra Mountains,

Dr. R. CoUelt (p).

91. 1. 21. 1.

Hungary.

174 CHIEOPTEEA

MYOTIS NATTERERI Kuhl.

1818. Vespertilio nattereri Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv

(= Neue Ann., i), pt. 1, p. 33.
1857. Vespertilio nattereri Blasius, Siiugethiere Deutsclilands, p. 88.
1863. [Isotus nattereri'] var. typhis Koch, Jahrb. des Vereins fiir Natur-

kunde im Herzogthum Nassau, xviii, p. 430 (Wiesbaden, Nassau,

Germany).
1863. [Isotus nattereri] var. spelxus Koch, Jahrb. des Vereins fiir Natur-

kunde im Herzogthum Nassau, xviii, p. 430 (Erdbach, Nassau,

Germany) .
1878. Vespertilio nattereri Dobson, Catal. Chiropt. Brit. Mus., p. 88.
1900. Myotis nattereri M6hely, Monogr. Chiropt. Hungarise, p. 179.
1904. Myotis escalerai Cabrera, Mem. Soc. Espau. Hist. Nat., ii, p. 279

(Bellver, Lerida, Spain).
1910. Myotis nattereri and M. escalerai Trouessart, Faune ilamm. d'Europe,

pp. 29-30.

Type locality. — Hanau, Hessen, Germany.

Geographical distribution. — Central and southern Europe,
west to Ireland, north to southern Sweden.

Diaijnosis. — Size medium among the European species (forearm
about 38 mm., longest linger about 70 mm., condylobasal length
of skull, 14 "0 to 14 '6 mm.); ear elongated, extending about
5 mm. beyond tip of muzzle when laid forward, the conch narrow
(about 10 mm.), obscurely emarginate on upper half of posterior
border, the tragus relatively longer than in any other European
species, its height distinctly more than half that of conch ; foot
about half as long as tibia ; wing membrane extending to base of
outer toe ; edge of interfemoral membrane fringed.

External characters. — General form essentially as in Myotis
mystacinus, but differing in the following particulars : ear much
longer and relatively narrower, extending conspicuously beyond
nostril when laid forward, its extremity more broadly rounded
off (owing to more uniform convexity of anterior border), and
concavity on posterior border even less evident ; antitragus
about as large as in 31. mystacinus but less well defined ; tragus
relatively longer and more attenuate than in any other European
bat, its height conspicuously more than half that of conch, its
gi'eatest width contained about 3^ times in length of anterior
Iwrder, its terminal third almost linear, sometimes faintly
recurved, the basal lobe usually small and ill defined ; insertion
of wing membrane as in 31. mystacinus, but membrane at first
very narrow, so that in some specimens, particularly those that
have been hardened in strong alcohol, the point of insertion
appears to be on side of metatarsus ; * tail rather shorter than
head and body, only the cartilaginous extreme tip free ; calcar
l)etter defined than in 31. mystacinus, about as long as the
distinctly fringed free border of interfemoral membrane.

* Such specimens seem to have formed the basis of the Myotis escalerai
of Cabrera.

MYOTIS 175

Fur and colour. — Quality and distribution of fur essentially as
in M. mydacinus, but free border of interfemoral membrane
distinctly though not very densely fringed with hairs about
1 mm. long. Colour of upper parts a lighter and less yellowish
brown than in M. mystacinua, the exact shade intermediate
between the wood-brown and broccoli-brown of Ridgway, the
longer hairs with faintly darker tips visible in certain lights ;
underparts rather sharply contrasted whitish buffy grey ; a well
defined line of demarcation extending from shoulder to base of
ear ; basal portion of hairs clove-brown ; muzzle and cheeks dusky,
but not so dark as in M. mi/stncinus ■ ears and membranes dark
brown.

Shill. — In all its dimensions the skull is appreciably larger
than that of Myotis mysiacinus. In general form it is slightly
less slender. Forehead rising moi'e abruptly but occipital region
not higher than main portion of brain-case, so that dorsal profile
shows a stronger concavity in interorbital region than that of
M. mystacinus, while over greater extent of brain-case it is
essentially flat, or slightly falling away posteriorly, instead of
rising by two well defined curves to lambdal region. Posterior
palatal region about as in M. mystacinus, except that mesopterygoid
space extends further forward, and median spine is better
developed. Auditory bulla relatively smaller than in M. mysta-
cinus, its greatest diameter decidedly less than distance between
bullae.

Teeth. — In general the teeth resemble those of Myotis
mystacinus apart from their greater size. They differ, however,
in certain details of form : crown area of outer upper incisor
appreciably greater than that of inner tooth ; angle at front of
lower incisor row wider, though evident ; upper canine relatively
weaker, its posterior cutting edge less developed, the cross section
of its shaft half-terete owing to the obsolescence of postero-
external and anterior longitudinal furrow ; lower canine with
length of base much greater in proportion to height of shaft, and
cingulum more oblique ; second upper premolar nearly equal to
first in cross section ; crown area of large premolar not so great
relatively to that of first molar, the cusps on inner border
obsolete or absent ; lower premolars with less slender crowns,
the cingulum of the third forming a very low, sometimes obsolete
antero-internal cusp ; molars, especially m^, with narrower
crowns.

Measitrenwnts. — External measurements of adult male from
Colpin, Brandenburg, Germany, and adult male from Magdeburg,
Germany: head and body, 50, 44; tail, 41, 40; tibia, 17, 16 ;
foot, 8-2, 9-0; forearm,"' 39-2, 40-2 ; thumb, 7-0, 6-2; third
finger, 71,71 ; fifth finger, 56, 56 ; ear from meatus, 18, 18 ; tragus,
11*0, 11 • 2. Two adult females from Spain (Xo. 94. 1 . 1.8, Se-\-ille,
and No. 8. 7. 23. 4 from Deliver, Lerida, paratype of Jlf. escalerai) :
head and body, 46, 46 ; tail, 4 1 , 43 ; tibia, 1 5, 1 7 ; foot, 7 • 6, 9*2;

176

CHIROPTERA

forearm, 36 -4, 40" 6 ; thumb, 7 -2, <S'4 ; third finger, 69, 72 ;
fifth finger, 52, n6 ; ear from meatus, 16, 16" 6; tragus, 10*2,
10±. For cranial measurements see Table, pp. 182-183.

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

Scotland : Inverary, Argyllshire, 1.

England: Harlech, Merionethshire, .3; Arrow Church, G; Alcester,
Warwickshire, 8; Lilford Hall, Northamptonshire, 5; Henley-on-Thames,
Oxfordshire, 3 (B.M. and U.S.N.M.) ; Bradfield, Berkshire, 1 ; Queen
Camel, Somersetshire, 1 ; Devonshire, 1 ; no exact locality, 3.

Ireland: Co. Longford, 1; Woodpark, Co. Galway, 1.

Germany: Oberlausitz, Silesia, 1 ; Colpin, Brandenburg, 1 ; Magdeburg,
Saxony, 2; Moritzburg, Saxony, 3 (U.S.N.M.).

Austria-Hungary: Haida, Arva, Bohemia, 1.

Switzerland: Canton Thurgau, 3 (U.S.N.M.); St. Gallen, 9 (B.-M.
and U.S.N.M.).

Italy: Arezzo, 4 (B.M. and U.S.N.M.) ; Valesia, 1 (U.S.N.M.) ; Siena, 1
(Mottaz) ; no exact locality, 4 (U.S.N.M.).

Spain: Bellver, L^rida, 2 (B.M. and Genoa; paratypes of escalerai
Cabrera) ; Seville, 1.

lii'hKi.rJis. — This species is easily recognized by its rather small
size, large ear, and very long, attenuate tragus. The exact point
of insertion of wing membrane on side of foot is probably in all
specimens the base of outer toe, but owing to a peculiar narrow-
ing of tlie membrane along edge of metatarsal, the point of
insertion sometimes appears to be at middle of side of foot. This
effect is often increased by the action of strong alcohol.

9. Inverary, Argyllshire,

Scotland.
1 al. Longford, Ireland.

9. Woodpark, Galway.

9, 9 juv. Harlech, Merioneth-
shire, Wales.
G. Arrow Church, Warwick-

shire, England.
Alcester, Warwickshire.
Lilford Hall, Northamp-
tonshire.
Bradfield, Berkshire.
Henley, Oxfordshire.
Henley, Oxfordshire.
Queeji Camel, Somerset.
England.
9. O be rlaus it z, Silesia,

340 m. Germany.
(W. Baer.)
6 al. Colpin, Brandenburg,

Prussia.
6 al. 3Iagdeburg, Saxony.

Duke of Argyll (p)

58. 8. IG. 1.

Dr. G. E. Dobson 76. 11. 3. 2.

(c & P).

R. F. Hibbert (p). 11. 1. 3. 20.

J. Backhouse (p). 11. 1. 3. 16-18.

8 al.

3 9 al

9.

9 St.

6 St.

(Jal.

1 al.

Tomes Collection.

Tomes Collection.
Lord Lilford (c & p).

N. H. Joy (c & p).
J. G. Millais (c & p).
Heatley Noble (c & p).
R. H. Read (p).
Dr. J. E. tTray (p).
Lord Lilford (p).

7.1.1.488-493.

7.1.1.736-743.
72. 8. 21. 3-.5.
72. 11. 12. 13.
11. 1. 3. 19.

I. 11. 2. 1.
0. 3. 23. 1.

II. 1. 3. 21.
•51. 1. 29. 12.
99. 1. 9. G.

Dr. H. Gadow (c & p). 82. 7. 31. 2.
Dr. W. Wolterstorff 92. 12. 1. 1.

3 S, 9.

jMagdeburg, Saxony.
{Wolterstorff.)

Haida, Bohemia. (ll'oZ-
terstorff.)

St. C^allen, GOO m. Swit-
zerland. {E. H. Zolli-
kofer.)

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

11. 1. 1. 8-9.

11. 1. 1.

4. 4. 5. 4-8.

MYOTIS 177

2 al. Arezzo, Italy. Florence Museum (e). 85. 7. 6. 4-5.

9 al. Bellver, Lc^Tida, Spain. A. Cabrera (p). 8. 7. 23. 4.

(Paratype of M. escalcrai, Cabr.)
1 al. Seville. Dr. V. L. Seoane 94. 1. 1. 8.

(c & p).

MYOTIS EMARGINATUS Geoffroy.

1806. Vesp{ertilio] emarginatus Geofiroy, Ann. Mus. d'Hist. Nat., Paris,
VIII, p. 198 (Charlemont, Givet, Ardennes, Prance).

1844. Vesp\crtilio'] rufescens Crespon, Paune Meridionale, i, p. 20 (near
Nimes, Gard, Prance). Type in Nimes Museum.

1853. V[espertilio] ciliatus Blasius, Wiegmann's Archiv fiir Naturgesch.,
1853, I, p. 287 (near Cologne, Germany).

1856. V[es2}ertilio] schrankii Kolenati, Allgem. deutsche Naturhist. Zeitung,
Dresden, neue Polge, ii, p. 178. A nomen nudum (ex Koch and
Giebcl) cited as sj'nouym of emarginatus. Not of Wagner, 1843.

1858. Vespertilio ciliatus Blasius, Siiugethiere Deutschlands, p. 91.

1878. Vespertilio emarginatus Dobson, Catal. Chiropt. Brit. Mus., p. 303.

1880. M)/otis ciliata var. buclapcsticnsis Margo, " INIagyar orv. es term^s-
zetvizsg. xx,nagygyul. munk, p. 255" (Budapest, Austria-Hungary).
See M6hely, Monogr. Chiropt. Hungarife, p. 170.

1890. Vespertilio neglectus Patio, Arch. Sci. Phys. et Nat., Geneve, 3rd ser.,
XXIV, p. 512, November 15, 1890 (Valavran, near Geneva, Swit-
zerland). Type in Geneva Museum.

1900. Mijotis emarginatus Mehely, Monogr. Chiropt. Hungarise, p. 170.

1910. Mrjotis e^narginatus Trouessart, Paune Mamm. d'Europe, p. 28.

Tj/jje locality. — Charlemont, Givet, Ardennes, France.

Geographical distribution. — Central and southern Continental
Europe.

Diagnosis. — Size essentially as in 3Ti/otis naitereri (forearm
about 40 mm., condylobasal length of skull about 15 mm.) ; ear
moderately long, extending 2 to 3 mm. beyond tip of muzzle
when laid forward, its posterior border with a deep almost
angular emargination slightly above middle ; foot about half as
long as tibia ; wing membrane extending to base of outer toe : free
margin of interfemoral membrane sometimes fringed ; fur some-
what woolly in texture, the hairs of back tricolored, drab at
base, yellowish at middle and dark brown at extreme tip.

External characters. — AVhole animal larger and more robust
than Mijotis miistacinus, though of essentially the same form and
proportions. Ears and membranes relatively thick and leathery,
not semi-transparent as in the related small species. Ear
moderately long, extending about 2 mm. beyond nostril
when laid forward, its size and general form much as in M.
mystacinus except that emargination of posterior border is deep
and conspicuous, in most specimens forming an evident angle
below. Inner siu'face of conch marked bv seven or eight short
but well developed cross ridges. Antitragus small and ill-defined.
Tragus slightly more than half as high as conch, its greatest
width contained nearly three times in length of' anterior border,
both its margins essentially straight from level of anterior base

N

178 CHIROPTERA

to rather acute tip. Wing as in the related species ; metacarpals
falhng short of elbow by about 3 mm. ; membrane inserted at
base of outer toe. Calcar slender, its termination usually marked
by a distinct lobe, its length about equal to that of free border
of interfemoral membrane. Tail relatively shorter than in 31.
mystacinus, extending to between ears when laid forward, onh'
the minute cartilaginous tip free from membrane.

Fur and colour. — Fur shorter and more dense than in M.
mystacinus and M. naUereri, and of a slightly woolly texture
unique among the European species, the hairs at middle of back
about S mm. in length. Distribution of fur not peculiar ; free
border of uropatagium sometimes with evident fringe. General
colour buff, light and clear on underparts, dulled and irregularly
clouded by darker brownish hair tips throughout upper parts ;
basal half of hairs drab ; muzzle and cheeks dusky ; ears and
membranes an indefinite rather light brown.

SJiulI. — In form the skull does not differ appreciably from
that of M. naUereri, except that the rostral portion and palate are
relatively more elongate and interorliital concavity is less strongly
pronounced ; occipital region similarly low as compared with that
of M. nnjsfacinns. Mandible slightly more robust than in 31.
naUereri, but of similar form.

Teeth. — The teeth are larger than those of 3Iiiot!s mystacinus,
in this respect agreeing with those of M. natteieri. Upper
incisors higher and more slender than in the preceding species,
their crowns sub-equal in cross section ; lower incisors very
slightly imbricated, forming a broadly and e^•enly convex
(U-shaped) row without anterior angle, the crowns of (j and i.,
alike in form and distinctly 4-cusped, that of i^ not thickened
posteriorly and with no trace of postero-internal cusp. Canines
about etiual to those of 31. nattereri in size, but shaft of upper
tooth with evident postero-external longitudinal groove. First
and second upper premolars even more strongly contrasted in
size than those of 31. mystacinus, the cusp of second only a little
exceeding cingulum of first ; the two teeth less crowded between
canine and large premolar than in 31. mystacinus and M. nattereri ;
lower premolars essentially as in 3T. nattereri, but less closely
crowded, and second relatively larger. Upper molars as in
M. nattereri, but crowns less narrowed ; a rudimentary commissure
extending outward from base of hypocone and another extending
inward from base of paracone ; lower molars not peculiar.

3'leasurements. — Two adult females from Florence, Italy :
head and body, 46-6 and 50 ; tail, 40 and 4-2 ; tibia, 19 and 19 ;
foot, 8-4 and 8-6; forearm, 40 and 41 ; thumb, 8 and 7-8;
third finger, 67 and 70 ; fifth finger, 56 and 57 ; ear from meatus,
16-6 and 17 ; width of ear, 11-4 and 12. For cranial measure-
ments see Table, p. 183.

Specimens examined. — Six, from the following localities : —
Holland : Maastricht, 1 (U.S.N.M.).

MYOTIS 179

Prance : Near Nimes, Gard, 1 (Nimes : type of rufcscens Crespon).
AusTBiA-HuNGARY : Herkulesbad, 2.

Switzerland: Valavran, near Geneva, 1 (Geneva: type of nenlectus
Fatio). -^

Italy: Florence, 2 (U.S.N.M.) ; no exact locality, 1.

Remarks. — The peculiar form of the ear, the short, somewhat
woolly fur, and the yellowish colour are highly characteristic of
this well-detined species.

2 9. Herkulesbad, Hungary. Hon. N. C. Roths- 7. 9. IG. 9-10

child (p).
skeleton Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 73:3.

without parte.)
skull.

MYOTIS BECHSTEINII Kuhl.

1818. Vespertilio bechsteinii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv

(= Neue Ann., i), pt. 1, p. 30 (Hanau, Hessen, Germany).
1857. Vespertilio bechsteinii Blasius, Siiugethiere Deutschlands, p. 85.
1878. Vespertilio bechsteinii Dobson, Catal. Chiropt. Brit. Mus., p. 308.
1900. Myotis beclistcinii M6hely, Monogr. Chiropt. HungariiB, p. 184.

1905. Vesplertiliol bechst{einii\ ghidinii vel Vcs2}[ertilio'] ghidinii Fatio,

Arch. Sci. Phys. et Nat., Geneve, 4th ser., xix, p. 511, May 15,
1905 (Lugano, Ticino, Switzerland). Type in Geneva Museum.

1906. Myotis beclistcinii favonicus Thomas, Ann. and Mag. Nat. Hist.,

7th ser., xviii, p. 220, September, 1906 (La Granja, Segovia,
Spain). Type in British Museum.
1910. Myotis bechsteini and M. bechstcini favonicus Trouessart, Fauue
Mamm. d'Europe, pp. 30-81.

Ti/pe localiti/. — Hanau, Hessen-Nassau, Germany.

Geographical distrihution. — Central and southern Europe,
west to England, north to southern Sweden.

Biagnods. — Size slightly greater than that of M. nattererl
(forearm about 40, condylobasal length of skull, 16 to 17) ; ear
elongated, extending about 8 mm. beyond tip of muzzle when laid
forward, the conch broad (about 15 mm.), its posterior margin
obscurely emarginate above, the tragus scarcely half as high
as conch ; foot about half as long as tibia ; wing membrane
extending to base of outer toe.

External characters. — In general like Myotis nattereri, but with
broader ears and relatively as well as actually larger legs and
feet (combined length of tibia and foot about oO mm. instead of
less than '25). Muzzle with moderately developed glandular
swellings (these less evident than in M. mystacinus). Ear extend-
ing considerablj' beyond nostril when laid forward, its length thus
about as in M. nattereri, but its breadth so much greater (about
17 mm. instead of about 10 ram.) that the ear is relatively larger
than in any other European bat except Plecotus auritns ; form
of ear essentially as in M. mystacinus, the upper half of posterior
border faintly concave ; inner surface of conch with about eight
rather ill-delined transverse ridges near posterior border ; anti-
tragus about '1 ' 5 mm. in length along base, rather abruptly

N 2

180 CHIROPTERA

rounded off above, its inner margin not continuous with posterior
border of conch. Tragus about half as high as conch, its form
essentially as in M. mystacinus though a little more slender
(greatest width contained nearly three times in height of
anterior border) ; basal lobe small but well developed. Wing
essentially as in M. mystacinus ; third, fourth and tifth meta-
carpals sub-equal, falling short of elbow by about 5 mm. ;
membrane inserted at base of outer toe. Calcar and free border
of interfemoral nieml^rane as in M. mystacinus. Tail shorter
than head and body (laid forward it extends to middle of crown),
the terminal vertebra free.

Fm- and colour. — Quality and distributicjn of fur as in Myotis
iity.^tacinas and M. nattereri. Upper jaarts uniform wood-brown
with a slight tinge of umber, the exact shade almost impossible
to describe exactly ; underparts a huffy grey in strong contrast,
irregularly clouded by the slaty brown of underfur ; ears and
membranes an indefinite dark brown.

Skull. — In general aspect the skull resembles that of ilf//o//.s'
myotis, due allowance being made for its much smaller size, since
it is much more slender than in any of the other small species ;
breadth of brain-case equal to about one-half distance from
lambda to posterior border of narial emargination. Dorsal
profile rising gradually above rostrum and forming a strong
convexity over anterior jiortion of brain-case, behind which it is
essentially flat to lambda ; ventral profile nearly flat, slightly
elevated posteriorly ; occipital region slightly projecting, just
sufficiently to conceal condyles when viewed from above. Brain-
case ovate, noticeably wider than rostrum ; sagittal crest low but
evident ; lambdoid crest moderately develo2:>ed at sides, obsolete
at middle ; greatest deptli of brain-case about two-thirds mastoid
breadth ; floor of brain-case with obscurely defined lateral grooves.
Interorbital region not evidently hour-glass shaped, owing to the
slight widening at lachrymal level ; anterior upper border of
orbit slightly ridged. Rostrum slender, widening a little at
anterior extremity, the dorsal surface smoothly rounded off at
sides, though with indication of a slightly developed median
groove ; narial emargination narrow but deep, extending back
about half way to level of anterior rim of orbit ; rostral depth
at front of orbit about equal to distance from orbit to middle of
canine ; palate long and narrow as compared with that of the
other small species, both transverse and lateral concavities evident,
especially just behind middle ; anterior emargination rather large,
sub-circular in general outline, extending back about to level of
space between canine and first premolar ; posterior extension of
palate about as broad as long ; interpterygoid space wider than
long, encroached on by blunt median palatal spine. Mandible
essentially as in Myotis myotis, the coronoid process higher and
with more obli(iue posterior border than in the other small species.
Teeth. — The teeth are rather small relatively to size of skull.

MYOTIS 181

Upper incisors essentially as in 31. mijutaciniis but more slender.
Lower incisors slightly imbricated, the row as a whole (J -shaped
or very broadly V-shaped, the form of the individual teeth not
peculiar. Canines with no special peculiarities. Small upper
premolars completely in tooth-row, not crowded, the crown area of
first eijual to about half that of canine, that of second to about
two-thirds that of first, the height of first a little less than half
that of canine, that of second about two-thirds that of first ;
cingulum complete but not forming secondary cusps. Large
upper premolar with crown area nearly three-(|uarters that of
first molar, its antero-internal cusp well developed. Lower
premolars essentially as in 31. nattereri. Upper molars with no
special peculiarities except that wi^ is more reduced than in the
other small species, its tranverse diameter through metacone
noticeably less than half length of anterior border. Lower
molars normal, but second triangle of m^ a little more reduced
than in the other small species, though much less so than in
3/1. myotis and M. oxygnatlius.

Measurements. — Adult female from Sweden : head and body,
46 ; tail, 38 ; tibia, 19 • 6 ; foot, 9 • 8 ; forearm, 42 ; thumb, 9 • 6 ;
third finger, 67 ; fifth finger, 56 ; ear from meatus, 22 ; ear
from crown, 19; width of ear, 13-6; tragus, 10. Two adults
from the New Forest, Hampshire, England : tibia, 18 '8 and 20 ;
foot, 9 • 8 and 10 • 6 ; forearm, 39 and 42 • 4 ; thumb, 8 • 4 and 10 ;
third finger, 64 and 68 ; fifth finger, 53 and 57. Adult male
from Lugano, Switzerland (Geneva : type of ghidinii Fatio) :
tibia, 19-4; foot, 10; forearm, 41-2; thumb, 10-4; third
finger, 65 ; fifth finger, 53 ; ear from meatus, 21 ± ; ear from
crown, 19; width of ear, 13; tragus, 11±. Adult female from
Ste. Baume, Var, France : head and body, 53 ; tail, 37 '4 ; tibia,
20 ; foot, 10' 4; forearm, 42-6 ; thumb, 9 ; third finger, 70 ;
fifth finger, 57 ; ear from meatus, 25*6 ; ear from crown, 24;
width of ear, 16*2 ; tragus, 11 "8. Adult male from Cadillac,
Gironde, France : head and body, 45-|- ; tail, 42 ; tibia, 20 ;
foot, 9; forearm, 41 ; thumb, 8-8; third finger, 67; fifth
finger, 56; ear from meatus, 26; width of ear, 15 "6. Adult
male from La Granja, Segovia, Spain (type of favoniciis Thomas) :
head and body, 48 ; tail, 38 ; tibia, 20 ; foot, 8 • 4 ; forearm, 40 ;
thumb, 8-2 ; third finger, 66 ; fifth finger, 55 ; ear from meatus,
23; ear from crown, 19; width of ear, 14-6; tragus, 10 '4.
Adult female from Zay-Ugrocz, Hungary : head and body, 46 • 6 ;
tail, 44 ; tibia, 22 ; foot, 9 ; forearm, 44 ; thumb, 9 ; third
finger, 73 ; fifth finger, 59 ; ear from meatus, 25 ; ear from
crown, 22"6; width of ear, 16; tragus, 11 '8. For cranial
measurements see Table, p. 183.

Specinmns examined. — Seventeen, from the following localities : —
England : Henley-on-Thames, Oxfordshire, 1 ; New Forest, Hamp-
shire, 6.

Sweden : Skdne, 1 (U.S.N.M.).

182

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184

CHIROPTERA

France : Etupes, Doubs, 1 (Mottaz) ; Ste. Baume, Var, 1 (Genoa) ;
Cadillac, Gironde, 1 (Lataste).

Switzerland: Lugano, Tioino, 1 (Geneva: type of gJiidinii Fatio).
Spain: La Granja, Segovia, 1 (type oi favonicu^ Thomas).
Austria-Hungary: Zay-Ugrocz, Trencsen, 4 (B.]\L and U.S.N.M.).

Bemarks. — This animal is recognizable among European bats
by its medium size and very large ears. On the basis of the
material examined I am unable to distinguish a Spanish or
western geographical race.

6. Henley-on-Thames, Ox- J. G. Millais (p). 6. 9. 14. 1.

fordshire, England.

1. New Forest, Hampshire. Dr. W. E. Leach (p). 56. a.

2. New Forest, Hampshire. G. W.H.Blagg(c&p). 7.7.16.1-2.
6 al. La Granja, Segovia, M. de la Escalera (c). 6. 11. 4. 1.

Spain. {Type oi M. favoiiiciis Thos.)

6, ? al. Zay-Ugrocz, Trencsen, Budapest IMuseum G. 6. 20. 1-2.
Hungary. (e).

1. Europe. Leydeii Museum. 37. 4. 28. 23.

MYOTIS DAUBENTONII Kuhl.

1819. Vcspcrtilio danbcntonii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv

(= Neue Ann., i), pt. 2, p. 195 (Hanau, Hessen, Germany).
1839. Vespcrtilio xdilis Jenyns, Ann. Nat. Hist., in, p. 73, April, 1839

(Aukland St. Andrew, Durham, England).
1844. V[espertilio] lanatus Crespon, Faune Meridionale, i, p. 15 (South of

Nimes, Gard, France). Type in Nimes Museum.
1857. Vcspcrtilio daubcntonii Blasius, Stiugethiere Deutschlauds, p. 98.
1871. Vcspertilio capucincUus "Koch, Bayr. Fauna," Fitzinger, Sitzungsber.

kais. Akad. Wissensch. Wioi, Math.-Naturwiss. Classe, lxii, pt. 1,

p. 206 (Bavaria?).
1871. Vcspcrtilio minutcllus " Koch, Bayr. Fauna," Fitzinger, Sitzungsber.

kais. Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1,

p. 206 (Bavaria?).
1871. Vcspertilio daubcntonii, albus Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1, p. 210

(Kenaming of V. xdUis Jenyns).
1878. Vcspertilio daubentonii Dobsou, Catal. Chiropt. Brit. Mus., p. 297.
1890. Vcspcrtilio staufferi Fatio, Faune Vert. Suisse, v, 3"'" suppl. aux

JIamm., p. 6 (Lucerne, Switzerland). jNIS. synonym; repudiated.
1900. Myotis daubcntonii M6hely, Monogr. Chiropt. Hungaria", p. 164.
1910. Myotis daubentoni Trouessart, Faune Mamm. d'Europe, p. 27.

Type locality. — Hanau, Hessen-Nassau, Germany.

Geo(jrapliical dhtrihution. — Europe from the Mediterranean
north to central Norway, west to Ireland, east into Asia.

Diagnosis. — 8ize nearly as small as in Myotis viystacinas
(forearm about 35 mm., longest finger about 60 mm., condylo-
basal length of skull, 1.3-4 to 13 '8); ear moderately long,
extending about 2 mm. beyond tip of muzzle when laid forward,
its posterior border with shallow inconspicuous emargination ;
foot decidedly more than half as long as tibia ; wing membrane
extending to side of metatarsus ; last caudal vertebra free.

MYOTIS 185

External characters. — Smallest of the large-footed European
species. General form less slender and delicate than in M.
mystacinun, the tail and legs relatively shorter. Ear rather
short, extending, when laid forward, about to tip of muzzle ;
anterior border faintly and evenly convex from basal lobe to
narrowly rounded-off tip ; posterior border with shallow though
evident concavity above ; antitragus small and ill-defined ; tragus
about half as high as conch, its greatest width contained three
times in length of anterior border, the posterior border
moderately and evenly convex, the anterior border nearly
straight, the tip rather blunt ; posterior basal lobe relatively
large and well defined. Wing slightly broader than in 31. mi/sta-
cinus, the three main metacarpals evidently graduated from
third to fifth, the third very slightly shorter than forearm ;
membrane inserted at middle of metatarsus. Foot large,
appreciably more than half as long as tibia ; calcar slender,
very long, without keel on posterior border and with slight
terminal lobe, its length fully double that of free border of
interfemoral membrane. Tail about as long as body without
head, the terminal vertebra free from membrane except for an
exceedingly narrow strip extending outward along each side.

Fur and colour. — Fur slightly shorter and more dense than
that of 31. mystac'mus, but with no peculiarities of distribution
except that it tends to spread farther outward along dorsal
surface of interfemoral membrane ; free border of urojiatagium
not fringed, but a slight fringe is usually present along basal
half of calcar. Colour above essentially as in ilf. nattereri,
though usually inclining more definitely toward wood-brown ;
underparts buffy grey visually less contrasted than in 31. nattereri,
though sometimes pale enough to produce a distinct line of
demarcation along sides of neck. Muzzle and cheeks dusky.
Membranes and ears an indefinite brown.

Skull. — The skull of Myotis dauhentonii is smaller tlian that
of any other European species except 31. niystacimifi. From this
it is immediately distinguishable by its noticeably greater breadth
both of rostrum, palate and brain-case, by the relatively lower
occipital region, and relatively deeper rostrum. Posterior exten-
sion of palate short and broad, the width just behind molars
greater than length to tip of hamular ; median projection angular,
seldom forming a distinct spine. Greatest breadth of brain-case
slightly though appreciably more than greatest length of skull.
Mandible with coronoid process low, scarcely rising above level
of condyle.

Teeth. — Teeth relatively smaller than those of 3lyotiS mysta-
cinus, a difference particularly noticeable in the crown areas of
the first and second upper molars. Upper incisors as in mystacinus,
but with cingulum less developed. Lower incisors very slightly
imbricated, tlie form of the row vacillating between (J-shaped
and broadly V-shaped, the cusps as in 31. mystacinus but less

186

CHIROPTERA

Fig. 31.
Miiotif: dauheidonii. Teetli X 10.

developed. Canines both above and below weaker and less
trenchant than in M. mijist acinus, their form essentially as in

M. nattereri. Premolars
and lower molars not
obviously diflerent from
those of M. Dii/stacinus.
Upper molars peculiar as
compared with those of
the small-footed European
species in the presence of
an evident protoconule on
anterior commissure of
protocone, the small cusp
provided with a small
but distinct secondary
commissure extending to
base of paracone ; m^ and
fii^ with a small commis-
sure extending from base
of hypocone to base of
metacone and partly fill-
ing depression lying be-
tween these cusps.

Measurements. — Two
adult males from Upsala,
Sweden : head and body, 43 and 44 ; tail, 34 and 36 ; tibia, 17
and 17 '4 ; foot, 11 and 11 ; forearm, 37 and 37 ; thumb, 8 '4
and 8 ; third finger, 62 and 59 ; fifth finger, 49 and 49 ; ear
from meatus, 13 and 13 ; width of ear, 10 and 9 "6. Adult male
and female from Lecco, Italy : head and body, 42 and 45 ; tail,

36 and 39; tibia, 16 and 17; foot, 10'6 and 11; forearm,

37 and 38 ; thumb, 8 and 8 ; third finger, 62 and 62 ; fifth finger,
51 and 51 ; ear from meatus, 13 '6 and 13' 6 ; width of ear, 9*6
and 9 "4. For cranial measurements see Table, p. 190.

Specimens examined. — Seventy-three, from the following localities : —

Scotland : No exact locality, 1.

England : Bowdon, Cheshire, 1 ; Knutsford, Cheshire, 1 ; Stratford-
on-Avou, Warwickshire, 2 ; Hillingdon, Middlesex 1, ; Epping, Essex, 2 ;
Northamptonshire, 1 ; Henley-on-Thauies, Oxfordshire, 2 (B.M. and
U.S.N.M.); Christchurch, Hampshire, 1; Devonshire,!.

Sweden: Upsala, 3 (B.M. and U.S.N. M.); upland, 1; no exact locality, 2.

Switzerland : Geneva, 1 (INIottaz).

Fbance : Near Nimes, Gard, 1 (Nimes : type of lanatus Crespon).

Italy : Lecco, Lombardy, 21 (U.S.N.M.) ; Pavia, 1 (U.S.N.M.);
Florence, 30 (Mottaz).

Heniarks. — This species is immediately recognizable among
the European members of the genus by its small size, large foot,
and naked upper surface of legs.

1. Scotland. Dr. J. !Macgiilivray (p).

Bowdon, Cheshire, Eng- T. A. Coward (c&p). 11. 1. 3. GO.
land.

6.

MYOTIS 187

9. Knutsford, Cheshire. T. A. Coward (c & r). 11. 1. 3. 22.

2. Stratford-on-Avon, War- Tomes Collection. 7.1.1.486-487.

wickshiro.

i al. Hilliugdon, Middlesex. O. Thomas (c & p). 84. 1. 29. 1.

2 al. Epping, Essex. H. Doubleday (c &p). 44. 10. 21. 1-2.

al. Northampton. Mrs. Jenyns (p).

9 St. Henley, Oxfordshire. J. G. Millais (c & p). 1. 11. 2. 2.

1 al. Christchurch, Hamp- Lord Lilford (p). 87. 'J. 1. 1.

shire.

6. Upland, Sweden. Lord Lilford (p). 11. 1. 1. 25.

(G. KoUhoff.)

2. Sweden. Stockholm Museum 46. 6. 2. 15.

(e). 48. 6. 28. 3.

MYOTIS CAPACClNll Bonaparte.

1837. Vespertilio capaccinii Bonaparte, Iconogr. Faun. Ital., i, fasc. xx

(Sicily). Type in British Museum.
1839. Vespertilio megapodius Temmiuck, Monogr. de Mamm., ii, p. 189

(Sardinia).
1841. Vesp[crtilio'] dasypiis de Selys-Longchamps, Atti della seconda

Riunione degli Scienziati Italiani, Torino, 1840, p. 249 (Published

as synonym of capaccinii).
1844. Vesplcrtilio] pellucens Crespou, Fauna Meridionale, i, p. 16 (Cave

near Pont-du-Gard, Gard, France).
1857. Vespertilio capaccinii Blasius, Saugethiere Deutschlands, p. 101.

1877. Vespertilio blasii Major, Atti Soc. Tosc. Sci. Nat., Pisa, iir, p. 108

(New name for the capaccinii of Blasius should it prove to be
different from that of Bonaparte).

1878. Vespertilio majori Ninni, Atti Reale Instit. Veneto, 5th ser., iv,

pt. 1, p. 721 (Substitute for blasii Major).
1878. Vespertilio capaccinii Dobson, Catal. Chiropt. Brit. Mus., p. 293.
1901. Myotis capaccinii Thomas, Proc. Zool. Soc. London, p. 37.
1910. Myotis {Leuconoe) capaccinii Trouessart, Faune Mamm. d'Europe, p. 26.

Type locality. — Sicily.

Geographical (ligtrihiition. — Mediterranean region, north to
Italian Switzerland, east into A.sia.

Diagnosis. — Not so small as Myotis dauhentonii (forearm about
42 mm., longest linger about 68 mm., condylobasal length of
skull, 14"0 to 14*8 mm.), but similar in form and proportions;
wing membrane extending to ankle ; last caudal vertebra free ;
tibia and adjacent membrane densely furred.

External cltaracters. — In all essential features the external
form is as in M. dauhentonii. The foot, however, is relatively
larger, and the wing membrane is inserted at the ankle.

Fur and colour. — Fur rather dense and short, the hairs at
middle of back about 6 mm. in length ; distribution peculiar
among the European members of the genus in its tendency to
spread on membranes, forming a tlistinct patch on upper surface
of wing at elbow and extending over entire uropatagium to level
of feet, the furry covering of tibia and immediately adjacent
membrane (both above and below) especially dense. Colour

188

CHIROPTERA

above a light drab tinged with grey or with ecru-drab, the
general effect jjaler and more greyish than in any of the other
European species ; underparts pale butfy grey, rather strongly
contrasted and with moderately well-defined line of demarcation
along sides of neck. Underfur slaty black. Muzzle and
cheeks faintly dusky. Ears and membranes an indefinite rather
Hght brown.

Skull. — In all resi^ects the skull so closely resembles that of
Myotls daubentonii that it is only distinguishable by its larger
size. From the skulls of M. nattereri and M. emarginatus it
differs in its greater breadth, a character perhaps most readily
appreciable in the form of the jjost-palatal region.

Teeth. — Except for their larger size the teeth I'esemble those
of Myotls daubentonii. The crown area of upper molars is
relatively less than in M. nattereri and 31. einanjinatus, though
»r retains the broader outline characteristic of M. daubentonii
and M. mystacinus. Upper molars with protoconule and its
accessory small commissure, and m^ and ?n'- with commissure
between hypocone and metacone as in jW. daubentonii.

Measureynents. — Average and extremes of forearms in four
males and eight females from Lugano, Ticino, Switzerland :
males, 39-9 (38-8-41); females, 39-4 (38-8-40-4). Sicily
(type): tibia, 15* 6 ; foot, 12 ; forearm, 39-2; thumb, 10 ; thii-d
finger, 56-2 ; fifth finger, 50. Adult male from Corleone, Sicily :
head and body, 49 ; tail, 38 ; tibia, 16 ; foot, 10 ; forearm, 41 ;
thumb, 8 • 2 ; third finger, 68 ; fifth finger, 55 : ear from meatus,
15 ; width of ear, 10 '4. Two adult males from Sassari, Sardinia :
head and body, 50 and 51 ; tail, 37 and 37 ; tibia, 16 '4 and 17 ;
foot, 10-6 and 11 ; forearm, 39-4 and 41*6 ; thumb, 8-4 and
8"6 ; third finger, 66 and 69 ; fifth finger, 51 and 56 ; ear from
meatus, — and 14*4; width of ear, 10 and 10. For cranial
measurements see Table, p. 191.

Specimens e.vamincd. — Seventy-two, from the following localities: —

Austria-Hungary: Herkulesbad, 1.

Switzerland : Near Lugano, 49 (B.M., U.S.N.M., and Mottaz).

Italy : Finalborgo, Liguria, 2 (Genoa) ; Pavia, 1 (U.S.M.M.) ; Ostia,
Rome, 2; Marsala, Sicily, 1; Corleone, Sicily, 1 (U.S.N.M.); Sicily, no
exact locality, 1 (type).

Sardinia : Cagliari, 4 (B.M. and U.S.N.M.) ; Sassari, 5 (U.S.N.M.) ;
Grotte de Sardale, 2.

France: Marseilles, 1 (U.S.N.M.).

Spain : Inca, Majorca, Balearic Islands, 1 ; Elche, Alicante, 1.

Itemarlis. — In general appearance this species resembles
Myotic daubentonii ; but it is immediately recognizable by its
even larger foot, and by the densely pubescent upper surface of
leg, the latter character unique among European bats.

4 6, 9. Near Lugano, Ticino, 0. Thomas (p). 2. 8. 4. 10-14.

Switzerland.

(E. H. Zullikofer.)
2. Ostia, Rome. Dr. L. Samhon (c&p). 1.1.2.3-4.

189

skeleton Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 734.

without parte.) {Type of species.)

skull.

9.

2al.
2al.

Marsala, Sicily. O. Thomas (p). 6. 8. 4. 21.

(.4. Robert.)

Cagliari, Sardinia. Florence Museum (e). 85. 7. 6. 6-7.

Grotte de Sardale, Sar- O. Thomas (p). 0. 12. 3. 1-2.

dinia.

Inca, Majorca, Balearic O. Thomas and R. I. 0. 7. 1. 3.

Islands. Poooek (c & p).

Elche, Alicante, Spain. G. S. Miller (c). 8. 8. 4. 13.

MYOTIS DASYCNEME Boie.

1823. Vcspertilio mijstaciniis Boie, Isis, p. 9G5. Not Y. viystacinus Kuhl

(Jutland, Denmark).
1825. Vespertilio dasycneme Boie, Isis, p. 1200 (Renaming of 7ni/slacinns

Boie).
1839. Vcspciiilio limnopldlus Temmiuck, Monogr. de Mamm., ii, p. 176

(Holland).
1857. Vespertilio dasycneme Blasius, Siiugethiere Deutschlands, p. 103.
1878. Vespertilio dasycnetne Dobson, Catal. Chiropt. Brit. Mus., p. 295.
1904. Myotis dasycneme Trouessart, Catal. Mamm. Tam viv. quam foss.,

suppl., p. 88.
1910. Myotis dasycnetne Trouessart, Faune Mamm. d'Europe, p. 27.

Type locality. — Chalk quarries at Dagbierg, near Wiborg,
Jutland, Denmark.

Geographical distribution. — Central and southern Europe,
west to the Atlantic coast,* north to Sweden, east into Asia.

Diagnosis. — Form and proportions essentialh' as in 3Iyotis
dauhentonii and M. capaccinii, but size much larger (forearm
about 47 mm., longest linger about 75 mm., condylobasal length
of skull about 16 mm.) ; tibia and adjacent membrane naked.

External characters. — Form not essentially different from the
other European members of the large-footed group. Tragus
relatively shorter than in any other European Mj/otis, its height
distinctly less than half that of conch, its anterior border slightly
concave, its posterior border slightly convex below, then more
abruptly convex to bluntly rounded ofi' tip, the two margins
essential!}' parallel through lower half. Free border of uro-
patagium without fringe.

Fur and colour. — Distribution of fur as in Myotis dauhentonii ;
quality not peculiar, the longest hairs on middle of back about
8 mm. in length. Colour of upper parts a light yellowish wood-
brown ; underparts strongly contrasted greyish white with a
tinge of but!', the line of demarcation along sides of neck well
defined. Muzzle and cheeks scarcely contrasted dusk\-. Under-
fur slaty black. Ears and membranes an indefinite dark brown.

* The supposed British record is probably erroneous (see Barrett-
Hamiltou, Hist. Brit. I\Iamm., i, pp. 157-158. December, 1910).

190

CHIROPTERA

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192 CHIROPTEEA

Skull. — The skull is large, sligbtly exceeding that of Myotis
hecltsteinii in length and noticeably surpassing it in breadth and
robustness. Its general appearance is the least typically Myotis-
like of any European member of the genus, a peculiarity
heightened by the crowding of the small premolars and conse-
quent shortening of anterior portion of tooth-row. Allowance
being made for the great difference in size the skull is not very
unlike that of Myotis dcmbentonii, but the rostrum is relatively
shorter and the brain-case broader and more depressed. Sagittal
crest barely indicated : auditory bull* proportionately as in the
smaller animal, less inflated than in 3/. bechsti'ini! ; lachrymal
ridge well developed, its lower extremity separated from lachrymal
foramen by an evident notch.

Teeth. ^In form the teeth resemble those of the small Myotis
dauhentonii ,■ molars decidedly more robust than those of M.
berlisteinii, the width of the crown diminishing much less rapidly
toward inner border ; small premolars much crowded, the second
barely or not visible from outer side, the diameter of its crown,
however, not much less than that of anterior tooth ; upper
molars as in M. dauhentonii and M. capaecinii ; mandibular teeth
with no special peculiarities, the premolars and outer cusps of
molars more slender than in M. hechsteinii and 31. dnuhentonii.

Measurements. — Two adult males from Maastricht, Holland :
head and body, 57 and 58 : tail, 49 and 51 ; tibia, 19-8 and
20-2; foot, 11-8 and 12 ; forearm, 43-6 and 44 ; thumb, 9-6
and 9 ; third finger, 72 and 77 ; fifth finger, 57 and 62 ; ear
from meatus, 17 and 17-2; width of ear, 10-6 and 11. Two
adult females from the same locality : head and body, 60 and 61 ;
tail, 47 and 46 ; tibia, 18 and 18 ; foot, 11-4 and 11 ; forearm,
44 and 44 ; thumb, 9 and 9 : third finger, 76 and 75 ; fifth
finger, 61 and 60 ; ear from meatus, 16-6 and 17 ; width of ear,
11 and 11. For cranial measurements .see Table, p. 191.

Specimens cxcanincd. — Six, from tlie following localities: —
Holland: Leyden, 1; Maastricht, 4 (U.S.N.M.).
Belgium : Near Namur, 1.

1. Leyden, Holland. Tomes Collection. 7. 1. 1. 501.

(H. Schlcgel.)
6 al. Namur, Belgium. Re%-. D. B. Lebbe 9. 1. 11. 1.

(c & p).

MYOTIS MYOTIS Borkhausen.

1775. Vespertilio murinus Schreber, Saugthiere, i, i>. 1G5 (Not of Linnfeus,

1758).
1797. Vespertilio myotis Borkhausen, Deutsche Fauna, i, p. 80 (Germany).
1797. Vespertilio myosotis B[orkhause]n, Der Zoologe (Compendiose

Bibliothek gemeinniitzigsten Kenntnisse fiir alle Stiinde, pt. xxi),

Heft v-viii, p. 46 (Germany).
1800. Vespertilio myosotis Bechstein, Pennant's Allgemeine Uebersicht der

Vierfiissigen Thiere, p. C32 (Germany).

MYOTIS 193

1801. Vespertilio viyotis Bechstein, Gemeinn. Naturgesch. Deutschl., i,

2ud ed., p. 1164 (Described but not named in 1st ed., 1789, x'. 1G4)

Thiiriugen, Germany.
1827. Vespertilio subvmrinus Brehm, Ornis, Heft iii, p. 24 (Renthendorf,

Tliiiringcn, Germany).
1844. V[espertilio] latipinnis Crespon, Faune Meridionale, i, p. 17 (Near

Nimes, Gard, France).
1857. Vespertilio murinus Blasius, Saugethiere Deutschlands, p. 82 (Not of

Linnteus, 1758).
1863. [Myotus tnurinus] var. typus Koch, Jahrb. des Vereins fur Natur-

liunde ini Herzogthum Nassau, xviii, p. 415 (Wiesbaden, Nassau,

Germany).
1863. [Myotus vmriuus] var. alpinus Koch, Jahrb. des Vereins fiir Natur-

kunde im Herzogthum Nassau, xviii, p. 415 (St. Gothard, Uri,

Switzerland).
1878. Vespertilio murinus Dobson, Catal. Chiropt. Brit. ]\Ius., p. 309 (Not

of Linnseus, 1758).
1886. Myotis murina var. spelxa Bielz, Verhandl. u. Mittheilungen des

Siebenbiirgischen Vereins fiir Naturwissensch. in Hermannstadt,

XXXVI, p. 83 (Homorod-Almas cave, Hungai-y).
1897. Myotis myotis Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 383,

October, 1897.

1909. Myotis myosoiis Miller, Ann. Mus. Zool. R. Univ. Napoli, N.S., iii,

No. 3, p. 1, April 26, 1909.

1910. Myotis myotis and M. m. spelasa Trouessart, Fauna Mamm. d'Europe,

p. 32.

Type locality. — Thliringen, Germany.

Geographical distribution. — Central and southern Continental
Europe, west to Portugal, north to southern Sweden, eastward
into Asia. One record of its occurrence in England."'

Diagnosis. — Largest species of European Myotis (forearm,
57 to 64 mm.; longest linger, 100 to 110 mm.; condylobasal
length of skull, 22 to 23 '6 mm.) ; form heavy, membranes thick
and leathery ; ear moderately long, extending about 5 mm.
beyond tip of muzzle when laid forward, its posterior margin
scarcely or not emarginate above middle ; foot slightly more than
half as long as tibia ; wing membrane extending to base of
outer toe.

External characters. — Although one of the largest European
bats Myotis myotis does not differ conspicuously in form from
the small M. mystacinus, except that its tail and legs are relatively
shorter. The general build is not I'emarkably heavy as compared
with other European species of approximately the same size, but
the ears and membranes are rather thick and leathery. Ear
moderately long, extending about 5 mm. beyond nostril when
laid forward ; anterior mai'gin moderately convex from base
nearly to rather narrowly rounded-off tip ; posterior border wath
shallow ill-defined concavity above ; antitragus low and long,
marked off posteriorly by a well-deHned notch and not continuous
with posterior border of conch ; tragus about half as high as

* Bell, Hist. British Quadrupeds, p. 88, 1836 : "But in England it . . .
has hitherto only been taken in the gardens of the British Museum."

O

194

CHIROPTERA

conch, its greatest width (slightly above level of antei'ior base)
contained about '2h times in length of anterior margin, the
posterior border convex to just below rather bluntly rounded tip,
the anterior margin straight, posterior basal lobe small but well
developed ; inner surface of conch with seven or eight ill-defined
transverse ridges near posterior border. Wing rather broad but
with no special peculiarities of form ; metacarpals somewhat
elongated relatively to jahalanges as compared with the smaller
species, the thii-d, fourth and fifth slightly but evidently
graduated, the third scarcely shorter than forearm ; membrane
inserted at side of metatarsus, but with a narrow strip extending
to base of outer toe as in M. nattereri. Foot slightly more than
half as long as tibia ; calcar heavy at base but tapering rapidly
and terminating obscurely, its posterior border with slightly
indicated keel, its length about equal to that of free border of
interfemoi-al membrane. Tail about as long as body without
head, only the extreme cartilaginous tip free from membrane.

Fur and colour. — Relatively to size of animal the fur is rather
short (longest hairs of back about 10 mm.); in distribution it
shows no peculiarities ; free border of uropatagium without
fringe. Colour an indefinite brown much like that of Mi/otis
nattereri, the exact shade intermediate between the wood-brown
and broccoli-brown of Ridgway, usually paler on head and neck
than on back, and in immature individuals than in adults, the
hairs slate-black through basal half, then light wood-brown
followed by a darker though not strongly contrasted terminal
area. Underparts strongly contrasted greyish white with a

slight bufly tinge ; a well-defined
line of demarcation along sides
of neck to ear, emphasized in
region of shoulder by a slight,
diffused blackening of edge of
dark area. Muzzle and cheeks
dusky. Ears and membranes an
indefinite brown.

Skull. — Though much larger
than that of any of the other
European .species of Myotu, M.
oxi/gnathus excepted, the skull of
Ml/of is myotis is one of the most
slender in general outline. . The
brain-case is longer proportion-
ately to its breadth and is less
contrasted with rostrum than in
the small members of the group ;
greatest breadth of brain-case
about one-half distance from
lambda to posterior margin of nares. Rostrum relatively deep
and interorbital concavity relatively shallow ; occipital region

Fig. 32
Myolis inyoth.

195

about on level with main portion of brain-case, and very slightly
overhanging foramen magnum ; \ antral profile scarcely elevated
posteriorly ; palate rather narrow (essentially as in M. mystaciiins) ;
width of posterior extension of palate less than its length, median
spine short but well developed ; posterior border of anteorbital
foramen over anterior root of first molar ; mandible with coronoid
process relativelv higher and narrower than in the small species,
its posterior border much more oblique.

Teeth. — The dentition is of a less primitive type than in the
small European species of Mijotls. This is indicated by the
general tendency to reduction shown especially in the lower
incisors, the small premolars and the posterior lower molar.
Upper incisors rather high and slender but not essentially
different in form from those of M. mystacinus, the cingulum of
inner tooth obsolete, that of outer slightly developed. Lower
incisors very strongly im-
bricated, the outline of the
row U-shaped or broadly
V-shaped ; cutting edge of
/j and /^ trifid, but decidedly
oblique owing to reduction
in size of outer cusp ; a
minute cingulum cusp usu-
ally present at extreme outer
edge ; i^ with small postero-
internal tubercle ; ^3 sub-
terete, 4-tuberculate, the me-
dian outer tubercle largest.
Canines relatively smaller
than in M. mystacinus and
with less developed posterior
cutting edge, their form
essentially as in M. nnttenri
and M. emarr/inatus. Small
upper premolars much
crowded, the second usually
forced inward from line of tootli-row, sometimes so much so that
the first is practically in contact with large premolar. In form
they show no special peculiarities ; shaft subterete ; cingulum
well developed. Large upper premolar more reduced than in the
small species, the inner margin of crown a mere cingulum at base
of main cusp, and without trace of secondarj' cusps. Lower
premolars not peculiar in form, but middle tooth usually more
crowded between first and third than in the small members
of the group. Molars .showing no special peculiarities of form ;
thickening that represents hypocone in iit^ and m- barely
indicated ; »(^ more reduced than in the small .species, its meta-
cone scarcely more than a slight widening of terminal portion of
third commissure. First and second lower molars essentially as

Fig.
Mijotis myotis.

196

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198

CHIROPTERA

in the small species, except that cingulum does not form a postero-
internal cusjj behind entoconid ; ^3 with hypoconid and ento-
conid more reduced than in the small species, the hypoconid
displaced further inward, so that second triangle is barely half
as large as first and conspicuously difierent from it in form.

Measurements. — Average and extremes of four adult females
from Tagerwilen, Thurgau, Switzerland: head and body, 75 '7
(72-79); tail, 56-3 (54-60); tibia, 26-1 (25 -4-26 -6); foot,
14-7 (13-16); forearm, 63 (63); thumb, 13-1 (12 -8-13 -4) ;
third finger, 107-7 (106-109); fifth finger, 85-2 (84-86); ear
from meatus, 27-6 (27-28) ; width of ear^lS-l (17-6-19). Two
adult females from Mte. Generoso, Ticino, Switzerland : head
and body, 72 and 76 ; tail, 50 and 51 ; tibia, 25 and 25 '4 ; foot,
15 and 14-4; forearm, 62 and 63-6 ; thumb, 13 and 12 ; third
finger, 104 and 107 ; fifth finger, 83 and 84 ; ear from meatus,
27 and 28; width of ear, 18 and 18-6. , Adult male from
Florence, Italy : head and body, 68 ; tail, 55 : tibia, 25 ; foot,
13; forearm, 61 ; thumb, 12; third finger, 100 ; fifth finger, 78 ;
ear from meatus, 27-6; width of ear, 17-6. For cranial
measurements see Table, p. 196.

Spec'uncns examined. — Ninety-five, from the following localities : —

France : Cadillac, Gironde, 1 (U.S.N.M.) ; Nimes, Gard, 2 (B.M. and
Nimes ; rexjresenting latiinnnis Crespon, but not type).

Germaky : Hamburg, 1 ; Niesiiy, Silesia, 11 ; Strass, near Burgheim,
Bavaria, 8 ; Heidelberg, Baden, 1.

Austria-Hungary : Herkulesbad, 1 ; Fiinfkircben, S.W. Hungary, 2.

RouMANiA : Bustenari, 3; Sinaia, 1 (U.S.N.JI.).

Switzerland : Geneva, 7 (Mottaz) ; Grotte de Vallorbe, Vaud, 1
(Mottaz) ; Boudry, Neucbatel, 2 (Mottaz) ; St. Moritz, 1 ; Thayngen, Schafif-
hausen, 2 (U.S.N.M.); Canton Thurgau, 5; Tiigerwilen, Thurgau, 10
(U.S.N.M.); Andermatt, Uri, 1 (U.S.N.M.); St. Gothard, Uri, 3 (B.M.
and U.S.N.M.) ; Jilonte Generoso, Ticiuo, 4 (U.S.N.M.).

Italy: Domodossola, 2 (U.S.N.M.); Fiualborgo, Liguria, 1 (Genoa);
Florence, 1 (U.S.N.M.) ; Rome, 2; Ostia, Rome, 2 ; ]Marsala, Sicily, G.

Sardinia : Oristano, Cagliari, 10.

Spain : Seville, 2.

Portugal: Cintra, 2.

Meniarhs. — Mijotis rityofis diflers strikingly from the other
European members of the genus, M. oxygnatJms excepted, in its
much larger size. From the large Vespertilionklse of other genera
it is immediately recognizable by its long ears, extending notice-
abl}' beyond nostril when laid forward, and by the greyish white
colour of the underpiarts. In the MediteiTanean region Myotis
myotis is associated with M. oxygnathus ; but it is the only large
species known to occur north of the Alps.

1 al.

Nimes, Gard, France.

G. E. Dobson (e).

80.

12. 14. 2.

1.

Hamburg, Germany.

Dr. J. E. Gray (p).

2 6, 8 V,

Niesky, Silesia. {Dr.

Dr. E. Hamilton (p).

97.

12. 4. 7-17.

9 juv.

W. Baer.)

0 9.

Strass, Burgheim, Ba-
varia. (Korbitz.)

Lord Lilford (p).

11.

1. 1. 10-15.

9.

Strass, Burgheim, Ba-

Lord Lilford (p).

11.

1. 1. 124.

varia. {Korbitg.)

11.

1. 1. 131.

MYOTIS

199

?. Heidelberg, Baden. Hon. N. C. Eoths- 10. 5. 29. 1.

child (p).

i juv. Herkulesbad, Hungary. Hon. W. Rothschild 7. 9. 16. 8.

[F. J. Cox.) (p).

2al. Fiintkirchen. BudapestMuseum(E). 94.7. 18. 11-12.

2 <J, 9. Bustenari, Prahova, Lord Lilford (p). 4.4.6.8-10.
840 m. Roumania.
(W. Dodson.)

1. St. Moritz.Grisons, Swit- Leon O. Galliard (p). 75. 9. 20. 4.

zerland.

5 ?, Thurgau. [E. H. Zolli- 0. Thomas (p). 4. 4. 5. 9-13.

kofcr.)

2 9. Rome. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. 26-27.

(!>)■

2. Ostia, Rome. Dr.L. Sambon(c&p). 1. 1. 2. 1-2.

6 9. INIarsala, Sicily. (.4. O. Thomas (p). 6. 8. 4. 17-20.

Bobcrt.) 8. 9. 1. .3-4.

10 al. Oristano, Cagliari, Sar- Hon. N. C. Roths- 7. 5. 24. 1-10.

dinia. child (p).

2 S al. Seville, Spain. (Dr. A. Lord Lilford (p). 73. 1. 8. 1-2.

Eitiz.)

2 i. Cintra, Portugal. O. Thomas (c & p). 98. 2. 2. 4-5.

MYOTIS OXYGNATHUS Monticelli.

1SS5. Vespcrtilio oxygnathus Monticelli, Ann. Accad. 0. Costa de Aspir.
Nat., I, p. 82. Type in Naples Museum.

1909. Mijotis oxygnathus Miller, Ann. Mus. Zool. R. Univ. Napoli, N.S.,

III, No. 3, p. 1, April 26, 1909.

1910. Myotis myotis oxygnathus Trouessart, Paune Mamm. d'Europe, p. 32.

Type locality. — Matera, Basilicata, Italy.

Geographical distribution. — Mediterranean region from Spain
to Greece, north to Italian vSwitzerland ; Sardinia ; Malta ;
Tunis.

Diagnosis. — Similar to Myotis myotis but smaller and with
shorter, narrower ears; condylobasal length of skull, 18 "6 to
21 • 4 instead of 22 to 23 • 6 mm. ; mandible, 15-2 to 17-2 instead
of 17 "8 to 19 mm.; maxillary tooth-row, 8' 2 to 9' 4 instead of
9-8 to 10-6 mm.

Measurements. — Type (adult male) : head and bod}', 63 ; tail,
54; tibia, 24-4; foot, 13; forearm, 57; thumb, 11-4; third
finger, 98 ; fifth finger, 76 ; ear 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, II and 11 "8 ; third finger, 93 and 97 ; fifth
finger, 74 and 78 ; ear from meatus, 23 and 22 ; width of ear,
13-6 and 13 "6. Adult male and female from Bozen, Tirol : head
and body, 68 and 71 ; tail, 53 and 53 ; tibia, 25 "4 and 24 ; foot,

200

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202

CHIROPTEEA

14 and 14 ; forearm, 59 and 60 ; thumb, 11 "4 and 11 -6 ; third
finger, 97 and 98 ; fifth finger, 76 and 78 ; ear from meatus, 26
and 25 ; width of ear. 17 and 15. For cranial measurements see
Table, p. 200.

Specimens examined. — Seventy-four, from the following localities : —

Spain : Near Burgos, 2.

Switzerland : Lugano, Ticino, 1 (U.S.N.M.).

AuSTBiA-HuNGARY : Bozeo, Tirol, 3 (U.S.N.M.).

Italy : Finalborgo, Liguria, 2 (Genoa) ; Isoverde, 1 (Genoa) : Vallom-
brosa, Florence, 1 (U.S.N.M.) ; Rome, 4; Velletri, Rome, 15 (U.S.N. M.) ;
ilatera, 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.JI.).

Malta : El Ghain, 5 ; Rubato, 3 ; no exact locality, 4.

Montenegro : Beri, 4.

CtREECe: Patras, 2; Corinth, 7 (U.S.N.M.); Nauplia, 1 (U.S.N.M.);
Lamia, Thcssaly, 1 (U.S.N.M.)

Crete : Labyrinth, 4.

Tunis: No exact locality, 3 (U.S.N.M.).

JReniarkn. — In general appearance Myotis oxygnathus resembles
M. miiofis, though the colour perhaps averages somewhat darker.
It is readily distinguishable, however, by its smaller skull
(distinctl}^ smaller head in spirit specimens), and shorter, narrower
ears. In form the skull is like that of 31. vu/otis ; and the
teeth are not peculiar except for their small size, a character
readily ajjpreciable 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 M. capaccinci and PipistreUus Imldii. North of the Alps
M. jiiyotis occurs alone.

?.

Burgos, Spain.

G. S. MUler (c).

8. 8. 4. 10.

4 i.

Rome, Italy. (Coli.)

G. Barrett-Hamilton

11. 1. 2. 22-25

3 6, 2 ?.

El Ghain, jNIalta.

Lord Lilford (p.)

11. 1. 1. 16-20

3 6, 9.

iMicallef.)
Rubato, Malta.

Lord Lilford (p).

11. 1. 1. 21-24

4.

2 (5, 2 9.

(Micallcf.)
Malta. (Micallef.)
Beri, 50 m. Montenegro.

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

95. 3. 2. 3-6.

5. 8. 4. 1-4.

2 9.

(L. Filhrer.)
Patras, Greece.

Hon. N. C. Roths-

S. 10. 2. 15-16

3 6, 9.

(C. Mottaz.)
Labyrinth, Crete.

child (p).
Miss D. Bate (c).

5. 12. 2. 5-8.

Genus PIPISTRELLUS Kaup.

1829. Pipistrellus Kaup, Enlw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt,
I, p. 98 [pipistrellus).

1837. Pipistrellus Bonaparte, Iconogr. Faun. Ital., i, fasc. xx.

1838. Bomicia Gray, Mag. Zool. and Bot., ii, p. 495, February, 1838

[calcarata = kulilii).

1839. Vespcrugo Kevserling and Blasius, Wiegmann's Archiv ftir Natur-

gesch., 1839, i, p. 312 (part).
1856. Hypsugo Kolenati, Allgein. deutsche Naturhist. Zeitung, Dresden,
neue Folge, ii, p. 131 (maurus and kraschcninikoivii).

PIPISTRELLUS 203

1S56. Nannugo Kolenati, Allgcm. deutsche Naturhist. Zeitung, Dresden

neue Polge, ii, p. 131 {nathusii, pipistrelhis, and kuhlii).
1857. Vesperugo Blasius, Saugethiere Deutschlands, p. 49 (part).
1878. Vesperugo Dobson, Catal. Chiropt. Brit. Mus., p. 183 (part).
1897. Pipistrellm Miller, Ann. and IMag. Nat. Hist., 6th ser , sx n 384

October, 1897. '

1899. Euvespemgo Acloque, Fauna de Prance, Mammiferes, p. 35 (part

included noctula, leisleri, maurus, kuhlii, pipistrellus, and

aoramusj.

1907. Piiyistrellus Miller, Families and Genera of Bats, p. 204 June 29
1907. ^ '

Tjipe speciefi.— Vespertilio pipistrelhts Schreber.

Geographical distribution.— Entire mainland of Eastern Hemi-
sphere to hmit.s of tree growth, also Malay Archipelai(o, New
Guinea, Solomon Islands, and northern Australia ; in America
trom_ northern United States (except in boreal zone) to southern
Mexico.

Characters.— Like Eptesicus (p. 224), but with 2-2 upper
premolars ; dental formula : i |f, c U, pm ||, m •"-■' = 34.

BemarTcs.—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
^urope. Externally these may often be confused with the smaller
Myotis, though they may usually be recognized by a certain
heaviness of form, and more especially by the shorter ear and
less slender tragus. Though often regarded as nearly related to
JSyctalus, on account of the similarity of dental formula, this
genus IS much the more primitive of the two, its members
showing no tendency to modify the ordinary vespertilionine wing
structure. It is in reality not much more than a sub-genus of
±.ptesicus (see remarks under the latter), though for the sake of
convenience the two groups are best treated as distinct.

KEY TO THE EUEOPEAN SPECIES OF PIPISTRELLUS.

Anterior upper premolar excessively minute, some-
tunes 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

. , *'.P^ P. savii, p. 219.

Anterior upper premolar not excessively minute,

never hidden by the gum, its crowu a'rea about

equal to that of outer incisor; anterior lower

premolar with crown area more than half that

of posterior premolar ; greatest width of tragus

much less than length of anterior border ; hairs

of back without noticeably contrasted light tips.

Outer upper incisor less than half as high as

inner ; large upper premolar almost or quite

m contact with canine, the small premolar

forced inwardfrom tooth-row and scarcelv or

not visible from outer side " p. kuhlii, p 215

204

CHIROI'TEEA

Outer upper incisor more than half as high as

inner ; large upper premolar separated from

canine by distinct space in which the small

premolar is clearly visible from outer side.

Lower canine robust, the length of base along

cingulum about equal to length of anterior

border of shaft ; 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. Yespertiiio xiiiyistrdlus Schreber, Saugthiere, i, pi. Liv. Described, i,

p. 167, 1775, under name Die Zwergfledermaus. (France, based

primarily on Daubenton.)
1776. yespertiiio pipistrcUe P. L. S. IMiiller, Natursyst. Suppl. u. Regist.-

Band, p. 16 (France, based on Schreber).
1825. Yespertiiio pugmieus Leach, Zool. Journ., i, p. 560, January, 1825

(Dartmoor, Devonshire, England).
1834. Vespertilio brachyotos Baillon, Mem. Soc. Royale d'Emulation

d'Abbeville, 1833, p. 50 (Abbeville, Somme, France).
1839. ? [Vespiertilio pipistrellus'] var. nigra de S61ys-Longchamps, Etudes

de Micromamm., p. 140 (nomen nudum).

1839. ? [Yespertiiio pipistrellus'] var. rnfescens de S61ys-Longchamps, Etudes

de Micromamm., p. 140 (nomen nudum).

1840. Y[esp)ertilio] pusillus Schinz, Europ. Fauna, i, p. 9 (Synonym of

pipistrellus; Brehm cited as authority).

1840. Y[espertilio] melanopterus Schinz, Europ. Fauna, i, p. 9. Brehm
cited as authority, but name apparently published here for first
time (Rhentendorf, Tliiiriugen, Germany).

1840. Y\_cspertilio] stenotus Schinz, Europ. Fauna, i, p. 9. Brehm cited as
authority, but name apparently publislied here for first time
(Rhentendorf, Thiiriugen, Germany).

1840. Vespertilio mintitissimus Schinz, Europ. Fauna, i, p. 9 (Ziirich,
Switzerland).

1845. P[ipistrellus'] nigricans Bonaparte, Atti dclla sesta Riunione degli
Scienziati Italiani, Milano, 1844, p. 340. Described but not named
in Atti della seconda Riunione degli Scienziati Italiani, Torino,
1840, p. 247, 1841. (Sardinia.)

1845. Pipistrellus genci Bonaparte, Atti della sesta Riunione degli Scien-
ziati Italiani, Milano, 1844, p. 340 (Alternative name for nigricans).

1845. P[ipistrcUus] typus Bonaparte, Atti della sesta Riunione degli Scien-
ziati Italiani, ililano, 1844, p. 340 (Substitute for pipistrellus
Schreber).

1867. Yesperugo pipistrellus Blasius, Siiugethiere Deutschlands, p. 61.

1862. Yesperugo pipistrellus var. macropterus Jeitteles, Verhandl. der k. k.

Zool. Bot. Gesellsch., Wien, xii, p. 250 (Kaschau, Hungary).

1863. \_Nannugo pipistrellus] var. typjis Koch, Jahrb. des Vereins fiir

Naturkunde im Herzogthum Nassau, xviii, p. 490. Not of
Bonaparte, 1845 (Wiesbaden, Hessen-Nassau, Germany).

PIPISTRELLDS 205

1863. [Nannugo pipistrellus] var. flavcscens Koch, Jahrb. des Vereins fiir

Naturkunde im Herzogthum Nassau, xviii, p. 491 (Nassau,

Germany).
1863. \_Nannugo piiiistrelhis] var. nigricans Koch, Jahrb. des Vereins fiir

Naturkunde im Herzogthum Nassau, xviii, p. 491. Not of

Bonaparte, 1845 (Nassau, Germany).
1868. {Nannngo pipistrellns] var. limbafiis Koch, Jahrb. des Vereins fiir

Naturkunde im Herzogthum Nassau, xviii, p. 491 (Siegen, Nassau,

Germany).

1878. Vcsjierugo jnpistreUus Dobson, Catal. Chiropt. Brit. Mus., p. 223.
1897. Pipistrellns pipistrcllus Miller, Ann. and Mag. Nat. Hist., 6th ser.,

XX, p. 384, October, 1897.
1904. Pipistrellus pipistrcllus mediterraneus Cabrera, Mem. Soc. Espaii.

Hist. Nat., II, p. 273 (Valencia, Spain).
1910. Pipistrellus piipistreUus and P . pipistrellus mediterraneus Trouessart,

Faune Mamm. d'Europe, pp. 14-15.

Type locality. — France.

Geographical distribution. — Europe from the Mediterranean
north to Scotland and Scandinavia, west to Ireland and the
Hebrides, east into Asia.

Diagnosis. — Smallest European member of the genus (forearm,
27-6 to 32 mm. ; condylobasal length of skull, 11 to 12 mm.);
outer upper incisor more than half as high as inner incisor ;
large upper premolar separated from canine by a distinct space,
the small tooth visible from outer side, its crown area about
equal to that of outer incisor ; anterior lower premolar with
crown area equal to more than half that of succeeding tooth ;
lower canine robust, the length of base along cingulum about
equal to length of anterior border of shaft ; tragus with greatest
width less than length of anterior border ; thumb short, its
length about equal to width of wrist ; length of fifth finger about
40 mm. ; posterior edge of wing membi-ane usually dark.

External characters. — General form robust, the tail and legs
rather short, the membranes relatively thick and opaque. Muzzle
with very noticeable glandular swellings extending back to
beneath eye. Ear extending about to nostril when laid forward,
its general form rather short and broad, though with narrowly
rounded tip ; anterior border abruptly convex at base, then
essentially straight almost to tip ; posterior border faintly and
irregularly concave above, evenly convex below, the antitragal
lobe represented by a thickened ridge extending along margin of
ear and turning abruptly inward without producing any notice-
able break in outline of conch. Inner surface of conch slightly
rugose, but without evident cross ridges. Tragus erect, scarcely
half as high as conch, its tip broadly rounded, its greatest width
(slightly above level of anterior base) about half length of anterior
border ; except near tip, both borders are nearlv straight or very
slightly convex ; posterior basal lobe small but well defined.
Wing rather nari'ow, with no special peculiarities of form, the third,
fourth and fifth metacarpals sub-equal (fifth slightly shorter than

206 CHIROPTERA

the others) and extending nearly to point of elbow ; fifth finger
extending beyond elbow to a distance equal to less than one-
third length of forearm ; thumb short, its length about equal to
width of wrist ; membrane inserted at base of outer toe. Foot
about half as long as the short, robust tibia ; calcar considerably
longer than free border of interfemoral membrane, robust at base,
but tajjering rapidly and terminating without lobe, its keel well
developed, with evenly convex margin. Tail about as long as
body without head and 2J times as long as tibia, the short
terminal vertebra free from membrane.

Fa7- and coluur. — The fur is closely confined to the bod}^
showing no tendency to spread on membranes. On wing it
extends, both above and below, to line joining knee and basal
third of humerus ; lower surface of interfemoral membrane
essentially naked except at extreme base, upper surface furred
nearly to middle. Colour of upper parts a uniform brown, in
most specimens nearly intermediate between the wood-bi"own and
cinnamon of Rid way but sometimes darker, with a strong tinge
of jirouts-brown or raw umber, this especially noticeable in
immature specimens, though occasionally evident in adults ;
under parts essentially like back though slightly less dark : hairs
everywhere slaty brown at base, those of upper parts with tips
darker than sub-terminal band, but not enough so to produce a
definitely tricolor effect. Ears and membranes blackish.

Skull. — Notwithstanding its small size, less than that of any
other European bat, the skull is robust and heavily built as
compared with that of the small species of Myotis. Dorsal profile
rising gradually from nares to lambda, with slight concavity in
interorbital region and slight convexity over middle of brain-case ;
occipital region scarcely produced backward except for a median
swelling between foramen magnum and lambda,
on each side of which a condyle is just visible
when skull is viewed from above ; ventral profile
nearly flat except for a slight upward bend pos-
teriorly. Brain-case ovate in general outline,
its region of greatest breadth distinctly beliind
middle, its surface smooth or with faintly indi-
cated sagittal crest and lateral portion of
lambdoid crest ; greatest breadth of brain-case
noticeably exceeding that of rostrum and slightly
Piput^l^'pivis- though evidently more than half greatest length
treiius. Nat. size, of skuU ; floor of brain-casc flat, without vacui-
ties ; a distinct groove between cochlea and
median portion of floor, this groove bounded antero-externally
by a slight though usually evident longitudinal ridge ; auditory
bulla' moderately large, not peculiar in form ; interorbital region
l^roadly hour-glass shaped, its least breadth about equal to
breadth across roots of canines ; between constriction and ante-
orbital foramen the orbital margin is slightly but evidently

PIPISTRELLUS

207

inflated, the inflated I'egion with a median angle suggesting
a rudimentary poster bital process ; rostrum short and broad,
narrowing gradually in fi'ont, a sliglit concavity at each side
bortlering lachrymal inflation, and an evident median longi-
tudinal groove, most noticeable posteriorly ; nasal emargina-
tiou slightly deeper than wide, extending less than half way to
interorbital constriction ; anteorbital foramen small, over point of
contact between large premolar and first molar ; palate broad,
distinctly concave both longitudinally and laterally ; anterior
emai'gination small, wider than deep, its posterior border on line
with posterior edge of canine ; mesopterygoid fossa squarish,
encroached on anteriorly by broadly triangular median palatal
spine ; hamulars slightly turned inward. Mandible robust, the
ramus much deeper at symjjhysis than behind tooth-row, the
coronoid process so low that upper edge of posterior portion of
mandible is squarely and horizontally truncate, parallel with
alveolar line ; angular process short but well developed, on level
with alveolar line, its extremity slightly bent inward.

Teeth. — Relatively to size of skull the teeth are rather large
and robust, though inclined to be low, tendencies especially
noticeable in the canines. Inner upper incisor robust, its shaft
nearly half as high as that of canine, and directed strongly
forward and slightly inward, its crown iiTegularly elliptical-oval
in outline, with main axis nearly in line of tooth-row ; secondary
cusp large and conspicuous, about half as high as main shaft,
from the postero-external surface of which it projects ; cingulum
well developed, often forming a minute postero-basal cusp. Outer
upper incisor slightly but evidently smaller than inner, its shaft
more than half as high as that of inner, to secondary cusp of
which its extremity is closely approximated ; crown outline
essentially as in inner tooth but main axis lying at right angles
to tooth-row ; posterior surface of shaft broadly concave ; inner
margin with small though distinct secondary cusp ; cingulum
moderately well developed. The main cusps of the two teeth lie
in line of general curve of anterior portion of tooth-row. Space
between outer incisor and canine about equal to greatest diameter
of incisor. Lower incisors forming a continuous, broadly (J'shaped
row between canines, their crowns very slightly imbricated ;
crowns much longer than high, trifid, that of i■^ narrowest,
longest and lowest, that of /._, and (3 widened posteriorly but
without additional cusps or tubercles. Upper canine robust,
the greatest diameter of its crown about three-quarters length
of anterior border of shaft, the cross section of shaft broadly
triangular with longest side formed by nearly flat postero-internal
surface ; a sharply defined antero-exteral longitudinal groove, and
less definite postero-external concavity ; anterior edge narrow-
but not strictly trenchant ; posterior edge trenchant, with well
marked angle slightly below middle, this angle frequently becom-
ing a distinct secondary cusp ; cingulum well developed but not

CHIROPTERA

forming true basal cusps. Mandibular canine low and heavy,
its apex scarcely rising above level of highest molar cusps, its
greatest diameter measured along cingulum nearly or quite equal
to length of anterior border of shaft ; cingulum well developed,
foi'ming a distinct antero-basal cusp, the apex of which rises to
level of middle of posterior border of shaft. Anterior upper
premolar with area of crown approximately equal to that of upper
incisor and about one-hfth that of canine. It is somewhat
crowded inward from tooth-row, though about half of its crown is
visible from outer side in space between canine and large
premolar ; main cusp short but well developed, lying somewhat
in front of middle of crown, the general form of the tooth much
like that of canine but proportionally lower. Large upper
premolar with crown area about equal to that
of canine or slightly greater, the inner portion
narrow and flattened-concave, with evident
elevated rim, the posterior border strongly
concave, the anterior border usually convex
' but occasionally a little concave ; height of
main cusp slightly greater than that of highest
Fig. 35. molar cusps and about equal to length of

AnterioTteeth X 5"* tooth along outer cingulum, posterior cutting
edge AVell developed ; secondary cusp low but
evident, rising from cingulum at antero-internal base of main
cusp. Lower premolars with crown areas not conspicuously
unequal, though that of second perceptibly greater than that of
first ; outline of crown of each tooth rhombic, the outer border
somewhat convex, the anterior border of second relatively shorter
than that of first ; cingulum well developed, forming a slight
antero-internal basal cusp ; main cusp triangular in outline when
viewed from the side, that of second as high as molar cusps, that
of first shorter, the antero-external surface of each tooth convex,
the internal and posterior surfaces concave. First and second
upper molars sub-equal, though transverse diameter is relatively
greater in latter than in former ; inner border rather narrowly
rounded, the region of greatest convexity a little in front of
middle ; anterior and posterior borders straight or slightly
concave ; protocone robust though rather low ; hypocone small
but well developed, though not completely distinct from posterior
commissure of protocone ; metacoue higher than paraccme ; styles
well developed ; VV'P^ttern normal ; in'^ with crown area about
two-thirds that of vi^, the hypocone absent, the metacone smaller
than paracone ; no trace of metastyle or fourth commissure.
Lower molars with no special peculiarities ; protoconid higher
than liypoconid in all thi'ee teeth ; hypoconid with greater basal
area than protoconid in j/^j and m.^, but with less in m.^ ; cingulum
well developed, forming a slight posterointernal cusp behind
entoconid.

Measureinenfs. — Adult male from Henley-on-Thames, Oxford-

PIPISTRELLUS

209

shire, England : hccad and body, 44 ; tail, 32 ; tibia, 10-4 ; foot,
6 ; forearm, ."50 -4 ; thumb, 4-4 ; third finger, 52 ; fifth finger,
.■}8. Two adult males from Horrento, Italy : head and body, 42
and 43 ; tail, 29 and 32 ; tibia, 10 -8 and 10 ; foot, 5-8 and 6-2 ;
forearm, 30-4 and 31 ; thumb, 4-8 and 4*2; third finger, 53
and 53 ; fifth finger, 39 and 38 ; ear from meatus, 11*4 and
1 1 • 4 ; width of ear, 8 • 2 and 8 • 2. Two adult females from the
same locality : head and body, 39 and 40 ; tail 32 and 33 ; tibia,
10 "8 and 10-6 ; foot, 6 and 6 ; forearm, 32 and 30-2 ; thumb,
5 and 4*4 ; third finger, 58 and 52 ; fifth finger, 41 '6 and 40 ;
ear from meatus, 1 2 and 1 1 " 2 ; width of ear, 8 • 4 and 8. Adult
female from l^urgos, Spain, and adult female from Ciudad Real,
Spain : head and body, 49 and 39 ; tail, 31 and 30 ; tibia, 10 '6
and 9-6 ; foot, 4-8 and 5-6 ; forearm, 32 and 28-8 ; thumb, 5
and 4-6 ; third finger, 57*6 and 51 ; fifth finger, 42 and 38 ; ear
from meatus, 10 '4 and 10 '4. Extremes of twenty males from
Florence, Italy : head and body, 33-38 ; tail, 26-31 ; tibia, 9'2-
9-6; foot, 5 • 0-5 • 2 ; forearm, 27 ■ 6-30 ; thumb, 5 • 0-5 • 0 ; third
finger, 49-53 ; fifth finger, 34 ■ 6-40. For cranial measurements
see Table, p. 210.

Speciviens examined. — Two hundred and seventy-nine, from the following
localities : —

Ireland : Co. Longford, 1 ; Co. Antrim, 1.

England : Alnwick, Northumberland, 1 ; Bowdon, Cheshire, 1 ; Great
Grimsby, Lincolnshire, 1 ; Henley-on-Thames, Oxfordshire, 1 (U.S.N.M.) ;
Tring, Hertfordshire, 4 : Lilford, Northamptonshire, 2 ; Aberia, Merioneth-
shire, 1; Chelmsford, Essex, 2; London, 1; Wimbledon, Surrey, 1; Twig-
worth, Gloucestershire, 1 ; New Forest, Hampshire, 1 ; Netley, Hamp-
shire, 1 ; Loddiswell, Devonshire, 1.

Sweden : Upsala, 6.

Denmark : Hillerod, Zealand, 1.

France : Boulogne -sur-Mer, Pas-de-Calais, 2 ; Etupes, Doubs, 1
(Mottaz) ; Nimes, Gard, 3 (Mottaz and Nimes, the last wrongly marked
type of nigrans Crespon) ; St. Geuies, Gard, 1 (Mottaz) ; Marseilles, 1
(U.S.N.M.)

Germany : Bonn, 4 ; Ingelhcim, Rheinhessen, 1 ; ^Magdeburg, Saxony, 1 ;
Berlin, 1 ; Rudolstadt, Bavaria, 3 ; Niesky, Silesia, 1.

Austria-Hungary : Transylvania, 1 ; Zara, Dalmatia, 1.

Switzerland : Geneva, 15 (ilottaz) ; Buchillon, Vaud, 4 (Mottaz) ;
Morat, Fribourg, 1 (Mottaz) ; Neuchatel, 1 (Mottaz) ; Cortivallo, Ticino,
1 (U.S.N.M.); Mt. San Salvatore, Ticino, 13 (U.S.N.M.).

Italy: Campiglio, Tirol, 1; near GVenoa, 10 (U.S.N.M. and Genoa);
Isola Giglio, 2 (CJenoa) ; Florence, 103 (U.S.N.M. and Mottaz) ; Rome, 2
(U.S.N.M.); Sorrento, 21 (U.S.N.M.); Mondulo, Sicily, 1 (U.S.N.M.);
Palermo, Sicily, 1 (U.S.N.M.) ; Sicily, no exact localitv, 1 ; Ustica Island, 1
(U.S.N.M.). ^ ^ ^-

Sardinia : Cagliari, 17 (B.IM. and Genoa) ; no exact locality, 2.

Greece: Athens, 1 (U.S.N.M.); Tatoi, near Athens, 11; Kephissia,
near Athens, 4.

Spain : Villalba, Lugo, 1 ; Burgos, 3 ; Silos, Burgos, 2 ; La Granja,
Segovia, 4; Ciudad Real, Madrid, 2; Granada, 1; Seville, 1; Aloala, near
Seville, 1.

Jiemarl's. — PipigtrelluH pipisttellns is the smallest as well as
one of the commonest and most generally distributed of European

p

210

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212

CHIROPTEEA

bats. Superficially it may be distinguished from the almost
equally small Mijotis miistacinua by its smaller ears and shorter
legs ; but for positive discrimination from the members of the
genus Pijjiatrellus recourse to the more technical characters of
skull and teeth is necessary. On superficial examination it
may usually be recognized among its congeners by its small size,
and by the shortness of the fifth finger.

<5al.

Co. Longford, Ireland.

Dr. G. E. Dobson (p).

76. 2. 12. 1.

9 St.

Co. Antrim.

Hon. N. C. Rothschild

P)-

1. 9. 3. 6.

?.

Alnwick, Northum-
berland, England.

W. E.de Winton (p).

11. 1. 3. 390.

2<5, 2 9.

Tring, Hertfordshire.

Hon. N. C. Rothschild

P)-

9. 2. 19. 1-4.

9.

Lilford, Northampton
shire.

Lord Lilford (p).

11. 1. 1. 125.

2 9.

Chelmsford, Essex.

M. Christy and E.
Thompson (p).

L.

11. 1. 3. 23-24.

9 juv. al.

London.

Dr. A. Giinther (p).

74. 7. 6. 1.

i.

Wimbledon, Surrey.

C. H. B. Grant (c & p)

11. 1. -6. 27.

juv.

New Forest, Hamp-
shire.

Col. J. W. Yerbury (c &

'p)-

11. 1. 3. 389.

9al.

Netley, Hampshire.

Dr. G. E. Dobson (c &

P)-

76. 11. 3. 1.

9.

Loddiswell, Devon-
shire.

Col. J. W. Yerbury (c &

P)-

11. 1. 3. 25.

3 9.

Up sal a, Sweden.
(Kolthoff.)

Lord Lilford (p).

11. 1. 1. 26-28.

S.

Hillerod, Zealand,
Denmark.

0. Thomas (c & p).

96. 6. 7. 1.

6, 9.

Boulogne, Pas - de -
Calais, Franco.

0. Thomas (c & p).

98. 1. 9. 1-2.

6, 9 al

Bonn, Rhineland,
Germany.

Dr. A. CUinther (p).

9.

Ingelheim, Rhein-
hessen.

C. H. Hilgert (c).

8. 11. 2. 3-4.

9al.

JMagdeburg, Saxony.

Dr. W. WolterstorfE (p

).

92. 12. 1. 2.

Sal.

Berlin.

Dr. A. Giinther (p).

66. 2. L22.

9 al.

Transylvania, Hun-
gary.

G. G. Danford and J.
Brown (c & p).

A.

74. 7. 4. 6.

6.

Zara, Dalmatia.

Lord Lilford (p).

11. 1. 1. 29.

2 al.

Ciudad Real, Spain.

X. Cabrera (p).

8. 7. 23. 2-3.

9.

Burgos, Prov. Burgos

. G. S. Miller (c).

8. 8. 4. 14.

9.

Silos, Burgos.

Ct. S. Miller (c).

8. 8. 4. 15.

1 al.

Campiglio, Tirol,

G. C. Champion (c & p).

96. 8. 7. 1.

Italy.

i juv. al
14 al.

Sicily.

Aristano, Cagliari,

46. 6. 15. P.
7. 5. 24. 11-24

Hon. N. C. Rothschild

(P)

Sardinia. (C.

Erausse.)

2 juv. al

Sardinia. (P. Bonomi.

) E. N. Buxton (p).

95. 4. 16. 4-5.

3 9.

Tatoi, Athens, Gi-reece
(C. Mottaz.)

Hon.N. C.Rothschild

(P)-

8. 10. 2. 19-21

6, 3 9.

Kephissia, Athens.

C. Mottaz (c).

8. 11. 3. 4-7.

PIPISTRELLUS 213

PIPISTRELLUS NATHUSII Ke3'.serling and Blasius.

1839. V[espciiilio] naflMsii Keyserling and Blasius, Wiegmann's Archiv

fiir Naturgesch., 1839, i, p. 320 (Berlin, Germany).
1857. Vesperngo nathusli Blasius, Saugethiere Deutschlands, p. 58.
1878. Vesperngo abranms Dobson, Catal. Chiropt. Brit. Mus., p. 226 (Part :

not of Temminck).
1900. Pipistrellus nathusii M6hely, Monogr. Chiropt. Hungariae, p. 276.
1905. Vesp[eritgo] nathusii var. unicolor Fatio, Arch. Sci. Phys. et Nat.,

Geneve, 4th ser., six, p. 510, May, 1905 (Geneva, Switzerland).

Type in Geneva Jluseum.
1910. Pipistrellus ahramus Trouessart, Faune Mamm. d'Europe, p. 16.

Type locality. — Berlin, Germany.

Geographical distribution. — Central and southern Continental
Europe ; exact limits of range not known.

Diagnosis. — Not so small as PipistreUus lApistrellus (forearm,
32 to 3-5 mm. ; condylobasal length of .skull, 12 '6 to 13 "4 mm.),
which in general it resembles, but : small upper premolar better
developed, the greatest diameter of its crown nearly half that of
canine ; canines both above and below much more slender, the
length of base of lower tooth measured along cingulum slightly
more than half length of anterior border ; tragus more slender,
its greatest width much less than length of anterior border ;
thumb long, its length much greater than width of wrist ; length
of fifth finger about 46 mm. ; posterior edge of wing membrane
always pale, though never sharply defined white.

E.dernal characters. — In general and apart from the animal's
less diminutive size, the external characters are essentially as in
Plj}i.^treUus pipistrellus. Ear larger and broader with more
obtuse apex, more evadently concave posterior border, the inner
surface of conch more rugose and with about four irregular cross
striations behind tragus ; antitragus small but well defined,
projecting distinctly beyond border of conch ; tragus about as
high as in P. pipistrellus and similarly blunt . at tip, but with
posterior border more evidently convex ; posterior basal lobe
small, usually le.ss well defined than in P. 2}!pistreUus. Wing
larger and relatively broader than in P. pipistrellus, the meta-
carpals as in the smaller animal, but fifth finger extending beyond
elbow to a distance equal to decidedly more than one-third length
of forearm ; thumb less shortened than in the other European
members of the genus, its length noticeably greater than width
of wrist ; membrane inserted at base of outer toe. Foot, calcar
and tail as in P. pipistrellus.

Fur and colour. — Fur slightly more loose in texture than that
of P. jripistrelhis, the individual hairs somewhat longer, those at
middle of back about 7 mm. in length. In distribution it shows
no peculiarities, though it extends perhaps less widely on dorsal
surface of interfemoral membrane. Colour essentially like that
of Pipistrellus pipistrellus, though usually distinguishable by a

214

CHIROPTERA

tendency aAvay from the cinnamon and raw-umber tints toward
a clearer brown more resembling Eidgway's mars-brown. Mem-
branes less blackish than in P. pipistrdlus, the wing from foot
nearly to fifth finger with a noticeable pale border about 1 mm.
in width, similar to that present in P. kultlii, but less sharply
defined and less nearly white.

Skull. — The skull is less diminutive than that of Pipistrellus
pipistrellus, its general size about as Myotis mystnciuus. General
form less robust than in P. pijristreUus, the width of brain-case
barely one-half greatest length, but more contrasted with that of
rostrum. Dorsal profile as in the smaller sjaecies, but with more
evident anterior concavity and posterior convexity, the anterior
edge of interparietal indicated bj'' a slight transverse constriction.
Other details of form essentially as in P. pipistrellus.

7'eetJi. — As compared with those of P'qnstrelhts pipistrellus the
teeth throughout show a tendency toward slenderness and height.
Inner upper incisor noticeabh' more slender than that of P.
pipistrellus, and with less well developed secondary cusp ; outer
upper incisor distinctly larger than inner, its apex extending
noticeably beyond secondary cusp of inner tooth, its general form
essentially as in P. pipistrellus, but inner margin without evident
secondary cusp. Lower incisors less crowded than in P. j^iipia-
trdlus, a slight space usually present in
median line, another between /^ and i^, and
another between i^ and canine ; outer edge
of /j slightly ovei'lapping i.^ ; in form the
teeth are not peculiar. Upper canine like
that of P. piinstreUus, except that the
greatest diameter of its crown is only about
half length of anterior border of shaft.
Mandibular canine high and slender, its apex
Pipistrcllus nathusii. rising distinctly above that of highest molar
Anterior teeth x 5. cusps, its greatest diameter measured along
cingulum equal to a little more than half
length of anterior border : apex of anterior cingulum cusp not
rising above level of basal third of posterior boi-der. LIpper
premolars as in P. pipistrelhis, except that the small tooth is
relatively higher and more perfectly in the tooth-row, and the
posterior border of its shaft usually shows some indication of an
angular secondary cusp corresponding to that of canine. Lower
premolars with crown area more nearly equal than in the
smaller species, but without special peculiarities of form. Molars
both above and below essentially similar to those of P. piplstreUus.
Measurements. — Adult male from Berlin, Germany (topotype) :
head and body, 45 ; tail, 35-4 ; tibia, 13 ; foot, 6*8 ; forearm,
33; thumb, 5*2; third finger, 65; fifth finger, 47; ear from
meatus, 12; width of ear, 11. Average and extremes of six
adults from Buchillon, Vaud, Switzerland : tibia, 1 2 • 9 ( 1 2 ■ 6-14) ;
foot, 7-3 (6 -8-7 -6); forearm, 33 (32 • 4-34 -G); thumb, 5-7

PIPISTRELLUS 215

(5-2-6); third finger, 61-3 (58-65); fifth finger, 44-6 (41-47).
Adult male and female from Florence, Italy : head and body,
46 and 47; tail, 40 and 38; tibia, 12-8 and 12-6; foot, 6-8
and 7 ; forearm, 35 and 33 ; thumb, 6 ■ 2 and 6 • 6 ; third finger,
63 and 62; fifth finger, 47 and 46; ear from meatus, 12-6 and
12-6; width of ear, 1 1 and 11-4. For cranial measurements see
Table, p. 222.

Specimens examined.— Thinx-thxee, from the following localities :—

Feance : St. Gilles, Gard, 1.

Geemany: Berlin, 1; Bavaria, 1 (U.S.N.M.) ; Ingelheim, Ehein-
hesseu, 1.

Switzerland: Geneva, 6 (Mottaz and Geneva, including type of
unicolor Fatio) ; Montreux, Vaud, 1 (Mottaz) ; Buchillon, Vaud, 6 (Mottaz) •
Neuchatel, 1 (U.S.N.M.) ; Canton Uri, 1 ; St. Gothard, Uri, 1 (U.S.N.M.).

Austkia-Hungary : Palics, Bacser, southern Hungary, 2.

Italy : Siena, 1 (U.S.N.M.) ; Florence, 3 (U.S.N.M.) ; Rome, 5 (B.M.
and U.S.N.M.) ; Borzoli, Liguria, 1 ; Catanzaro, Calabria, 1 (U.S.N.M.).

Bemarks. — Though readily distinguishable from the other
European members of the genus by its cranial and dental
characters, Pipistrelliis nathusii is superficially muoh like P. 2J'pis-
trcllus. It is usually recognizable, however, by its slightly less
diminutive size, more robust form, and by the constant presence
of an ill-defined light (though never actually whitish) border to
the wing.* As pointed out by Mehely in 1900 it has no very
near relationship to the Oriental P. abramus.

6. St. Gilles, Gard, France. G. S. Miller (c). 8. 8. 4. 128.

6 al. Berlin, Germany. Dr. Giinther (c & p). 66. 2. 1. 22.

?. Ingelheim, Rheinhessen. C. Hilgert (c). 8. 11. 2. 3.

2 al. Palics, Bacser, Hungary. Budapest :Museum (e). 0. 4. 9. 1-2.

S. Canton Uri, Switzerland. Tomes Collection. 7. 1. 1. 398.

6,9. Rome. {C. Coll.) G. Barrett-Hamilton (p). 11.1.2.20-21.

9. Borzoli, Liguria. Italy. Marquis G. Doria (c & p). 5. 12. 15. 7.

PIPISTRELLUS KUHLII Kuhl.

1819. Vespertilio kuhlii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv

{- Neue Ann., i), pt. 2, p. 199 (Triest).
1835. Yesp[ertiUo] albolimbatiis Kiister, Isis, p. 75 (Cagliari, Sardinia).
1837. Vespertilio vispistrellus Bonaparte, Iconogr. Faun. Ital., i, fasc. xx

(near Rome, Italy). Type in British Museum.

1837. Vespertilio alcythoc Bonaparte, Iconogr. Faun. Ital., i, fasc. xxi

(Sicily.) Type in British Museum.

1838. Romicia calcarata Gray, Mag. Zool. and Bot., ii, p. 495 (locality

unknown).
1841. Pipistrellus jnarginatus Bonaparte, Iconogr. Fauna. Ital., Indie.

distrib., nomencl. mod. (Substitute for albolivibatus).
1844. Vespertilio 7narginatus Wagner, Schreber's Siiugthiere, Suppl., i,

p. 503, pi. Lv A. No description. Name occurs in synonymy of

kuhlii with ^Michahelles as authority, and on plate. Apparently

not previously published.

* A light border sometimes occurs in P. pipistrellus, but is rare. That
constantly present in P. kuhlii is more sharply defined and more truly
whitish than in P. nathusii.

216

CHIROPTERA

1840. V[es2)ertilio] Ursula Wagner, Schreber's Saugthiere, SuppL, i, p. 505

(Morea, Greece).
1857. Vcsperugo kuhlii Blasius, Saugethiere Deutscblands, p. 63.
1878. Vesperugo kuhlii Dobson, Catal. Cbiropt. Brit. Mus., p. 230.
1886. [Vesperugo kuhlii] var. albicans Monticelli, Atti Soc. Ital. Sci. Nat.,

Milano, xxvii, p. 200, Marcb, 1886 (Caivano, Naples, Italy).
1886. [Vesperugo kuJilii'} v&t . imllatiis Monticelli, Atti Soc. Ital. Sci. Nat.,

Milano, xxvii, p. 200, March, 1886 (Bella Vista, near Portici,

Naples, Italy).
1900. Pipistrellus kuhlii M6hely, Monogr. Cbiropt. Hungarise, p. 261.
1910. Pipistrellus kuhli Trouessart, Faune Mamm. d'Europe, p. 17.

Type locality. — Trieste, Austria-Hungary.

Geographical distribution. — Mediterranean region and eastward
into Asia.

Diatjnoms. — Size about as in Pipistrellus natlmsii (forearm, 31
to 35 mm.; condylobasal length of skull, 12-0 to 13-2 mm.);
outer upper incisor less than half as high as inner incisor ; large
upper premolar almost or (}uite in contact with canine, the small
premolar forced inward from tooth-row and scarcely or not visible
from outer side, its greatest diameter about equal to that of
outer incisor ; canines less robust than in P. pipistrellHs, less
slender than in P.natluisii ; tragus with greatest width less than
length of anterior border ; thumb short, its length about equal to
width of wrist ; posterior edge of wing membrane with sharply
defined whitish border.

External eharacters. — General form very similar to that of
Pipistrellus pipistrellus, the wing similarly narrow as compared
with that of P. natlmsii. Ear narrowly rounded at tip, the posterior
border slightly concave ; antitragus slightly developed, producing
an evident break in contour of conch ; inner surface of conch
somewhat rugose, without well defined transverse striations ;
tragus essentially as in P. natlmsii, the posterior border noticeably
convex. Wing, foot, calcar and tail as in P. p>ip)istrellus.

Fur and colour. — Quality and distribution of fur essentially
as in Pipistrellus pipjistrellus, but dorsal surface of interfemoral
membrane haired scarcely beyond basal third. Colour not very
different from that of Pipistrellus pipistrellus, but somewhat
lighter and more yellow, often approaching raw-siena. Mem-
branes blackish, the wing between foot and fifth finger with a
sharply defined very narrow (less than 1 mm.) nearly white
border.

Skull. — The skull resembles that of Pipistrellus natlmsii in
size, but its form is even more robust than that of P. pipnstrellus.
Dorsal profile with very slight interorbital concavity and barely
perceptible convexity over middle of brain-case. Breadth of
brain-case about half greatest length of skull. Dorsal surface
of rostrum less rounded off" at sides than in P. pipistrellus and
P. natlmsii, but not .so much flattened as in P. savii. Narial
emargination more abruptly narrowed posteriorly than in the

PIPISTEELLUS

217

other European species. Mesopterygoid space slightly longer
than wide. Mandible with coronoid process distinctly higher
than articular process, so that upper edge of posterior portion is
oblique and not pai'allel with alveolar line.

Teeth. — Inner upper incisor essentially as in PipistreUus
yi pistrellus except that secondary cusp is reduced to a minute,
sometimes obsolete, projection from cingulum at posterior base of
shaft ; outer incisor very small, less than half as high as inner,
its apex about on level with highest point of cingulum of larger
tooth, its crown area about two-thirds that of latter, its structure
essentially as in P. pipistreUus, though with very small secondary
cusp ; small tooth situated directly exterior to
large, so that a line perpendicular to main
axis of skull would pass through middle of
all four incisors ; lower incisors essentially as
in P. pipistrellus, though relatively larger and
more strongly imbricated. Canines both abo^e
and below intermediate in form between those
of P. lAjjistrellus and P. nathusii. Anterior
upper premolar crowded inward from tooth- 1/ y,M -ry

row and closely wedged between canine and
large premolar which are nearly or quite in pipistrTiii^'kuhm.
contact ; the small tooth is usually though not Anterior teeth x .'>".
always invisible from outer side, its cusp is
very low, nearly terete, its crown area about equal to that
of outer incisor ; large premolar with no special peculiarities.
Lower premolars as in the related species, but disproportion in
size more marked, the crown area of first a little more than
half that of second. Molars both above and below essentially
as in P. piinstrelhis and P. natliusii, but somewhat more robust.

Measiiremeiifs. — Two adult males from near Genoa, Italy :
head and body, 43 and 46 ; tail, 37 "4 and 38 ; tibia, 12-4 and
13 ; foot, 6 and 6 ; forearm., 33-6 and 34 ; thumb, 5 and 5-2 ;
third finger, 60 and 60 ; fifth finger, 45 and 44 ; ear from meatus,

12 6 and 13; width of ear, 10-4 and 10. Two adult females
from the same locality : head and body, 44 and 47 ; tail, 39 and
40 ; tibia, 1 2 and 13 ; foot, 6 ■ 2 and 6 • 2 ; forearm, 33 • 4 and 35 ;
thumb, 5-2 and 5-4; third finger, 60 and 63 ; fifth finger, 45
and 47 ; ear from meatus, 13 and 13 ; width of ear, 10 and 10.
Adult male and female from Palermo, Sicily : head and body,
42 and 44 ; tail, 36 and 35-4 : tibia, 12-4 and 12-4 ; foot, 5-6
and 5 • 4 ; forearm, 3 1 and 34 ; thumb, 5 • 4 and 5 • 2 ; third finger,
56 and 61 ; fifth finger, 41 and 43 -6 ; ear from meatus, 12-4 and

13 ; width of ear, 10 and 10. Adult male and female from
Cagliari, Sardinia : head and body, 44 and 45 ; tail, 35 and 37
tibia, 12-6 and 12-4 ; foot, 6-2 and 6-8 ; forearm, 33 and 33
thumb, 5 and 5 ; third finger, 58 and 62 ; fifth finger, 43 and 45
ear from meatus, 13 and 12 ; width of ear, 10 and 10. For
cranial measurements see Table, p. 218.

218

CHIROPTERA

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Balearic Islands : San Cristol
France : St. Gilles, Gard
Italy : Perti, Finalborgo

Siena

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Catans^aro, Calabria

Sicily : Palermo .

Ustica Island .
Sardinia: Wt. Gennargcntu

Greece : Corfu

Argostoli, Cephaloni

PIPISTRELLUS 219

Specimens examined. — One hundred and eight, from the followino
localities : — "^

Feaxce: St. GiUes, Gard, 2; Nimes, Gard, 3 (Mottaz) ; St. Genies
Gard, 4 (Mottaz) ; Marseilles, 1 (U.S.N.M.).

Switzerland: Coremmo, Ticino, 1 (Mottaz) ; Lugano, Tioino, 1 (Mottaz).

Italy; Near Genoa, 14 (U.S.N.M. and Genoa); Siena, 6 (B.M. and
U.S.N.M.); Florence, 1 (U.S.N.M.); Rome, 6 (B.M. and U.S.N.M.);
near Rome 1 (type of vispistrellus Bonaparte) ; Sorrento, 3 (U.S.N.M.) ;'
Catanzaro, Calabria, 6 (U.S.N.M.) ; Palermo, Sicily, 23 (U.S.N.M ) '
Corleone, Sicily, 1 (U.S.N.M.); Ustica Island, Sicily, 4 (U.S.N.M.);
Sicily, 1 (type of alcythoe Bonaparte).

Sardinia: Cagliari, 4 (U.S.N.M.); Mt. Gennargentu, 7 (U.S.N.M.);
no exact locality, 2.

Greece : Corfu, 2 ; Cephalonia, 10 ; Patras, 3.

Spain : San Cristobal, Minorca, Balearic Islands, 2.

Bemarks. — PipistreUus huhlii is easily recognizable by the
form and relative size of the upper incisors. Externally it may
usually be known by the sharply defined whiti.sh border to the
wing membrane, though too much reliance should not be placed
on this character alone. Many specimens from Sardinia are
lighter in colour than those from the mainland. These represent
the albolimhatus of Kiister. Normally coloured examples also
occur ; and in the absence of adequate material it has seemed
preferable for the time being not to attempt to define the insular
form.

6. St. GiUes, Gard, G. S. MiUer (c). 8. 8. 4. 127.

France.
<^. Siena, Italy. Dr. E. Hamilton (p). 98. 10. 2. 2.

(S. Brogi.)
3 6, ?. Rome. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. IG-IP

(P)-
skeleton with- Sicily. {Prince Bona- Tomes Collection. 7. 1. 1. 780.

out skull. 2^a'-tc.) {Type of V. alcythoe Bonaparte.)

skeleton with- Near Rome, Italy. Tomes Collection. 7. 1. 1. 7"29.

out skuU. {Prince Bonaparte.) {Type oi F. vispisfeZ^MS Bonaparte.)

i, 9. Corfu, Greece. J. I. S. Whitaker (p). 8. 10. 1. 4-5.

(C. Mottaz.)
3 <;. Argostoli, Cephalonia. J. I. S. Whitaker (p). 8. 10. 1. 1-:3.

(C. Mottaz.)
6, 9. Patras. {C. Mottaz.) Hon. N. C. Rothschild 8. 10. 2. 17-18.

2 c?. San Cristobal, O. Thomas and R. I. 0.7.1.29-30.

Minorca, Balearic Pocock (c & p).
Islands.

PIPISTRELLUS SAVII Bonaparte.

1837. Vespertilio savii Bonaparte, Iconogr. Faun. Ital., i, fasc. xx (Pisa)

Type in British Museum.
1837. Vespertilio aristippe Bonaparte, Iconogr. Faun. Ital., i, fasc. xxi

(Sicily).

1837. Vespertilio leucippe Bonaparte, Iconogr. Faun. Ital., i,. fasc. xxi

(Sicily). Type in British Museum.
1833. Vespertilio bonapartii Savi, Nuovo Giorn. de' Lettcrati, Pisa, sxxvii

p. 226 (Tuscany).

220

CHIROPTEEA

18i4. Vesp[ertilio] nigrans Crespon, Faune Meridionale, i, p. 24 (Nimes,

Gard, France).
1853. Vesperugo manrus Blasius, Wiegmann's Arcliiv fiir Naturgesch.,

1853, I, p. 35 (Central chain of the Alps).
1857. Vesperugo maurus Blasius, Siiugethiere Deutschlands, p. 67.
1872. V[espertilio] agilis Fatio, Faune Vert. Suisse, Append, au vol. i,

p. iii (Alternative name for V. savii Bonaparte, ex Savi MS.).
1878. Vesperugo maurus Dobson, Gatal. Chiropt. Brit. Mus., p. 218.
1904. Vespertilio ochromixtus Cabrera, Mem. Soc. Espaii. Hist. Nat., ii,

p. 267 (Sierra de Guadarrama, Madrid, Spain).
1910. PipistreUus savii and P. savii oclironiixtus Trouessart, Faune Mamm.

d'Europe, pp. 13-14.

Tiipc locality. — Pisa, Italy.

Geographical distribution. — Southern Europe, west to the
Iberian Peninsula, north to the Alps ; also the Canary Islands,
northern Africa and southern Asia. Limits of range very
imperfectly known.

Diagnosis. — Largest European member of the genus (condy-
lobasal length of skull, 13 to 14 mm. ; forearm, 31 to 33 mm.) ;
outer upper incisor more than half as high as inner ; large
premolar broadly in contact with canine, the small tooth very
minute, crowded inward from axis of tooth-row, invisible from
outer side and occasionally covered by the gum, its diameter
much less than that of outer incisor ; anterior lower premolar
with crown area less than half that of succeeding tooth ; lower
canine robust ; tragus with greatest width nearly equal to length
of anterior border ; thumb short ; hairs of back usually with
contrasting light brown tips.

External characters. — Ear broad, its general form about as in
PipistreUus nathusii, the posterior border slightly but evidently
concave above middle, the inner surface of conch noticeably
rugose and with faint, irregular transverse ridges behind tragus ;
antitragus small aiid ill-defined, but producing an evident break
in outline of conch ; tragus less than half as high as conch, very
wide (greatest width, at level of middle of anterior border, nearly
equal to length of anterior border), the anterior border nearly
straight, the posterior border strongly and evenly convex from
tip to notch above small basal lobe. Wing, foot, calcar and tail
as in P. pipistrellus.

Fur and colour. — The fur resembles that of Piplstrellus Tcuhlil
in quality and distribution. Colour differing from that of the
other European species in the evident contrast between light
tips of hairs of back and darker ground tint. It is also the only
species in which there is much individual variation in colour.
Four specimens from the neighbourhood of Genoa are coloured as
follows : male, not fully adult, uniform very dark vandyke-brown,
the extreme tips of hairs of back faintly lighter, underparts a
light brown faintly overlaid on blackish under colour ; adult male :
light tips on back well developed, giving general colour to region,
between raw-umber and clay-colour ; adult male : light tips very

PIPISTRELLUS 221

conspicuous, a peculiar dull brownish ochraceous-bufF; adult female :
light tips as conspicuous as in last, dull brownish cream-buft".

SJatll. — The skull is slightly larger than that of Pipistrellus
nafhiisii and P. kuhVu, and is immediately distinguishable among
the European species by the flatness of dorsal surface of rostrum,
prominence of ridge along edge of orbit, and
relatively small size of narial emargination,
characters the first two of which suggest
Verpcrtilio murinus. Dorsal profile of skull as
in P. jiipistrellus and P. Icnldii, but brain-case
slightly more depressed. Breadth of brain-case
about half greatest length of skull. Rostrum
relatively broader than in the other European
species, its dorsal surface more flattened and
orbital ridges more prominent ; narial emar-
gination scarcely larger than in P. pipistrellus.
Mesopterygoid space about as wide as long, its „• V",,""
general outline, aside irom the notch caused Nat. size,

by median spine of palate, broadly barrel shape,
the hamulars distinctly turned inward. Mandible with coronoid
process and upper border of posterior portion as in P. hihlii,
but with angular process less curved and relatively longer than
in the other European species.

Tt'eth. — Incisors both above and below essentially as in
P/jjitifrellus pipistrellus, except that inner upper tooth has the
secondary cusp somewhat better developed.
Canines with no .sj^ecial peculiarities, not
essentially different from those of P. pipi-
strellus. Anterior upper premolar very
minute, sometimes hidden in the gum or
occasionally absent, its crown area never
much more one-sixth that of outer incisor ;
large premolar always strongly in contact
. .. with canine, its form peculiar in the absence

Anterior teeth, x's. Or slight development of the antero-internal
cusp. Lower premolars strongly contrasted
in size, the crown area of first decidedly less than half that
of second, its cusp relatively lower and less developed than in
anj' of the other European species. Molars both above and
below with no special peculiarities.

Measurements. — Two adult males from Palermo, Sicily : head
and body, 43 and 47 ; tail, 34 and 35; tibia, 12-6 and 12-8;
foot, 6-4 and 7; forearm, 31 and 32-6; thumb. Sand 5-6;
third finger, 54 and 56 ; fifth finger, 42 and 41 ; ear from meatus,
12-4 and 12-6; width of ear, 'l2 and 12. Two adult females
from the same locality : head and body, 46 and 47 ; tail, 35 and
39 ; tibia, 13 and 13-4 ; foot, 6-6 and 7 ; forearm, 33 and 33 ;
thumb, 5 and 5-4 ; third finger, 56 and 57 ; fifth finger, 42 and
43 ; ear from meatus, 12-4 and 13 ; width of ear, 12 and 11-6.

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224

CHIROPTEKA

Adult male from Escorial, Spain (paratype of ochromi.vtus Ca.hrern) :
head and body, -iS-G ; tail, 34 ; tibia, 13-2; foot, 5-6; forearm,
57 ; thumb, 5 "4; third finger, 43 ; ear from meatus, 13 ; width
of ear, 11. For cranial measurements see Table, p. 223.

Specimens examined. — Twenty-four, from the following localities : —

Switzerland : St. Gothard, Uri, 1 ; no exact locality, 1.

Italy: Near Genoa, 10 (B.M. and Genoa); Florence, 3 (Mottaz) ;
Sorrento, 1 (U.S.ISr.M.) ; no exact locality, 1 (type) ; Palermo, Sicily, 8
(U.S.N.^I.); Sicilv, no exact locality, 1 (type of leucippe Bonaparte);
Ustica Island, Sicily, 1 (U.S.N.M.).

FRA^'CE : Near Nimes, Gard, 2 (U.S.N.M. and Nimes ; the latter
agreeing with description of nigrans Crespon, though not marked type) ;
St. Gilles, Gard, 1 ; no exact locality, 1.

Spain: El Escorial, iladrid, 1 (paratype of ochromixtus Cabrera).

Be marks. — This species is readily distinguishable among the
European members of the genus Plpi-s-freUus by the peculiar form
of the tragus, apart from its very pronounced cranial and dental
characters. Its colour gives it a superficial resemblance to
Ejjtesicus nilssoni, a likeness that is so heightened by the exces-
sively small size of the anterior upper premolar that the animal
has been once and pierhaps twice described as a member of the
genus EptesicKs*

1. St. Gothard, Uri, Swit- Tomes Collection. 7. 1. 1. 397.

zerland.
Switzerland. Purchased (Brandt). 45. 11. 1. 3.

4 al. Genoa, Liguria, Italy. Genoa ^Museum (e). 86.11.3.14-

17.
i. Borzoli, Liguria. !Marquis G. Doria 5. 12. 15. 6.

(c & P).
skeleton with- Italy. {Prince Bona- Tomes Collection. 7. 1. 1. 7-32.

out skuU. parte.) [Type of species.)

skeleton with- Italy. [Prince Bona- Tomes Collection. 7. 1. 1. 731.

out skull. parte.) (Type of F. Z<;«cy)pt' Bonaparte.)

? al. France. Purchased (Lefebre). 46.1.2.12.

6. El Escorial, Madrid, A. Cabrera (p). 8. 7. 23. 1.

Spain. (Paratype of Y. ochromixtus Cabrera.)

Genus EPTESICUS Ratinesque.

1820. Eptesiciis Rafinesque, Annals of Nature, p. 2 [melanops = fitscus).
1S29. Cnephxus Kaup, Entw.-CTesch. u. Natiirl Syst. Europ., Thierwelt, i,

p. 103 [serotinus).
1839. Vespcrugo Keyserling and Blasius, Wiegmann's Archiv fiir Natur-

gesch., 1839, i, p. 312 (part).
1839. Vesp>crus Keyserling and Blasius, Wiegmann's Archiv fiir Natur-

gesch., 1839, i, p. 313 (Sub-genus of Vesperugo, part). Not Yesperns

Latreille, 1829.

* This is certainly the case with the Vespertilio ochromixtus of Cabrera.
In the paratype of this species (B.M. no. 8. 7. 23. 1), which I carefully
examined before removal of the skull, in company with Mr. Knud Andersen,
no trace of the small premolar could be found. Wbeu the skull was cleaned,
however, the presence of the tooth in its normal position was revealed,
thus showing the animal's true identity. It seems not improbable, so far
as can be judged from the original description, that Satunin's Ycspcrugo
caucasicus (Zool. Anzeiger, xxiv, p. 462, August 5, 1901) was based on
similar specimens.

EPTESICUS

225

I

1841. JSoctula Bonaparte, Iconogr. Faun. ItaL, i, fasc. xxi, in account of

Vespcrtilio alcijthoc {serotinus).
1856. Cateonis Kolenati, Allgeni. deutsche Naturhist. Zeituug, Dresden,

neue Folge, ii, p. 131 (serotinus).

1856. Meteorus Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden,

neue Folge, ii, p. 131 (part).

1857. Vesperus Blasius, Saugethiere Deutschlands, p. 51 (Sub-genus of

VesjJcrugo), part.

1858. Amblyotiis Kolenati, Sitzuugsber. kais. Akad. Wissensch. Wien,

Math.-Naturwissensch. Classe, xxix, p. 252 (atratiis — nilssoni).
1863. Aristijipc " Kolenati, Beitriige zur Kenntniss der Phthiriomyiarien,

Petersburg, 1803" (part, included both discolor = vmrinus and

7iilsso?ii).
1866. Pachijomus Gray, Ann. and Mag. Kat. Hist., 3rd ser., xvii, p. 90,

February, 1863 (ixiclii/omus).
1870. Nyctiptenus Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien,

Math.-Naturwiss. Classe, lxii, p. 424 (smithii).
1878. Tesj^enis Dobsou, Catal. Chiropt. Brit. Mus., p. 184 (Sub-genus of

Vespcriigo), part.
1892. Adelomjcteris H. Allen, Proc. Acad. Nat. Sci. Philadelphia, 1891,

p. 466, January 19, 1892 (part; substitute for Vesperus, pre-
occupied).
1897. Yespcrtilio Miller, Ann. and Mag. Nat. Hist., 0th ser., xx, p. 384,

October, 1897 (part).
1900. Eptesicus Mehely, Monogr. Chiropt. Hungarise, p. 219 (part).
1907. Eptesicus Miller, Families and Genera of Bats, p. 207, June 29, 1907.

Type species. — Eptesicus melanops Rafinesque = Verpertilio
fuscus Beauvoi.s.

Geographical distribution. — Europe, Asia (except Malay region),
Au.stralia, Africa, Madagascar ; America from southern Canada
southward (except Lesser Antilles).

Characters. — Dental formula : / |-^, c J--i, pm j^,, m p", = 32.
Teeth strictly normal throughout, and showing no special pecu-
liarities. Both upper incisors well developed, the inner larger
than the outer and usually with distinct secondary cusp, the
outer separated from canine by a space equal to its greatest
diameter ; m^ variable in form, usually with well developed
metacone and three commissures in the smaller species, but with
raetacone and third commissure obsolete in larger forms. Skull
without special peculiarities of form or structure, the rostrum
tlattish or more usually rounded ofl' above, the nares and palatal
emargination not specially enlarged, the latter at least as deep
as wide. Ear of moderate size, not peculiar in form : wing
broad (normal).

BemarJcs. — Among European bats the members of the genus
Eptesicus may be distinguished by their dental formula combined
with a simple Pij>i.'<trelliis-]\ke ear and not specially modified
skull. The group is nearh' related to PipistrelJus through
P. snvii, in which the small premolar is occasionally absent and
not infreciuently so minute as to be concealed by the gum. About
forty-five species are known, three of which occur in Europe.

226 CHIROPTEEA

KEY TO THE EUBOPEAN SPECIES OF EPTESICUS.

Forearm less than 40 mm. ; coudylobasal length of
skull less than 16 mm. ; a distinct line of demar-
cation between colours of upper and lower surfaces
of neck E. nilssonii, p. 2.34.

Forearm more than 45 mm. ; condj'lobasal length of
skull more than 17 mm. ; no line of demarcation
between colours of upper and lower surfaces of
neck.

Coudylobasal length of skull 19 to 21*6 mm E. serotinus, p. 226.

Condylobasal length of skull about 18 mm E. sodalis, p. 231.

EPTESICUS SEROTINUS Schreber.

1774. Vcspertilio serotinus Schreber, Siiugthiere, i, pi. liii (Description, i,

-p. 167, 1775, under name : Die Blasse Fledermaus). France,

based primarily on "La Serotine " of Daubenton, Hist. Acad.

Royale des Sci., 1759, p. 377. 1765.
1776. Vcspertilio serotine P. L. S. Miiller, Natursyst. Suppl. u. Regist.-

Band, p. 16 (Based on "die Blasse Fledermaus" of Schreber).
1827. Vcspertilio u-iedii Brehm, Ornis, Heft in, p. 24 (Renthendorf,

Thiiringen, Germany).
1827. Vespertilio okcnii Brehm, Ornis, Heft in, p. 25 (Renthendorf,

Thiiringen, Germany).
1844. Vcspjlertiliol incisivus Crespon, Faune M6ridiouale, i, p. 26 (Nimes,

Gard, France).
1857. Vcsperugo serotinus Blasius, Siiugethiere Deutschlands, p. 76.
1863. [Cateorus serotiniis'\ var. typus Koch, Jahrb. des Vereins fiir Natur-

kunde im Herzogthum Nassau, xviii, p. 466 (Wiesbaden, Nassau,

Germany).
1863. \_Cateorus serotinus'] var. rufesccns Koch, Jahrb. des Vereins fiir

Naturkjude im Herzogthum Nassau, xviii, p. 466 (Freiburg,

Breisgau, Germany).
1878. Vesiwrugo serotinus Dobson, Catal. Chiropt. Brit. Mus., p. 191.

1885. Vespierus serotinus var. transylvanus Daday, Orvos-Termeszettudo-

manyi Erteset(5, Kolozsvar, x, p. 275 (Also-Szocs, Szolnok-Doboka,
Hungary).

1886. Vesperus serotinus var. transsylvanus Daday, Verhandl. u. Mittheil-

ungen des Siebenbiirgischen Vereins fiir Naturwissensch. in

Hermannstadt, xxxvi, p. 81.
1900. Eptesic.us serotinus Mehely, Monogr. Chiropt. Hungarise, p. 209.
1904. Vespertilio serotinus insularis Cabrera, Mem. Soc. Espan. Hist. Nat.,

II, p. 263 (Minorca, Balearic Islands).
1904. Vcspicrtilio isnhellinus Cabrera, Mem. Soc. Espan. Hist. Nat., ii,

p. 264 (southern Spain). Not of Temminck.
1904. Vcspertilio hoscai Cabrera, Mem. Soc. Espan. Hist. Nat., ii, p. 265

(Muchamiel, Alicante, Spain).
1910. Eptcsicus serotinus, E. serotinus transsylvanus, and E. boscai Troues-

sart, Faune Mamm. d'Europe, pp. 20-22.

Tj/jye locality. — France.

GeograjMcal distrihution. — Central and southern Europe
from England and Denmark to the Mediterranean ; eastward
into Asia.

Diaynosis. — Size rather large (forearm more than 45 mm.,

EPTESICUS.

227

condylobasal length of skull more than 19 mm.) ; colour of upper
parts a yellowish brown without noticeably contrasted light tips
to the hairs ; under parts essentially similar, so that there is no
line of demarcation along sides of neck.

External characU'rs. — General form robust, though less so
than in Nyctalus, the tail and legs rather short, the membranes
thick and opaque. Muzzle with moderately prominent glandular
swelHngs, its greatest width across this region less than distance
from nostril to ear ; nostrils projecting very slightly, the concavity
between them not conspicuous, the orifice crescentic. Ear
moderately long, extending slightly more than half way from eye
to nostril when laid forward, its breadth when flattened about
equal to height above crown ; anterior border of conch abruptly
convex below, then nearly straight to narrowly rounded olT tip ;
posterior border straight or irregularly concave from just below
tip to level of anterior base, then convex to abrupt angle under
meatus marking posterior limit of small but well defined anti-
tragus, the anterior border of which terminates obscurely about
3 mm. behind angle of mouth ; inner surface of conch obscurely
papillose, the region behind tragus marked by about six faint
cross ridges ; tragus short, its height less than half that of ear
conch, its anterior border straight, its posterior border gently
convex from narrowly rounded tip to upper edge of small but
distinct basal lobe, its greatest width, at level of middle of anterior
border, equal to slightly more than half length of anterior border.
Wing broad, the fifth finger exceeding forearm by one-quarter
to one-third length of forearm, the membrane leathery and
opaque, though perhaps less so than in Nyctahis noctula, joining
leg at base of outer toe ; third and fourth metacarpals sub-equal,
nearly as long as forearm, fifth about 2 mm. shorter. Leg rather
slender, the foot less than half as long as tibia ; calcar about
four-fifths as long as tibia and slightly exceeding length of free
border of uropatagium, its keel ill-detined, its terminal lobe well
developed though small. Tail extending to between shoulders
wlien laid forward, the last vertebra and distal tliird of penulti-
mate vertebra free from membrane.

Far and colour. — Fur soft and dense, the longest hairs on
back about 10 mm. in length, those of underparts shorter. It
is strictly confined to body, only extending as a thin pubescence
on extreme base of membranes and along a narrow line bordering
under surface of forearm ; free edge of uropatagium naked,
(i round colour of upper parts ranging from prouts-brown to a
light wood-brown, the basal portion of the hairs not essentially
different, the tips of the hairs of back behind shoulders ^\•ith
inconspicuous lighter (bufi'y) tips ; underparts slightly paler,
sometimes approaching ochraceous-bufi", but never sufficiently
contrasted to produce a line of demarcation along sides of neck.
Muzzle, cheeks, ears and membranes blackish. The variation
in general colour appears to be strictly individual.

Q 2

228

CHIEOPTERA

SJcuH. — General aspect of skull robust and flattened, with
widely spreading zygomata, but rather narrow brain-case and
rostrum. Dorsal prolile rising gradually from nares to over-
hanging lambda, essentially straight throughout, though with
slight concavity over lachrymal region. Ventral profile very
slightly elevated posteriorly. Brain-case ovate in general outline,
narrower than in Ni/ctahis noctula, the straight, well developed
lambdoid crests which form its posterior border meeting in median
line almost at right angles, depth at middle about half mastoid
breadth ; sagittal crest low but evident, the region at each side
of it not depressed ; floor of brain-case smooth, with no evident
ridges or depressions, a very narrow slit between cochlea and
basioccipital ; auditory bulhe small, the transverse diameter con-
siderably less than distance between bulla?. Interorbital region
moderately constricted, hour-glass shaped, the lachrymal region
decidedly less wide than brain-case,
with slight tubercular projection close
to anterior rim of orbit ; rostrum
flattened, with shallow but evident
lateral concavity on each side, dis-
tinctly narrower anteriorly than pos-
teriorly, the narrowly obovate narial
emargination extending about half
way back to level of lachi'ymal fora-
men ; rostral depth at front of orbit
less than distance from orbit to outer
incisor ; anteorbital foramen less re-
duced than in Ni/ctalus noctula, its
posterior border over region of con-
tact between large premolar and first
molar, lachrymal foramen directly
behind it, on inner side of orbital
rim. Palate long and narrow, slightly concave both laterally and
longitudinally, the anterior emargination small, squarish, extend-
ing back to level of middle of canine. Posterior extension of
palate nearly parallel sided, though narrowing a little posteriorly,
its width at level of posterioi' molar considerably less than its
length ; hamulars slightly turned inward ; median spine well
developed. Mandible robust, but with lower border nearly
parallel to alveolar line ; posterior portion high in front, low
behind, the height of coronoid process above level of alveolar
line about equal to horizontal diameter, the upper border sloping
abruptly from coronoid to articular process ; angular process
moderately long, about on level with alveolar line, its main axis
directed gradually outward and downward, its distal extremity
slightly expanded and hooked upward.

Teeth. — Relatively to size of skull the teeth are large and
robust, rather more so than in Nyctalini noctula. Inner upper
incisor about half as high as canine, its crown area about one-

Fia. 40.
Eptesicus serotinus.

EPTESICUS

229

quarter that of canine, the subterete shaft directed inward and
slightly forward, its secondary cusp large and prominent, situated
on outer side of shaft near tip ; ciugulum well developed, but
without CUSJ3S. Outer upper incisor much shorter than inner
and with about half its crown area, the apex of its shaft slightly
exceeding level of cingulum of larger tooth ; outer and posterior
surfaces flattened or double-concave, the two concavities some-
times separated by a low but evident ridge ; inner margin with
a low secondary cusp on well developed cingulum. A line
perpendicular to main axis of skull would pass through centre
of shafts of all four incisors ; the outer tooth is separated from
canine by a space about equal to breadth of its own crown. Lower
incisors large, much crowded, and very conspicuously imbricated,
i\ and ?.j overlajjping more than half of front surface of the
succeeding tooth, the general outline of the entire series V-shaped ;
front surface of crowns about as high as wide, the edge obliquely
trifid (occasionally a low, rudimentary fourth cusp at outer
margin of i.^) ; cross section of crown somewhat triangular, the
posterior angle occupied by a low tubercle in i., and ^'3. Upper
canine large, its shaft decidedly the highest of the upper series,
its cross section sub-triangular, the posterior cutting edge well
developed, the anterior less trenchant than in Nyctalns noctula ;
inner surface divided by a low ridge into two shallow con-
cavities, the posterior of which is the larger ; antero-outer surface
convex ; postero-outer surface with deep longitudinal groove ;
cingulum narrow but complete, without cusps. Lower canine
very robust, the diameter of crown noticeably greater than least
distance between canines, the shaft decidedly higher than main
cusps of molars, smoothly rounded in front, flattened-concave
behind and on inner side, the cingulum narrow but complete
except at point of contact with jj, where it becomes abruptly
obsolete, terminating in a slight tubercle corresponding to the
cusp present in Ni/ctalus and Pipistrellus. Upper j^remolar with
crown area about ecjual to that of canine and about two-thirds
that of first molar, its main cusp robust, nearly as long as canine
(measured along cingulum), sharply trenchant posteriorly, flcxt-
tened-concave on inner side, a well developed external and antero-
internal longitudinal groove ; crown with slight anterior and
more marketl posterior emargination, the inner side narrow, with
slight concave crushing surface and well developed cingulum,
which rises to a small cusp anteriorly. Lower premolars closely
crowded, the crown area of first about half that of canine, that
of second nearly four-fifths that of canine ; cusp of first a little
more than half as high as second, which slightly exceeds main
cusps of molars ; cingulum of each tooth well developed, tending
to form a slight postero-internal tubercle. First and second
upper molars sub-equal in crown area, the second wider but more
coustricted at middle ; protocone robust, not very high ; no true
hypocone, but region which it would occupy indicated by slight

230

CHIEOPTERA

columnar thickening of posterior base of protocone ; paracone
lower and smaller than metacone, the contrast unusually notice-
able ; styles and commissures well developed, the W-pattern
normal ; m'' with crown area less than half that of 711^, its longi-
tudinal diameter through metacone much less than half transverse
diameter, the mesostyle, metacone, and second and third com-
missures greatly reduced, though not sufficiently to lose their
identity. Lower molars with no special peculiarities ; angles in
commissures between outer and inner cusps rather wide and
shallow, especially that between protoconid and metaconid ; area
of second V in "'3 scarcely half that of first.

Measurements. — Adult female from Herrnhut, Saxony : head
and body, 62 ; tail, 54 (its free tip, 6-6) ; tibia, 20 ; foot, 10-4 ;
forearm, 50-4; thumb, 9 ; third finger, 84 : fifth ringer, 63; ear
from meatus, 18; width of ear, 15. Adult male from Barsac,
Gironde, France : head and body, 69 ; tail, 54 (free tip, 7 ■ 6) ;
tibia, 22 ; foot, 11 -4 ; forearm, 51 ; thumb, 9 ; third finger, 91 ;
fifth finger, 67 ; ear from meatus, 18 '4 ; width of ear, 16. Adult
male and female from Seville, Spain : head and body, 67 and 64 ;
tail, 46 and 51 ; tibia, 21 -6 and 22-4 ; foot, 11 and 10'4 ; fore-
arm, 49 and 50 ; thumb, 9 • 2 and 8 • 4 ; third finger, 89 and 90 ;
fifth finger, 64 and 68; ear from meatus, 17 '6 and 18. Adult
male and female from Rome, Italy : head and body, 72 and 73 ;
tail, 54 and 54 ; tibia, 21-4 and 21 ; foot, 11 and 11 -4 ; fore-
arm, 51-6 and 53 ; thumb, 9 • 6 and 9 ■ 8 ; third finger, 91 and 92 ;
fifth finger, 66 and 67 ; ear from meatus, 19 and 19 ; width of
ear, 15 '6 and 15 -4. For cranial measurements see Table,
p. 232.

Specimens examined. — Seventy-eight, from the following localities: —

England: Kenley, Surrey, 1 ; Hawkhurst, Kent, 1; Whitstable, Kent, 2 ;
Wingbam, Dover, Kent, 1 ; Yalding, Kent, 5 (B.M. and U.S.N.JI.) ; Isle
of Wight, 5.

France : Barsac, Gironde, 1 (U.S.N.M.) ; near Nimes, Gard, 2 (Nimes ;
type of inc'isivus Crespon, and a specimen wrongly marked type of palusiris
Crespon).

Geemany : Ingelheim, Rheinhessen, 2 ; Magdeburg, Saxony, 2 (B.M.
and U.S.N.M.); Herrnhut, Saxony, 1 (U.S.N.M.); Strass, near Burgheim,
Bavaria, 3; Bavaria, no exact locality, 2 (U.S.N.jNI.) ; Tubingen, Silesia, 2.

Austria-Hungary : Moravia, 1 ; Csallokoz-Somorja, Pressbnrg, Hun-
gary, 1 ; Budapest, Hungary, 2 ; Transylvania, 5 ; Zara, Dahuatia. 2.

Boumania : Bustenari, Prahova, 3.

Greece: Patras, 14 ; near Athens, 1.

Italy : Siena, 3 (B.M. and U.S.N.M.) ; Florence, 2 (B.M. and U.S.N.M.) ;
Vallombrosa,! (U.S.N.M.); Volterra, 1 (U.S.N.M.) ; Rome, 3 ; San Martino
al Cimino, Rome, 1 (Genoa) ; Ustica Island, Sicily, 1 (U.S.N.^I.).

Sardinia : Cagliari, 2 (Genoa).

Spain : Pajares, Leon, 1 ; Seville, 3 ; Muchamiel, Alicante, 1 (Madrid ;
type of boscai Cabrera).

Bemarks. — Among the bats of Europe this species is recogniz-
able by its rather large size, noticeable free tip to the tail,
moderately long, narrow ear, and straight, erect tragus. With
the material now available for study it seems impossible to

EPTESICUS

231

distinguish any local geographical forms. Specimens from Seville
representing the isabelUnus of Cabrera I am unable to separate
from true serotlHuii ; the type of huscai Cabrera is a young of the
same animal ; insularis I have not seen, but there is nothing in
the original description to indicate that it is distinct.*

2 9.
$al.

1 St.

3 9.

9al.
2.
1.

1.

6,9.

2 J, 9.

2al.

lal.

2.

2al.
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6, 9.
6, 9.

1 9.

9.
9.

1 al.

6.
9.

9al.

lal.

1.

Kenley, Surrey, England.

Whitstable, Kent.
Wingham, Kent.
Yalding, Kent.
Yalding, Kent.

Isle of Wight.

Freshwater, Isle of Wight.

Freshwater, Isle of Wight.
(F. Bond.)

Bembridge, Isle of Wight.

Ingelheim, Bheinhessen,
Germany.

Magdeburg, Saxony.
(Wolterstorff.)

Strass, Burgheim, Bavaria.
(Korbitz.)

Tiibingen, Silesia.

Moravia, Hungary.

Csallokoz-Somorja, Press-
burg.

Budapest.

Transylvania.

Zara.Dalmatia. (K.Blos.)

Bustenari, Prahova, 840 m.

Roumania. ( W. Dodson.)

Patras, Greece. {C.Motfaz.)

Patras.

Athena. (C. Mottaz.)

Florence, Italy.
Rome. (C. CoU.)
Pajires, Leon,

(N. Gonzalez.)
Seville.
Seville.
Europe.

Spain.

W. R. Ogilvie-Grant

(c & p).
C. H. B. Grant (c & p).
G. Donker (c & p).
H. Reid (c & p).
W. R. Ogilvie-Grant

(c & p).
Rev. C. Bury (c & p).
F. Bond (c & p).
Tomes Collection.

Tomes Collection.
C. Hilgert (c).

Lord Lilford (p).

Lord Lilford (p).

Dr. A. Giinther (p).
Purchased (Parreys).
Budapest Museurq. (e).

Budapest Museum (e).
C. G. Danford and J. A.

Brown (c & v).
Lord Lilford (p).
Lord Lilford (p).

Hon. N. C. Rothschild

C. Mottaz (c).

Hon. N. C. Rothschild

Florence Museum (e).
G.Barrett-Hamilton (p).
O. Thomas (p).

Dr. V. L. Seoane (p).
Seville Museum (e).
Leyden Museum (e).

11. 1. 3. 45.

11.1.3.46-47.
90. 4. 17. 1.
97. 8. 27. 1.
11.1.3.42-44.

44. 6. 16. 7.
61. 11. 5. 1-2.
7. 1. 1. 352.

7. 1. 1. 353.

8. 11. 2. 1-2.

11. 1. 1. 38.

11. 1. 1. 33-.35.

66. 2. 1. 7-8.
46. 6. 15. 54.
94. 3. 1. 12-13.

94. 7. 18. 6-7.
74. 7. 4. 1-5.

11. 1. 1. 36-37.
4. 4. 6. 2-3.

8. 10. 2. 1-13,

15.
8. 11. 8. 3.
8. 10. 2. 14.

85. 7. 6. 13.
11. 1. 2. 28.
8. 2. 9. 1.

94. 1. 1. 6.
94. 5. 8. 1.
37. 4. 28. 58.

EPTESICUS SODALIS Barrett-Hamilton.

1910. Vespertilio sodaU.v Barrett-Hamilton, Ann. and Mag. Nat. Hist.,

8th ser., v, p. 291, March, 1910. Type in British Museum.
1910. Eptesicus sodalis Trouessart, Faune !!Mamm. d'Europe, p. 22.

Tj/pc locality. — Bustenai'i, Prahova, Koumania (in Carpa-
thians, alt. 840 m.).

(h'0(iraphical (listrihufidu. — Known only from the type locality
and St. Gothard, Switzerland.

* Mr. Cabrera has come to the same conclusion (Bol. Real Soc. Espafi.
Hist. Nat., VI, p. 449, December, 1908).

232

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234

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Diagnosis. — Similar to Eptesims serotinus but smaller, condylo-
basal length of skull about 18 mm. instead of 19 to 21-6 mm.

Measurements. — Type (young-adult male) : head and body,
63; tail, 42; tibia, 18-6; foot, 9-8; forearm, 45-4; third
finger, 79-0; fifth finger, 58-0 ; ear (fresh), 18. Adult from
St. Gothard, Switzerland : tibia, 19 ; forearm, 48. For cranial
measm-ements see Table, p. 233.

Specimens examined. — Two, from the following localities : —
Switzerland: St. Gothard, 1 (U.S.N. M.).
RouMANiA : Bustenari, Prahova, 1 (type).

RemnrJcs. — The two specimens on which this species is based
indicate the existence of an animal bearing much the same
relationshij) to Eptesirus serotinus as Nyctalus nocfula to N.
maximus.

S. Bustenari, Prahova, 840 m. Lord Lilford (p). 4. 4. 6. 1.

Roumauia. (W. Dodson.) [Type of species.)

EPTESICUS NILSSONII Keyserling and Blasius.

1836. Vespertilio kuhlii Nilsson, Ilium. Fig. Skand. Fauna, pt. 17, pi. .34

upper figure. Not of Kuhl, 1819.
1888. Vespert\ilio'] borealis Nilsson, Ilium. Fig. Skand. Fauna, pt. 19,

pi. 34 (renumbered 36) upper figure. Not of P. L. S. Miiller, 1776

(Scandinavia).
1839. V[csp}crfilio] nilsso7iii Keyserling and Blasius, Wiegmann's Archiv

fiir Naturgesch., 1839, p. 315 (Mountains of Scandinavia. Based

on the V. kuhlii of Nilsson, 1836).

1857. Vesperugo nilssonii Blasius, Saugethiere Deutschlauds, p. 70.

1858. Amhlyotas atratus Kolenati, Sitzungsber. kais. Akad. Wissensch.

Wien, Math.-Naturwissensch. Classe, xxix, p. 252 (Altvater,

Austrian Silesia, alt. 2400-4600 ft.).
1878. Vesperugo burealis Dobson, Catal. Chiropt. Brit. Mus., p. 203.
1894. Vesperugo nillsoni (sic) Rhoads, Reprint Ord's N. Amer. Zoology,

Append., p. 3.
1907. E['ptesicus'] nilsso7ii Miller, Families and Genera of Bats, p. 209,

June 29, 1907.
1910. Eptesicus nilssoni Trouessart, Faune Mamm. d'Europe, p. 23.

Type localiUj. — Sweden.

Geographical distribution. — Continental Europe, from northern
Norway to the Alps.

Diagnosis. — Size medium (forearm less than 40 mm., condylo-
basal length of skull less than 16 mm.) ; colour of upper parts a
rich dark brown, the hairs of back with noticeably contrasted
light tips ; underparts light yellowish brown ; a well defined
line of demarcation along sides of neck.

External form. — In general the external form agrees with
that of Eptesicus serotinus, due allowance being made for the less
robust stature of the smaller animal. Ear relatively Ljnger,

EPTESICUS

235

extending to nostril when laid forward, though of the same
general form as in E. serotinus ; tip less narrowly rounded oft',
and flattened or concave portion of posterior border less con-
spicuous ; tragus relatively shorter and wider, though not
essentially difterent in form, its greatest width nearly equal to
length of anterior border ; transverse striations on inner surface
of conch oljsolete. Wings and feet essentially as in E. serotinus.
Tail slightly longer than in the related animal, extending nearly
to head when laid forward, its terminal vertebra free.

Fur and colour. — In quality the fur resembles that of E.
serotinus except that it is softer and more silky in texture, the
hairs fullj- as long as in the larger animal. In distribution it
shows no special peculiarities ; dorsal surface of uropatagium
thinly furred to about middle instead of on extreme base only.
Colour above a rich daik brown, ranging from burnt-umber nearly
to seal-brown, the hairs everywhere with slightly darker, faintly
slaty bases, those of median dorsal region from crown to base of
tail tipped with light glossy ochraceous-buft' in evident contrast
with ground colour, the light tips most numerous behind shoulders,
and forming a noticeable mantle over middle of back ; under-
parts rather strongly contrasted light yellowish brown, between
the wood-brown and ochraceous-buiF of Ridgway, the basal
portion of the hairs similar to ground colour of back, the yellowish
brown area extending over sides of head and completely encircling
base of ear, the line of demarcation between it and the dark
brown of upper parts sharply defined along sides of neck. Muzzle,
cheeks, ears and membranes blackish.

Skull. — In general the skull difiers from that of Eptesicus
serotinus, apart from its smaller size, in a general tendency to
greater depth, less elongation, and smoother, more evenly
rounded surfaces. Dorsal profile with evident convexity at middle.
Lambda not overhanging ; low, ill-defined
lambdoid crests curving slightly toward point
of contact at middle, the rounded 2:)osterior
outline of the occiput plainly visible behind
them when skull is viewed from above ;
sagittal crest essentially absent. Brain-case
sub-spherical or broadly ovate in outline,
its depth slightly more than half mastoid
breadth ; floor of brain case marked by a
wide lateral groove between each cochlea
and the median line ; auditory bulla? slightly
larger than in E. ,sr, o//hhv. Interorbital fig. 41.

region relatively less constricted than in Epte^icus nU:<xonu.
E. serotinus, but of essentially the same form ; ^at- si^e.

lachrymal swelling present but less noticeable

than in the larger animal. Rostrum rounded off" at sides, with
only the faintest trace of lateral concavities ; narial and palatal
emarginations essentially as in E. serotinus. Palate showing no

236

CHIROPTERA

noteworthy i^eculiai'ities, its general outline less narrow than in
the related species.

Teeih. — Inner upper incisor as in Epfesicus serotinus, but more
robust in proportion to its height ; outer incisor noticeably higher
than in the related species, its apex reaching level of secondary
cusp of larger tooth, its secondary cusp more prominent ; no
marked contrast between crown areas of the two teeth ; each
pair in line of general curve of anterior jjortion of palate, instead
of at right angles to main axis. Lower incisors less crowded
than those of E. serotinus, their imbrication distinct but not
unvisual, the general outline of the row broadly V-shaped ; in form
the individual teeth show no special peculiarities ; /^ without
postero-internal tubercle. Canines and premolars with no special
peculiarities. First and second upper molars essentially as in
Eptesicus serotinus, but disproportion between paracone antl
metacone less evident ; m^ with crown area about two-thirds that
of wi', its longitudinal diameter through metacone slightly more
than half trans^■erse diameter, the mesostyle and metacone well
developed, the second and third commissures more than half as
long as first ; lower molars like those of E. serotinus in form, but
angles in commissures between outer and inner cusps deeper ;
area of second V ^^ '"3 nearly equal to that of first.

Measurements. — Adult male from Grotte de Vallorbe, Vaud,
Switzerland: head and body, 68 • 5 ; tail, 47; tibia, 17; foot, 10;
forearm, 38 • 2 ; thumb, 9 • 8 ; third finger, 68 ; fifth finger, 49.
Forearm in adult male from Upsala, Sweden, 38-4. Three
females from the same locality : forearm, 39, 39, and 39 ■ 6 ;
third linger in the four specimens from Upsala : 66, 68, 70 and
68. For cranial measurements see Table opposite.

Specimens examined. — Fifteen, from the following localities : —

Norway : No exact locality, 1.

Sweden : Upsala, 8 (B.M. and U.S.N.M.) ; Upland, 2 (U.S.N.M.).

Germaky : Wernigerode, Saxony, 2.

Austria-Hungary : Csallokoz-Somorja, Pressburg, 1.

Switzerland : Grotte de Vallorbe, Vaud, 1 (Mottaz).

Itemarlcs. — This species is readily distinguishable from Epfesicus
sn-otinus by its smaller size and by the conspicuous pale tips to
the hairs of back. From Vespertilio murinus, which resembles it
in colour jiattern, the narrow ear, more yellowish (less whitish)
hair-tips on hack, and slightly smaller size distinguish it super-
ficially. Pipistrellus sarii, which often has almost exactly similar
coloration, is a much smaller animal (forearm, 31 to 33 instead
of 38 to 40).

1. Norway. {Collctt.) E. E. Alston (p.) 81. 6. 9. 2.

4 9. Upsala, Sweden. {EoUhoff.) Lord Lilford (p). 11.1.1.39-42.

6, ?. Wernigerode, Saxony, Ger- Dr. W. WolterstorfE 0. 2. 8. 3-4.
many. (c&p).

1. Csallokoz-Somorja, Press- Budapest IMuseum (e.) 94.3.1.14.
burg, Hungary.

EPTESICUS

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Genus VESPERTILIO LinnEeus.

1758. Vespertilio Linnseus, Syst. Nat., i, lOtb ed., p. 31 (nmriims by

tautonymy).
1839. Vesperugo Keyserliug and Blasius, Wiegmann's Archiv fiir Natur-

gesch., 1839, i, p. 312 (part).
1839. Vesperiis Keyserling and Blasius, Wiegmann's Archiv fiir Naturgesch.,

1839, I, p. 313. Sub-genus of Vesperugo (part).

1856. Meteorus Kolenati, Allgem. deutscbe Naturbist. Zeitung, Dresden,

neue Folge, ii, p. 131 (part).

1857. Vesperus, misprinted Vesperugo Blasius, Saugetbiere Deutscblands,

p. 69 (Sub-genus of Vesperugo).
1863. '^ Aristippe Kolenati, Beitrage zur Kenntniss der Pbtbiriomyiarien,

Petersburg, 1863" (part).
1872. Marsipolxmus Peters, Monatsber. k. preuss. Akad. Wissenscb.,

Berlin, p. 260 (Sub-genus of Vesperugo for albigularis = murinus).
1878. Vesperus Dobson, Catal. Cbiropt. Brit. Mus., p. 183. Sub-genus of

Vesperugo (part).
1897. Vespertilio Miller, Ann. and Mag. Nat. Hist., Ctb ser., xx, p. 384,

October, 1897 (part).
1900. Vespertilio Mebely, Monogr. Cbiropt. Hungariaj, p. 219 (part).
1907. Vespertilio Miller, Families and Genera of Bats, p. 209, June 29, 1907.

Type species. — Vespertilio 'murinus Linn^^^us.

Geograpliical distribution. — Forested northern portion of the
Paltearctic region from the Atlantic coast eastward through
Continental Asia.

Characters. — Like Eptesicus, but ear much shortened and
broadened, rostrum flattened above, with deep concavity on
each side between nares and lachrymal region ; nares very large,
extending back nearly half way to interorbital constriction, and
palatal emargination extended so far laterally that its width is
distinctly greater than its depth.

Remarlcs. — The genus Vespertilio as now restricted contains
only two species, V. murinus Linnaeus, and V. superans Thomas,
the former European, the latter Asiatic. The form of the skull
.suggests in certain respects that of Nyctalus, and in others the
North American Lasionj/cteris.

VESPERTILIO MURINUS Linn;eus.

1758. [Vespertilio] murinus Linnaeus, Syst. Nat., i, 10th ed., p. 32 (Sweden).

1819. Vespertilio discolor Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv
(= Neue Ann., i), pt. 2, p. 187 (Vienna, Austria).

1857. Vesperugo discolor Blasius, Saugetbiere Deutscblands, p. 73.

1872. Vesperus [Marsipolsemus) albigularis Peters, Monatsber. k. preuss.
Akad. Wissenscb., Berlin, p. 260 (Type supposed to have been
taken in Mexico ; for its reference to this species see Mebely,
Monogr. Cbiropt. HungariEe, pp. 229, 341, 1900).

1878. Vesperugo discolor Dobson, Catal. Cbiropt. Brit. Mus., p. 204.

1885. Vesperus siculus Daday, Orvos-Termeszettudomduyi Erteseto, Kolozs-
var, X, p. 275 (Homorod-Almas cave, Hungary). See Mebely,
Monogr. Cbiropt. Hungarise, pp. 229, 346, 1900.

VESPERTILIO

239

}

188G. Vesperus siculus Daday, Verhandl. u. Mittheilungen des Siebenbiirg-
ischen Vereinsfiir Naturwissensch. in Hermauustadt, xxxvi, p. 82.

1897. Vcs2)crtilio murinus Miller, Ann. aud Mag. Nat. Hist., 6th ser., xx,
p. 382, October, 1897.

1910. Eptesicus siculus and Vcspertilio murinus Trouessart, Paune INIamm.
d'Europe, pp. 23 and 25.

Type locality. — XJpsala, Sweden.

Oeographical distribution. — Northern and central Europe,
from southern Norway and central Sweden south to the Alps.
One record of its occurrence in England.*

Diagnosis. — General characters as in the genus ; length of
forearm about 43 to 45 mm.

External characters. — General appearance much as in Eptesicus
nilssonii but slightly larger and more robust. Ear extending
nearly to nostril when laid forward, its width when flattened
about li times height from crown ; anterior basal lobe so much
reduced as to suggest the keel of the Molossidse, the anterior
border of conch nearly straight from forehead to broadly rounded
tip ; posterior border shallow ly sinuous-concave from just below
tip nearly to level of meatus ; here it turns forward almost at
right angles and extends nearly to angle of mouth, where it
terminates at a well developed wart, the terminal portion often
forming a slight fold or pocket ; antitragus low and keel-hke,
nearly 2 mm. from margin of conch ; inner surface of conch
slightly papillose, without evident cross ridges ; tragus low,
scarcely rising above level of outer angle of anterior lobe, its
greatest width (slightly above middle) nearly equal to length of
straight or slightly concave anterior border, its tip broadly
rounded, the convexity continuous behind with that of upper
portion of posterior border ; posterior basal lobe obsolete. Wing
rather narrow, the fifth finger exceeding forearm by one-sixth to
one-fifth length of forearm, the membranes thin, extending to
base of outer toe ; third and fourth metacarpals sub-equal, nearly
5 mm. shorter than forearm, fifth about 3 mm. shorter than
third. Leg rather robust ; foot about half as long as tibia ;
calcar slightly longer than tibia and free border of uropatagium,
its keel ill defined, its termination obscure. Tail extending to
between shoulders when laid forward, the last vertebra free.

Fur and colour. — The fur is rather short and dense, less silky
than that of Eptesicus nilssonii, the hairs at middle of back about
7 mm. in length. In distribution it shows no special peculiarities ;
upper surface of interfemoral membrane furred on basal third.
Colour above essentially as in Eptesicus 7iilssonii, but liglit tips to
hairs very pale, almost whitish crcam-bufl", producing a decidedly
" frosted " appearance ; underparts creain-buft", the dusky under
colour showing through on chest and anterior portion of belly ;
line of demarcation on sides of neck sharply defined ; muzzle and
chin dusky ; ears and membranes blackish.

* A single specimen, undoubtedly a straggler, taken at Plymouth.

240 CHIROPTEEA

ShiU. — The skull combines the short, smooth brain-case of
Ej^tesicHS nihsonii with the broad, flat rostrum of E. serotinus, but
differs conspicuously in aspect from both in the very deep,
N>/cfahis-like narial and palatal emarginations. Dorsal profile
essentially straight from nares to rounded, not overhanging
lambda. Sagittal crest obsolete ; lambdoid crest low but evident.
Depth of brain-case at middle decidedly more than half mastoid
breadth ; floor with wide conspicuous slit between cochlea and
basioccipital : a flattened pit-like depression at front of each slit ;
auditory bullae moderately large, their trans-
verse diameter about equal to space between
them. Interorbital region broadly hour-glass
shaped, flattened above ; edge of orbit from
lachrymal region nearly to most constricted
portion distinctly and irregularly swollen.
Dorsal surface of rostrum flattened, with broad,
shallow lateral depressions ; narial emargina-
tion broadly triangular, its apex extending
back to level of lachrymal region ; rostral depth
at front of orbit equal to distance from orbit
"'r' '"" . to front of inner incisor ; anteorbital foramen

\ es'perlilio murvaux. . „

Nat. size. moderately large, over point ot contact between

large premolar and first molar ; lachrymal fora-
men directly behind it on inner rim of orbit. Palate broad and
short, noticeably concave both laterally and longitudinally, the
anterior emargination large, much wider than deep, but extend-
ing back to level of middle of premolar ; posterior extension of
palate parallel sided, its length slightly greater than width
behind molars ; hamulars slightly bent inward ; median spine
well developed. Mandible robust, much deeper at symphysis
than behind tooth-row, the coronoid process low (height less than
least width of posterior section) with gradually sloping upper
border ; angular process rather slender, its extremity slightly
curved inward.

Teeth. — While in general resembling those of the European
species of Eptesicus, especially those of E. nilssonii, the teeth of
Vesjpertilio murinus show certain notable peculiarities. Inner
upper incisor similar to that of Epfesicus serotinus when viewed
from in front, but with well developed postero-basal cusp which
in some specimens rises nearly to half the height of main cusp,
l)roducing a distinctly trifid tooth ; outer incisor with crown area
slightly less than that of inner, the cusp rising nearly to middle
of shaft of inner tooth ; secondary cusp slightly developed ;
position of incisors relatively to tooth-row about as in E. nilssonii.
Lower incisors as in E.nil.'^.'ionii. Upper canine with transverse
diameter slightly greater than longitudinal diameter, the reverse
of the condition in the two species of Ej^tesicus ; lower canine
with same peculiarity in outline of crown though to a less degree.
Upper premolar with longitudinal diameter of crown relatively

VESPERTILIO

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less than in the European species of Eptesicus, and antero-interual
basal cusp better developed : lower premolars more crowded and
compressed than in E. nihsonii, and crown area of first relatively
smaller. Molars as in E. nilssonii except that columnar hypocone
of m^ and m? stands out more prominently from outline of
protocone.

3Ieasiirements. — Three adult males from Denmark : head and
body, 62, 59 and 62; tail, 43, 40 and 44 ; tibia, 17-4, 16-8 and
16 • 8 ; foot, 9 • 6, 9 • 2 and 10 ; forearm, 43 • 4, 44 and 43 ; thumb,
7, 7 '4 and 7' 6; third linger, 73, 76 and 74; fifth finger, o2, 54
and 53; ear from meatus, 15, 15 and 15' 6; width of ear, 16,
16 and 17. Adult female from the same locality: head and
body, 63 ; tail, 41 ; tibia, 16 • 4 ; foot, 9 ; forearm, 45 ; thumb, 8 ;
third finger, 78; fifth finger, 54: ear from meatus, 16; width
of ear, 17. Adult male from Morat, Fribourg, Switzerland:
head and body, 56 ; tail, 42 ; tibia, 16 : foot, 8*8 ; forearm, 45 :
thumb, 5'4 ; third finger, 76 ; fifth finger, 55 ; ear from meatus,
12; width of ear, 12 "4. For cranial measurements see Table,
p. 241.

Specinieyis examined. — Twelve, from the following localities : —

England : Plymouth, 1.

Sweden : No exact locality, 1.

Denmark: No exact locality, 4 (U.S.N.M.).

Germany: Gross Hennersdorf, Saxony, 1; Bavaria, 2 (U.S.N.M.).

AnsTRiA-HuNGARY : Csallokoz-Somorja, Pressburg, 1.

Switzerland : Morat, Fribourg, 1 (Mottaz).

Locality unknown: One; type of aifcir/aiaris Peters (Berlin).

Remarks. — Superficially this bat somewhat resembles Ejjtesicns
nilssoni. It is readily distinguishable, however, by its slightly
larger size (forearm, 43 to 45 instead of 38 to 40), low, rounded
ear, and l)y the whitish rather than yellowish wash on back.

1. Plymouth, Devonshire, Dr. Leach (p). 37. a.

England.
— Sweden. Stockholm Museum (e). 46. 6. 2. 21.

?. Gross Hennersdorf, Lord Lilford (p). 99.1.9.5.

Saxony {W. Baer.)
1. Csallokoz-Somorja, Press- Budapest ^luseum (e). 94. 3. 1. 1-5.

burg, Hungary.

Genus NYCTALUS Bowdich.

1825. Nijctalus Bowdich, Excursions in ^Madeira and Porto Santo, p. 36

(verrucosus).
1829. Pten/gistes Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt,

I, p. 99.
1839. Vesperucjo Keyserling and Blasius, Wiegmann's Archiv fiir Natur-

gesch., 1839, i, p. 312 (part).
1842. Noctulinia Gray, Ann. and Mag. Nat. Hist., x, p. 258 (part ; contained

inoterus = noctula and fulvus = Scotophilus kulilii).

1856. Fanugo Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden,

neue Folge, ii, p. 131 {noctula and leisleri).

1857. Vesperugo Blasius, Siiugethiere Deutschlauds, p. 49 (part).

NYCTALUS 243

1878. Vesperugo Dobson, Catal. Chiropt. Brit. Mus., p. 183 (part).

1893. Noctulinia II. Allen, Proc. U.S. National Museum, xvi, p. 30,

June 13, 1893.
1897. Pterygistes Miller, Ann. and Mag. Nat. Hist., Gth ser., sx, p. 384,

October, 1897.
1899. Euvcspenigo Acloque, Faune de Prance, Mammiferes, p. 32 (part,

included twctula, leisleri, maurus, kuhlii, pipistrellus, and

abramus).

1907. Pterygistes Miller, Families and Genera of Bats, p. 207, June 27,

1907.

1908. Nyctalus Andersen, Ann. and Mag. Nat. Hist., 9th ser., i, p. 434,

May, 1908.

Type species. — Nyctalus verrucosus Bowdicli.

Geographical distribution. — Northern portion of Eastern Hemi-
sphere from the Azores and Madeira to Japan.

Characters. — Dental formula as in Pipistrellus ; skull with
nares extending unusually far back, half way to interorbital con-

a b c d

Fig. 43.
Xi/ctahis maximus (a), N. noctula (b), N. leiilen (c), and X. azoreum (A). Jfat. size.

striction, and with large anterior palatal emargination ; fifth finger
much shortened, scarcely exceeding metacarpal of fourth or third.
Hemarhs. — This genus is well diflerentiated from allied groups
by the peculiar narrowing of the wing due to the shortness of the
fifth finger. Three of the half dozen known species occur on the
continent of Europe, while a fourth inhabits the Azores. The
most obvious character by which the European forms are dis-
tinguished is the size of the skull (see fig. 43).

KEY TO THE EUROPEAN SPECIES OP NYCTALUS.

Condylobasal length of skull more than 17 mm. ; fore-
arm more than 45 mm. ; hairs of back without
dark bases.
Condylobasal length of skull 22 to 23 mm. ; forearm

64 to 68 mm. (Southern Europe) N. maximus, 'p. 244.

,Condylobasal length of skull 17 ■ 4 to 19 • 4 mm. ; fore-
arm 47 to 55 mm. (Distribution general) N. noctula, p. 245.

Condylobasal length of skull less than 17 mm. ; fore-
arm less than 45 mm. ; hairs of back with notice-
able dark bases.
Condylobasal length of skull 15 to 16 mm. ; forearm

39 to 43 mm. (Distribution general) N. leisleri, -p. 252.

Condylobasal length of skull 13 to 14-2 mm. ; fore-
arm 35 to 42 mm. (Azores) N. asorcum, p. 254.

R 2

244

CHIROPTERA

NYCTALUS MAXIMUS Fatio.

1781. ? Vespe^'tilio lasioptcrns Schrober, Siingethiere, pi. lvii b. See

Thomas, Ann. and Mag. Nat Hist., 8th ser., viii, pp. .379-380,

September, 1911.
1827. ?■? Vespertilio ferrufjineus Brehm, Ornis, Heft iii, p. 26 (Renthendorf,

Thiiringen, Germany).
1869. [Vcsperugo noctula] var. viaxima Fatio, Faune Vert. Suisse, i, p. 67.

Co-type in Geneva Museum.
1900. Pterygistes maximus Miller, Proc. Biol. Soc. Washington, xiii,

p. 156, June 13, 1900.
1910. Nydalus maximus Trouessart, Faune JIamm. d'Europe, p. 19.

Ty2)e locality. — Amsteg, Uri, Switzerland.

Geographical distribution. — At present known from a few
localities in Switzerland and Italy.

Diagnosis. — Essentially similar to Nyctalus noctula but much
larger; condylobasal length of skull, 22 to 23 mm. ; forearm, 64
to 68 mm.

External cltarncters. — Except for the conspicuously greater
size and consequent more robust form, there appears to be no
tangible character by which the animal can be distinguished
from N. noctula.

Fur and colour. — In the few specimens examined the furred
area on under surface of wing membrane behind forearm appears
to be better defined than in N. noctula. Colour as in the common
species.

Skull and teeth. — The skull and teeth so closely resemble those
of Nyctalus noctula that in general they differ in their greater
size only. The posterior portion of occiput is, however, more
elevated above base of cranium than in the smaller animal, so
that lower edge of condyle is about on level with anteorbital
foramen and alveolus of canine instead of distinctly below them.
Correlated with this character is a more abrupt I'ising of lambdal
region above level of anterior portion of brain-case, and a less
nearly horizontal truncation of occipital region. Palate appar-
ently more concave longitudinally than in any of the smaller
European species.

Measurements. — Adult male and female from Pisa, Italy :
head and body, 78 and 87 ; tail, 59 and 66 ; tibia, 23 • 6 and 24 ;
foot, 12 and 13 ; forearm, 65 and 68 ; thumb, 11 and 11 ; third
linger, 119 and 123 ; fifth fing(!r, 76 and 77 ; ear from meatus,
21 and 22 ; width of ear, 23 and 24. Adult male from Venice : *
head and body, 92 ; tail, 65; tibia, 26; foot, 14 ; forearm, 67.
Adult female from Amsteg, Uri, Switzerland : head and body, 90± ;
tibia, 21-6 ; foot, 13 ; forearm, 64 ; thumb, 9-4 ; third finger,
114 ; fifth finger, 72. For cranial measurements see Table, p. 250.

Specimens examined. — Five, from the following localities : —
Switzerland: Amsteg, Uri, 1 (Geneva; co-type).
Italy : Pisa, 3 (B.M. and U.S.N.M.) ; Ravenna, 1.

Ninni, Atti Soc. Ital. Sci. Nat., Milano, xxvi, p. 109, 1883.

NYCTALUS 245

Remarks. — Ni/ctalus maximus is readily distinguished among
the bats of Eui-ope by its large size. It is at present a rare and
little known animal.

9 al. Pisa, Italy. Dr. A. Senna (e). 94. 6. 3. 3..

6 al. Ravenna. Florence iluseura (e). 85. 7. C. 9.

i al. Lidth dc Jeude Coll. 67. 4. 12. 337.

NYCTALUS NOCTULA Schreber.

1774. Vcspcrtilio nochda Sohreber, Saugthiere, i, pi. lii ; description, i,

p. 16G, under name : Die Speckmaus (France ; based primarily

on " La Noctule " of Daubenton, Hist. Acad. Royale des Sci.,

Paris, 1759, p. 376. 1765).
1776. Vespertilio lardarius P. L. S. MiiUer, Natursyst. Suppl. u. Regist.-

Band, p. 15 (France; based primarily on Schreber's Speckmaus).
1789. [Vespertilio] magnus Borkenhaut, Syn. Nat. Hist. Great Britain and

Ireland, i, p. 1 (Cambridge, England; based on Pennant, Brit.

Zool., No. 38).
1789. Vespertilio altivolans White, Nat. Hist, and Antiquities of Selborne,

p. 93 (Selborne, Hampshire, England).
1816. ? Vespertilio major Leach, Syst. Catal. Spec. Indig. Mamm. and

Birds Brit. ]Mus. (Willoughby Society reprint, 1882), p. 5 Nomen

nudum : " Great Bat."
1818. Vespertilio proterus Kuhl, Ann. Wetterau, Gesellsch. Naturk, iv

(= Neue Ann., i), pt. 1, p. 41 (Substitute for noctula).
1829. Vespertilio rufesccns Brehm, Isis, p. 643 (Jena, Thiiringen, Germany).
1841. Vespertilio noctula Bonaparte, Iconogr. Faun. Ital., i, lud. Distrib.

(sp. iUustr.).
1841. Noctula serotina Bonaparte, Iconogr. Faun. Ital., i, Ind. Distrib.

(uomencl. moderna).
1844. Vcsp[ertilio] palustris Crespon, Faune M6ridionale, i, p. 22 (marshes

near Nimes, Gard, France).
1857. Vesperugo noctula Blasius, Saugethiere Deutschlands, p. 53.
1869. [Vesperugo noctula] var. minima Fatio, Faune Vert. Suisse, i, p. 58

(Geneva, Switzerland).
1878. Vesperugo noctula Dobson, Catal. Chiropt. Brit. Mus., p. 212.
1897. Ptenjqistes noctula Miller, Ann. and Mag. Nat. Hist., 6th ser., xx,

p. 384, October, 1897.
1910. Nyctalus noctula Trouessart, Faune Mamm. d'Europe, p. 18.

Tjipe locality. — France.

Geographical diMrihutian. — Europe from southern Norway
and central Sweden to the Mediterranean, and from England
eastward into Asia.

Diagnosis. — Condylobasal length of skull, 17 '4 to 19 '4 mm. ;
forearm, 47 to 55 mm. ; general colour dark j'ellowish brown, the
hairs not darker at base.

External characters. — General form robust and heavy. Muzzle
broad, with conspicuous glandular swelling between eye and
nostril, the greatest width across this region decidedly more than
distance between nostril and ear ; nostrils projecting forward and
outward with evident median concavity between them, the orifice
crescentic. Ear short, extending when laid forward about half

246 • CHIROPTERA

way from eye to nostril, its breadth when flattened decidedly
greater than height above crown ; anterior border of conch
abruptl}' convex below, then nearly straight to broadly rounded
off extremity ; posterior border convex throughout, most strongly
at middle ; antitragus long and low, well marked off posteriorly,
its anterior border extending to just below angle of mouth ; inner
surface of con^h finely papillose, without evident cross ridges ;
tragus very short, scarcely rising above level of anterior base of
conch, much wider above than below, its greatest width about
equal to height ; anterior and posterior borders concave, the
anterior more so than posterior, their length about equal to that
of expanded upper portion, which is more abruptly rounded
anteriorly than posteriorly. Wing long and slender, the iifth
finger exceeding forearm by only ^ to ^ length of latter, the
membrane leathery and opaque, joining leg at ankle ; third and
fourth metacarpals sub-equal, the third slightly the longer and
about equal to forearm, fifth abruptly shorter by slightly more
than one-fifth forearm. Leg short and strong, the broad foot
about one-half as long as tibia ; calcar 1 ^ to twice as long as
tibia or as free border of interfemoral membrane, its distal
termination obscure but basal portion strong, well defined and
with keel about 2-5 mm. wide supported by a well defined
thickening and terminating in an abrupt convexity on side
nearest heel. Tail rather short, extending to between shoulders
when laid forward, the tip of last vertebra (about 2 ram.) free.

Fiu- and colour. — Fur dense and velvety, the hairs at middle
of back only about T) mm. in length ; on dorsal surface of wing
it extends to line joining knee with middle of humerus, on ventral
surface to line from elbow to knee ; beyond this line it spreads
thinly on under surface of antebrachial membrane and also
behind forearm and across bases of metacarpals along an area
about 10 mm. wide. Interfemoral membrane furred both above
and below at extreme base only. Colour of upj^er parts a rather
dark yellowish brown, near the wood-brown and cinnamon of
Ridgw ay, the hairs showing a faint clouding of prouts-brown in
certain lights, their basal portion a light dull isabella-colour ;
underparts scarcely difierent from back, though usually a little
lighter and more dull ; muzzle and cheeks dusky ; ears and
membranes blackish.

Skull. — General aspect of skull broad and robust, rather high
posteriorly, low anteriorly, with conspicuous narial emargination.
Dorsal profile rising rather rapidly from incisors to lambda, with
slight convexity over lachrymal region and a slighter though
more abrupt concavity just in front of lambda. Ventral profile
elevated posteriorly. Brain-case ovate, but with wide mastoid
and paroccipital region and squarely ti-uncate occiput which
together produce a distinctly truncate-cuneate outline, slightly
though evidently wider than long ; depth at middle distinctly
more than half mastoid breath : sagittal crest low but evident,

247

I

Nyctalus noctiUa.
Nat. size.

the region on each side of it flat or sHghtly depressed ; lambdoid
crest high, curved abruptly forward at point of junction with
sagittal crest ; floor of brain-case with small but evident lateral
pits, well defined anteriorly but communicating posteriorly with
vacuity between cochlea and basioccipital ; auditory bulhe well
developed but of moderate size, the transverse diameter al>out
equal to distance between bulhe. Interorbital region deeply
constricted, short hour-glass shaped, the lachiymal region widen-
ing abruptly to a breadth nearly equal to that of brain-case, and
forming a slight though evident tuber-
cular projection over anterior rim of orbit ;
rostrum squarish, somewhat narrower
anteriorly than posteriorly, the oval or
ovate narial emargination extending back
to level of lachrymal foramen ; rostral
depth at front of orbit about equal to
distance from orbit to front of inner
incisor ; anteorbital foramen small, over
space between large jiremolar and first
molai', lachrymal foramen slightly above
and behind it, on orbital rim. Palate
rather short, owing to the large size of
the anterior emargination, the posterior
edge of which is on level with middle of
large premolar, rather evidently concave

laterally, less so longitudinally. Posterior extension of palate
nearly parallel sided, its width at level of posterior edge of third
molar slightly less than length ; hamulars barely turned inward ;
median spine large, acute. Mandible robust, noticeably deeper
at symphysis than immediately behind tooth-row ; posterior
portion rather low and long, the height of coronoid process
above alveolus less than least length, the upper border sloping
gradually from coronoid to articular jsrocess ; angular process
moderately long, about on level with alveolar line, its main
axis directed rather abruptly outward, its distal extremity
oblicjuely widened.

Ti/efh. — Relatively to size of skull the teeth are large, though
the cusps are rather low. Inner upper incisor slender, with
slightly developed cingulum, its crown area barely one quarter
that of canine, the nearly terete shaft directed obliquely inward,
its apex extending slightly beyond level of cingulum of canine,
its .secondary cusp small but evident, situated on postero-external
surface of shaft ; outer upper incisor with crown area distinctly
greater than that of inner tooth, the shaft deeply concave, its
concavity directed outward and backward, the main cusp lying close
against secondary cusp of inner tooth, the well-defined se-.-ondary
cusp relatively much larger than that of inner incisor, cingulum
M-ell developed, bearing a small but distinct antero-external cusp
on opposite side of concavity from secondary cusp. The two

248

GHIROPTERA

incisors are closely crowded against each other ; outer tooth
separated from canine by a very narrow space. Lower incisors
rather strongly imbricated, forming a very broadly U'Stiap^d
row between canines, the crowns of /^ and /., compressed, trilid,
that of i^ sub-terete, flattened anteriorly, with two low posterior
tubercles in addition to the three cusps corresponding to those of
other teeth. Upper canine robust, with well developed anterior
and posterior cutting edge, the shaft triangular in cross-section ;
cingulum well developed but without true secondary cusps.
Lower canine with posterior surface of shaft strongly concave,
the well developed cingulum forming a postero-internal basal
cusj) and a conspicuous secondary cusp near middle of antero-
internal border of shaft ; diameter of crown distinctly less than
distance between canines. Anterior upper premolar minute, closely
wedged in space between canine and posterior premolar, its cross-
section less than half that of incisors, its crown reniform in
outline, its cusp low but distinct ; posterior upper premolar with
crown area somewhat more than half that of first molar, the
protocone well developed, nearly as high as in molars, the main
cusp rather high but much shorter than canine (measured along
cingulum), its two cutting edges well developed. Lower pre-
molars closely crowded, their ci-owns about equal in cross-section
and somewhat more than half that of canine, the shaft of first
lower and more robust than that of second and about half as
high as canine ; cingulum well developed, in each tooth forming
a small but evident antero-internal and postero-internal cusp.
First and second upper molars with crowns rather broad on
inner side, the posterior emargination slight ; protocone robust
but low ; hypocone small but well developed and distinct from
commissure of protocone ; outer cups without sjiecial peculiarities,
the styles low but well developed, the VVP'ittern normal, //t^ with
crown area distinctly more than half that of m}, its longitudinal
diameter through metacone about half transverse diameter, its
three cusps, two styles and three commissures well developed,
but proportion between paracone and metacone reversed as
co)npared with other molars. Lower molars with unusually
robust hypoconid and entoconid, producing a strong contrast in
size between the anterior and posterior V in «*i and in.^ and
approximate equality in m-^ ; cingulum well developed but not
forming basal cusjd behind entoconid.

Measurements. — Adult female from Kew Gardens, Surrey,
England: head and body, 71 ; tail, 51 ; tibia, 18; foot, 10;
forearm, 52 ; thumb, 9 ; third finger, 94 ; fifth finger, 58 : ear
from meatus, 15; width of ear, 16. Adult female from Herrn-
hut. Saxony, Germany : head and body, 79 ; tail, 53 ; tibia, 19 ;
foot, 10' 6; forearm, 53; thumb, 9*4; third finger, 97; fifth
finger, 60 ; ear from meatus, 17 ; width of ear, 17. Two adult
males from Pisa, Italy : head and body, 70 and 71 ; tail, 49 and
52-6 ; tibia, 18 and 19 ; foot, 10 and 11 , forearm, 51 and 53 :

NYCTALUS 249

thumb, 8 and 9 ; third finger, 92 and 97 ; fifth finger, 56 and 59 ;
ear from meatus, 14 "6 and 16 ; width of ear, 14 and 15 "6. Two
adult females from the same locahty : head and body, 74 and 75 ;
tail, 50 and 53 ; tibia, 18 '6 and 18 ; foot, 10 and 11 ; forearm,
51-6 and 52 ; thumb, 9 and 10 ; third finger, 93 and 97 ; fifth
finger, 57 and 58; ear from meatus, 16 and 16; width of ear,
16 and 16. Two adult females from Corinth, Greece: head and
body, 69 and 76; tail, 51 and 54; tibia, 18 and 19; foot, 11
and 10 ; forearm, 52 and 54 ; thumb, 9 • 4 and 9 • 4 ; third finger,
96 and 102 ; fifth finger, 58 and 62 ; ear from meatus, 15-6 and
16 ; width of ear, 15 and 16. For cranial measurements see
Table, p. 250.

Specimens examined. — One hundred and fourteen, from tlie following
localities : —

England : Bowdon, Cheshire, 1 ; Stoke, Staffordshire, 1 ; Oundle,
Northamptonshire, 4 ; Bedfordshire, 1 ; Trumpington, Cambridgeshire, 1 ;
Cambridgeshire, no exact locality, 1 ; Henley-on-Tliames, Oxfordshire, 1 ;
Sandringham, Norfolk, 1 ; Kingsbury, Middlesex, 1 ; Chelmsford, Essex, 1 ;
Wcllyn, Hertfordshire, 1 ; Tring, Hertfordshire, 3 ; Wandsworth Common,
Surrey, 1 ; Shalford, Surrey, 2 ; Earlsfield, Surrey, 5 ; Kew Gardens,
Surrey, 1 (U.S.N.M.) ; Pulham Park, London, 1 ; Eastwell, Kent, 2 ; St.
Leonards, Sussex, 1 ; Parnborough, Hampshire, 1 ; Wareham, Dorset-
shire, 1.

Holland: Oosterbeek, Guelderland, 1.

GEEJtANY: Moritzburg, Saxony, 3 (U.S.N.M.); Herrnhut, Saxony, 2
(B.M. and U.S.N.M.) ; Niesky, Silesia, 1 ; Schwarzburg, Thiiringen, 2 ;
Strass, near Burgheim, Bavaria, 3 ; Bavaria, no exact locality, 8 (U.S.N.M.) ;
Stuttgart, 1.

Austbia-Hungary : Csallokoz-Somorja, Pressburg, Hungary, 2.

RouMANiA : Gageni, 1 ; Bustenari, 1 ; no exact locality, 1 (U.S.N.M.).

Switzerland : Geneva, 4 (Geneva and Mottaz) ; St. Margarethen,
Appenzell, 8 (B.M. and U.S.N.M.).

Spain : Seville, 4 ; no exact locality, 2.

Italy : Pisa, 18 (U.S.N.M.) ; Plorence, 4 (Mottaz) ; Vallombrosa, near
Florence, 1 (U.S.N.M.) ; Siena, 1 ; Rome, 2.

Greece : Corinth, 12 (U.S.N.M.).

Bemarks. — This is one of the most widely distributed and
abundant of the larger European bats. From Eptesicits serotinus,
the only species with which it is likely to be confused, it is easily-
distinguished by the narrow wing, short, recurved tragus, and
low, rounded ear.

6. Aberia, Merionethshire, Col. Yerbury (c & p). 11. 1. 3. 26.

Wales.
9. Bowdon, Cheshire, Eng- G. Barrett-Hamilton 11. 1. 2. 100.
land. (T. A. Coivard.) (p).

jll.l. 1. 12G-
2 c5, 2 9. Oundle, Northampton- Lord Lilford (p). 127.

shire. 11. 1. 1. 129-

[ 130.
9. Bedfordshire. J. S. Elliot (c & p). 11. 1. 8. 391.
1. Trumpington, Cambridge- G. Barrett-Hamilton 11. 1. 2. 101.
shire. (p).
9 al. Cambridgeshire. Rev. L. Jenyns (c & p).

250

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CHIKOPTERA

Henley-on-Thames, Ox- Heatley Noble (c & p).
fordshire.

6 st. Sandringham, Norfolk.

?. Chelmsford, Esses.

?. Wellyn, Hertfordshire.

2 (5, ? st. Tring, Hertfordshire.

2 S. Shalford, Surrey.

3 d, 2 9. Earlsfield, Surrey.

<5st.
(5,9.

Fulham Park. Loudon.
Eastwell, Kent.

St. Leonards, Sussex.

Farnborough, Hampshire.

Wareham, Dorset.

Oosterbeek, Guelderland,
Holland.

Niesky, Silesia, Germany.
{W. Baer.)

Schwartzburg, Thtiringen,
200 m.

St r ass, Burgheim,
Bavaria. {Korhitz.)

Stuttgart, Wurtemburg.

Csallokoz-Somorja, Press-
burg, Hungary.
6. Gageui, Prahova, Kou-

mania. {W. Dodson.)
9. Bustenari, Prahova. (IT.
Dodson.)
2(5,3 9. St. Margaret hen,
Appenzell, Switzer-
land. (E.H.ZoUikofcr.)

Seville, Spain. {A. Ruiz.)

Spain,

3 6.
1 al.

4al.
2al.

i juv. al. Rome.

H.M. King Edward VII.

il. Christy and L. E.

Thompson (c & v).
C. H. B. Grant (c & p).
Hon. N. C. Rothschild

(c & p).
W. R. Ogilvie - Grant

(c & p).
C. H. B. Grant (c & p).

J. Saunders (c & p).
C. H. Caton - Haigh

(c & P).
R. Butterfield ^c & Pi.
H. S. Scott (c & p).
W. T. Blanford (c & p).
0. Thomas (c & p).

Lord Lilford (p).

Lord Lilford (p).

Lord Lilford (p).

Dr. A. Giinther (p). .
Budapest Museum (e).

Lord Lilford (p).

Lord Lilford (p).

0. Thomas (p).

Seville Museum (e).
Lord Lilford (p).
Florence Museum (e)

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NYCTALUS LEISLERI Kuhl.

1818. Yc&pcrtilio leisleri Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv

(= Neue Ann., i), pt. 1, p. 46.
1818. Vcspcrtilio dasycarpos Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv

(= Neue Ann., i), pt. 1, p. 49 (Alternative name for leisleri).
1839. Vcspcrtilio pachygnathus MichaheUes in Wagner, Schreber's Saug-

thiere, Suppl.', i, pi. lv b (Dalmatia. See Fitzinger, Sitzungsber.

kais. Akad. Wissensch. Wien, Math.-Naturwiss. Classe. lxii,

Abth. I, p. 222, 1870).
1857. Vesperugo leisleri Blasius, Saugethiere Deutschlands, p. 66.
1878. Vesperugo leisleri Dobson, Catal. Chiropt. Brit. Mus., p. 215.
1910. Nyctalus leisleri Trouessart, Faune Mamm. d'Europe, p. 19.

Tijjte localUii. — Hanau, Hesseu-Nassau, Germany.
Geographical distribution. — Central Europe, west to Ireland.
Diagnosis. — Like Nyctalus noctula but not so large ; condylo-

NYCTALUS 253

basal length of skull, 15 to 16 mm. ; forearm, 39 to 43 mm. •
basal portion of fur conspicuously darkened. '

Cohmr.— The colour is usually a brown, darker than in JV
nociula, nearly the prout-brown of Ridgvvay, with or without -i
lighter, wood-brown tinge, and usually showing drab or isabelk
reflections in certain lights, the underparts not so dark as back •
hairs everywhere blackish (seal-brown) through basal half Eai-s
and membranes (dry) blackish. Two skins are a pale bufty wood-
brown throughout except for the usual dark bases of the hairs

Slcull and teeth.—Except for its smaller size and more delicate
structure the skull does not differ appreciably from that of
Nyctahis noctula. Teeth essentially as in the larger species but
crown area of upper incisors nearly equal, and lower incisor row
forming a deeper, almost V-shaped convexity.

Measurements.— Externiil measurements of adult male from
Co. Wicklow, Ireland : head and body, 60 ; tail, 39 • tibia 16 • "^ ■
foot, 8-2 ; forearm, 42 ; third finger, 78 ; fourth finger, 48 • elv
from meatus, 13. Adult female from Co. Armagh, Ireland •
head and body, 63 ; tail, 42 ; tibia, 16-6 ; foot, 8 ; forearm 4"' •
third finger, 76 ; fifth finger, 49 : ear from meatus, 14 • width
of ear, 14. Two adults from Welford, Warwickshire, Ent^knd •
forearm, 40 ■ 4 and 41 • 4. Adult from Meiringen, Bern, Switzer-
land : forearm, 40. For cranial measurements see Table
p. 255. '

Spcciviens examined. ~^\e\m, from the following localities •—
Ieeland: Belfast, 1; Bray, Co. Wicklow, 2- Co Armacrh ">
England: Mexbro', Yorkshire, 1; Welford, Warwickshire! 2 ■ Cleave
Priory, 1 ; no exact locality, 1. ' ^^^"'^'^

RouMANiA : Bustenari, Prahova, 1.
Switzerland : Meiringen, Bern, 1.

JJemarfc.— Though its smaller size is the most obvious
character of this species as compared with JV. noctida the
bicolored fur, the difierent relative sizes of the upper incisors
and the more abrupt convexity of the mandibular incisor series
would by themsehes be quite sufficient to distinguish it.

9. Belfast Ireland. G. Barrett-Hamilton (p.) 1. 3. 15 1

6 a . Bray, Wicklow. J. E. Harting (c & p).' gO. 2. 14 1

9a. Armagh. Dr. G. E. Dobson (c & p). 89.11.12.5

2 9 wTf?!'^ w •,.,• R-M.Barrington(c&P). 74.5.28.8.'

2y. Welford, Warwickshire, Tomes CoUection 7 1 1 SSfi-

England. 'sS7

6. Cleeve Priory, Warwick- Tomes Collection. 7. 1. 1. 388.

1. England. Dr. Leach (p). 63 a

6. Bustenari, Prahova, 840 m. Lord Lilford (p) 44fifi

Roumauia. (ir.Z>o(/.>,o«.) *• ^. o. o.

1. Meiringen, Bern, Switzer- Tomes Collection. 7 1 1 sqq

land. I. J.. J., ooy.

254

CHIROPTERA

NYCTALUS AZOREUM Thomas.

1901. Pferygistes azoreum Thomas, Ann. and Mag. Nat. Hist., 7th ser., viii,
p. 33, July, 1901. Type in British Museum.

Type locality. — St. Michael, Azores.

Geograjjhical description. — Azores.

Diognosis. — Smaller than Nyctaliis leisleri (condylobasal length
of skull, 13 to 14' 2 mm. ; forearm, 35 to 42 mm.) ; colour darker
than in the Continental European species.

External characters. — Except for its conspicuously smaller
size Nyctalus azoreuni does not differ appreciably in external
features from N. noctula. The fifth finger shows a tendency to
be less shortened relatively to length of forearm, but this is appa-
rently not constant enough to be regarded as a specific character.

Colour. — Upper parts jjrout-brown, the hairs with darker,
ill-defined slaty basal area, and usually with lighter tips, the
lighter colour ranging in diflerent individuals from raw-umber
through wood-brown to a dull light buft', and usually producing
a noticeable wash from shoulders backward, particularly near
edge of membranes ; underparts not essentially different from
back ; muzzle, cheeks, ears, and wings blackish.

Skull and teeth. — The skull does not differ appreciably in form
from that of N. noctula, though it is perhaps slightly narrower
and deeper, and the surface is more smoothly rounded off;
lambdoid crest slightly developed. Teeth essentially as in the
larger species, but transverse diameter of m^ relatively greater,
crown area of upper incisors nearly equal, and lower incisor row
forming a more abrupt convexity.

Measurements. — Type (adult male, St. Michael) : head and
body, 54 ; tail, 42 ; tibia, 17 ; foot, 7 -7 ; forearm, 37 ; thumb, 6 ;
third finger, 62; fifth finger, 43 • 7 ; ear from meatus, 12. Adult
male from Terceira : head and body, 50 ; tail, 43 ; tibia, 1 6 ■ 6 ;
foot, 7 ; forearm, 42 : thumb, 6 ; third finger, 72 ; fifth finger,
48 ; ear from meatus, 12 ; width of ear, 1 1. For cranial measure-
ments see Table, p. 255.

Specimens examined. — Seventeen, from the following localities in the
Azores :— Terceira, 4 (B.M. and U.S.N.M.) ; above Magdalena, Pico, 5 ; St.
Michael, 6 (B.M. and U.S.N.M.) ; St. George, 2.

Remarks. — The Azorean Nyctalus is slightly more differentiated
from N. leisleri than the Continental forms are among themselves,
since it differs from the other members of the genus rather
noticeably in colour as well as in size. The specimens show
considerable variation in the extent of the light wash on upper
parts caused by the pale tips to the longer hairs, but this appears
to be independent of locality.

6 al. St. Michael, Azores. F. Du Cane Godman 65. 10. 2. 1.

(c & p). {Type of species.)
2d, 2 9. St. Michael. (W. R. 0. Hon. W. Rothschild 3.6.5.1-3,7.
Grant.) (i").

255

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,, not worn.

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,, not worn.
,, slightly worn.

„ not worn.

,, slightly worn.
,, moderately worn.

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Number.

90. 2. 14. 1
74. 5. 28. 9
98. 11. 12. 5
1. 3. 15. 1
7. 1. 1. 386
7. 1. 1. 388
7.1.1.389

G. 3. 5. 4

6. 3. 5. 5

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60645

3. 6. 5. 8

3. 6. 5. 9

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England: Welford,
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Switzerland : Meirii

N. azoreu

Azores: Terceira

San Jorge
Pico' .

St. Michael

256 CHIROPTERA

6,29. Reguinho, Terceira. Hon. W. Rothschild 3.6.5.4-6.
{W. R. O. Grant.) (p).

2^,3?. Magdalena,Pico. (T7.B.0. Hon. W. Rothschild 3.6.5.10-14.
Grant.) (p).

6,9. Calheto, St. George. Hon. W. Rothschild 3.6.5.8-9.
(TF. B. 0. Grant.) (p).

Genus PLECOTUS Geoffroy.

1816. ? Macrotiis Leach, Catal. Spec. Indig. INIamm. and Birds Brit. Mus.
(Willoughbv Society reprint, 1882), p. 1. Nomen nudum : "Euro-
pean Longear, Macrotiis eiirojuieus."

1818. Plecoiits Geofiroy, Description de I'Egvpte, ii, p. 112. For date see
Sherborn. Proc. Zool. Soc. London,' 1897, p. 288.

1857. Plccotus Blasius, Saugethiere Deutschlands, p. 38.

1878. Plecotus Dobson, Catal. Chiropt. Brit. Mus., p. 177 (part).

1907. Plecotus Miller, Families and Genera of Bats, p. 224, June 29, 1907.

Type species. — Vespertilio auritus Linnajus.

Geographical disfrihution. — Temperate Europe, Asia and
northern Africa.

Characters. — Dental formula : * ^^^, c Jf?, jj»i 70,, m ^^ = 36.
Skull with large, elongate and rounded brain-case, and slender
but normally formed rostrum ; auditory bulliB larger than in any
other European member of the family, their greatest diameter
equal to about three times the distance between them. Ears
very large, much longer than head, joined across forehead ;
no.strils opening upward, their orifices continued backward by
slit-like prolongations, the wart-like outgrowths on muzzle not
specially prominent.

Bemarks. — The genus Plecotus is at once recognizable among
European bats by the very large ears, joined across forehead.
In the only other European genus with 36 teeth {Miiiiopterus)
the ears are unusually short. Seven species are known, one of
which occurs in Europe.

PLECOTUS AURITUS Linnseus.

1758. Vespertilio auritus Linnseus, Syst. Nat., i, 10th ed., p. 32 (Sweden).

1816. ? Macrotus europieus Leach, Catal. Spec. Indig. Mamm. and Birds
Brit. Mus. (Willoughby Society reprint, 1882), p. 1. Nomen
nudum: " European Longear, Macrotus europpeus."

1818. Plccotus auritus Geoffrey, Description de I'Kgypte, ii, p. 118.

1825. Vcsiwrlilio otus Boie, Isis, p. 1206 (Copenhagen, Denmark).

1826. Vespcrtilio cornutus Faber, Isis, p. 515 (Jutland, Denmark).

1827. Plecotus communis Lesson, Man. de Mammal., p. 95 (France).

1828. P[lecotus'] brevivianus Jenyns, Trans. Linn. Soc, London, xvi, p. 55

(Grunty Fen, Isle of Ely, Cambridgeshire, England).

1829. Plecotus vulgaris Desmarest, Faune Frau(jaise, p. 18 (Prance).

PLECOTUS 257

1829. [VcsjKrtilio am-itus] y auslrincus Fischer, Synops. ]\Ianiin., p. 117.

Based on the "second variety" of Desmarest, Diet, des Sci. Nat.,

LViii, p. 51, 182'J (Vienna, Austria).
1832. riccotus velatus I. Geoffroy, Gu^rin's Mag. do Zool., ii, CI. i, pi. 2,

p. 5 (not numbered), footnote (Name api)licd by lapsus calami to

tlie common Plccotus of England).
1810. Plrcotiis megalotos Schinz, Europ. Fauna, i, p. 19 (Synonym of

auritus ; lirehm cited as authority).
1857. I'lecotus auritus Blasius, Siiugethiere Deutschlaiids, p. 39.
18G0. Plccotus kirschbanmii Koch, Achter Ber. Oberhess. Gesellsch.

Natur.- u. Heilkunde, Giesson, p. 40, May, 1860 (Dillenburg,

Oberhessen, Germany).
1863. [Plccotus auritus] var. ttjinis Koch, Jahrb. des Vereius fiir Natur-

kunde im Herzogthum Nassau, xviii, p. 406 (Wiesbaden, Nassau,

Germany).
1863. [Plccotus auritus] var. montanus Koch, Jahrb. des Vercins fiir Natur-

kunde im Herzogthum Nassau, xviii, p. 406 (Westerwald, Nassau,

Germany).
1863. [Plccotus auritus] var. brcvipes Koch, Jahrb. des Vereins fiir Natur-

kunde im Herzogthum Nassau, xviii, p. 407 (Substitute for kirscli-

bauniii Koch).
1878. Plccotus auritus Dohson, Gatal. Chiropt. Brit. Mus., p. 178.
1910. Plccotus auritus Trouessart, Faune Mamm. d'Europc, p. 12.

Type locality. — Sweden.

Geographical distribution. — From Ireland ea.stward into Asia,
and from the Mediterranean north to the Scandinavian Peninsula.

Diagnosis. — Characters as in the genus ; auditory bulhe
not excessively enlarged ; forearm about 37 mm.

External characters. — General form slender and delicate, the
most conspicuous feature the greatly enlarged ears, the super-
ficial area of which together when outstretched is considerably
greater than that of entire body. Muzzle rather nari'ow, its
width at middle about equal to distance from tip to point on
forehead at middle of base of joining membrane of ears ; glandular
swellings not conspicuous, but extending up over sides to median
line behind nostrils, and backward to eye. Nostrils opening
upward and slightly foi'ward, their orifices crescentic with slit-
like posterior elongation ; space between nostrils flat, crossed
at middle by a transverse groove with overhanging anterior edge.
Lips moderately full, smooth. Ear very large, extending about
20 mm. beyond tip of muzzle when laid forward, its length fi-om
meatus nearly equal to that of forearm : general outline of conch
a simple, rater elongate oval, the anterior and posterior borders
evenly convex, the tip rather narrowly rounded ; anterior bases
joined across forehead, the membrane at point of junction about
2 • 5 mm. high ; outline of anterior border broken about 3 mm.
above base by conspicuously projecting, evenly rounded lobe
2 mm. long by 3 mm. wide at base ; antitragus scarcely indicated,
the posterior border of conch terminating abruptly a little behind
level of eye ; inner surface of conch with about twenty faint
cross striations ; joining membrane thinly furred ; a hairy line

s

258 CHIROPTERA

parallel with anterior border, and minute scattered tufts on other
parts of inner surface ; tragus simple, erect, about half as high
as conch, the anterior border straight below, slightly convex
above, the tip narrowly rounded, the posterior border faintly
concave above, distinctly convex below, greatest width (through
anterior base) ecjual to a little less than half length of anterior
border ; posterior basal lobe well developed though not large.
Wing broad, the fifth finger exceeding forearm by more than
t)ne-third length of latter, the membrane thin and semi-ti'ans-
parent, joining foot at base of outer toe ; third and fourth meta-
carjials equal, about 2 mm. shorter than forearm, fifth slightly
shorter than fourth ; leg slender ; foot about half tibia ; calcar
slender but sharply defined, equal in length to tibia and to free
Ijorder of inteiiemoral membrane, its distal termination marked
by a slight lobe, its keel obsolete. Tail extending about to point
of juncture of ears when laid forward, the extreme tip (about
1 • 5 mm.) free.

ShuU. — Among the European members of the order the
skull of Plecotus is noticeable for its large, rather low, smoothly
rounded brain-case, relatively small, weak rostrum, and much
inflated auditory bulliie. Dorsal profile rising gradually from front
of nares to middle of anterior portion of brain-case, with slight
concavity at interlachrymal region, then falling gradually to
low, strongly convex, slightly overhanging lambda ; ventral
profile scarcely elevated posteriorly. Brain-case large but rather
low, the depth at middle only a little more
than half mastoid breadth, the general out-
line when viewed from above squarish with
rounded occipital projection and abrupt
anterior constriction, the surface smoothly
rounded, with slightly indicated sagittal ridge
anteriorly and very low, short lambdoid crest,
the latter situated unusually close to upper
edge of foramen magnum ; an irregular area
in supramastoid region thickly sprinkled with
small vacuities ; base of brain-case smooth,
without sj^ecial features, a narrow groove at
„, V" ' ■. each side of basioccipital ; auditory bulla

Nat. size. relatively larger than in any other European

bat, its diametei- about three times the
space between Inilhe. Interorbital constriction narrow, scarcely
hour-glass shaped, the anterior upper rim of orbit with narrow,
slightly jirojecting edge ; lachrymal region scai'cely half as wide
as brain-case. Rostrum both narrow and short, the narial
emargination, though not unusually large, extending slightly
more than half way back to lachrymal level, its general outline
oval, sometimes flattened posteriorly ; dorsal surface of rostrum
with slight median crease, the sides evenly rounded off ; rostral
depth at front of orbit about equal to distance from orbit to

PLECOTUS 259

aKoulus of inner incisor ; auteorbital foramen small, close to I'ini
of orbit, its orifice over parastyle of first molar ; lachrymal fora-
men slightly above and behind it, on inner edge of orbit. Palate
rather broad and short, strongly concave both laterally and
longitudinally, the anterior emargination small, scarcely extend-
ing back to level of middle of canine ; posterior extension of
palate squarish, slightly longer than wide, with slightly developed
median spine ; mesopterygoid space somewhat wider than deep.
Zygoma slightly but evidently expanded at middle. Mandible
slender, but noticeably deeper at symphysis than behind tooth-
row ; coronoid process moderately high, the upper margin of
posterior section of mandible oblique-concave ; angular process
ratlier heavy, scarcely or not expanded at tip.

Teeth. — The teeth are rather small relatively to size of skull.
Inner upper incisor about half as high as canine, with large
secondary cusp slightly above middle of shaft, the outline of
crown oval, its long axis in line of tooth-row ; outer upper incisor
about half as high as inner, its apex a little below secondary cusp
of larger tooth, its secondary cusp well developed, on inner side, the
outline of crown ovate, with long axis perpendicular to tooth-row
and narrower extremity directed inwards. Lower incisors closely
crowded but slightly imbricated, the row broadly (J shaped ; the
crown area increases regularly from first to third, by addition to
posterointernal portion ; cutting edge obscurely trifid ; /g with
well developed postero-internal tubercle. Upper canine small,
scarcely higher than main cusps of molars, its shaft nearly terete,
with slightly developed posterior cutting edge, on each side of
which lies an evident groove ; cingulum complete, without cusps.
Lower canine small, slightly exceeding molars in height, its
shaft concave posteriorly, flattened interiorly, evenly convex
antero-externally ; cingulum well developed, foi'ming a prominent
antero-internal secondary cusp. Anterior upper premolar
perfectly in tooth-row, about as large as outer incisor, though
more robust, its crown sub-terete, about one-third that of canine
in basal area, its shaft with evident posterior cutting edge ; large
premolar with crown area about two-thirds that of first molar,
the inner side very narrow and without crushing surface, the
antero-internal cusp slender but well developed. Crown area of
anterior lower premolar abovit half that of canine, that of
posterior premolar nearly ecjual to that of canine, that of middle
premolar a little more than half that of first ; first sub-terete,
second oval with long diameter of crown lying across tooth-row,
third scjuarish with antero-external corner rounded oflf ; cusp of
first about half as high as canine, that of second a little shorter,
that of third a little longer ; cingulum well developed but not
forming true secondary cusps. Upper molars narro\\' internally,
the jirotocone with rather short base, the hypocone absent or
barely indicated ; metacone decidedly higher than paracone ;
styles and commissures well developed, though mesostyle of j«'^

s 2

260

CHIKOPTERA

does not extuiul outward to level of parastyle and nietastyle ;
Wpattern normal ; m-' with three cusps and three commissures,
its crown area about half that of m'-', its transverse diameter
through metacone about half length of anterior border. Lower
molars with no special peculiarities except that protoconids and
inner cusps are unusually high and slender ; ciugulum forming a
barely indicated postero-internal cusp behind entoconid ; second
triangle of riu much narrower than tirst, but about equal to it
in area.

3Icnsurcmeiiis. — Adult female from Chelmsford, Essex,
England: head and body, 42; tail, 45; tibia, 17; foot, 9*6;
forearm, 39 ; thumb, 7 ; third hnger, 69 ; fifth finger, 52 ;
ear from meatus, .'56 ; width of ear, 23 ; tragus, 19. Two adult
males from Silos, Burgos, Spain: head and body, 50 and 51 ;
tail, 46 and 46 ; tibia, 18 and 17 ; foot, 8 and 8-4 ; forearm,
41 and 38; thumb, 7' 2 and 7 •4; third finger, 72 and 70; fifth
finger, 57 and 53 ; ear from meatus, 35 and 37 ; width of ear,
26 and 26 "4 ; tragus, 17 and 18. Adult female from La Granja,
S(•go^ ia, Spain : head and body, 45 ; tail, 44 ; tibia, 20 ; foot, 9 ;
forearm, 40 ; thumb, 9 ; third finger, 72 ; fifth finger, 53 ; ear
from meatus, 37 ' 4 ; width of ear, 23 ; tragus, 17 ■ 6. Adult female
from I'lorence, Italy : head and body, 49 ; tail, 48 ; tibia, 19" 2 ;
foot, 9; forearm, 39-4; thumb, 7-8; third finger, 69; fifth
finger, 54 ; ear from meatus, 34 ; width of ear, 23*4 ; tragus, 17.
For cranial measurements see Table, p. 262.

Specimens examined. — Eighty, from the following localities : —

Scotland : Montrose, Forfar, 4.

Ireland : Antrim, 1 ; Co. Longford, 2.

England: Alderley, Cheshire, 1; Bowdon, Cheshire, 1; Diss, Norfolk, 1;
Winfarthing, Norfolk, 1 ; Woburn Sands, Bedfordshire, 1 ; Pembrokeshire,
no exact locality, 1 ; Chelmsford, Essex, 1 (U.S.N.M.) ; Tring, Hertford-
shire, 1 ; Boxniore, Hertfordshire, 1 ; Hoddesdon, Hertfordshire, 1 ;
London, 4 ; Gtodalming, Surrey, 1 ; Surrey, no exact locality, 1 ;
Bonchurch, Isle of Wight, 1; near Honiton, Devonshire, 2 ; Devonshire,
no exact locality, 8 ; no exact locality, 1.

Holland : Oosterlieek, Guelderlaud, 4.

France : Near Barcelonnette, Basses- Alpes, 1 (Mottaz).

Germany : Wernigerode, 3 ; Ummerstadt, Thiiringen, 2 ; Magdeburg, 2 ;
Niesky, Silesia, 4 ; Damsdorf , Silesia, 1.

Austria-Hungary : Csallokoz-Somorja, Pressburg, 7.

Switzerland : Geneva, 2 (Mottaz) ; Lausanne, Vaud, 1 (Mottaz) ;
Morat, Fribourg, 1 (Mottaz) ; Boudrv. Neuchatel, 2 (Mottaz) ; Teufen,
Appenzell, 1 (U.S.N.M.); Thurgau, St. Gallen, 1; St. Gothard, Uri, 3
(U.S.N.M.) ; no exact locality, 1.

Italy: i3orzoli, near Genoa, 1 (Genoa); Florence, 1 (Mottaz) ; Kimini,!;
Rome, 1 ; no exact locality, 1 ; Sicily, 1.

Spain : Silos, Burgos, 2 (U.S.N.M.) ; La Granja, Segovia, 1.

IteniarJcs. — Plecotus auritus is so readily distinguished from all
other European bats by the great size of the ears that no sj^ecial
comparisons are required.

6, 3 ?. Montrose, Scotland. J. H. Coward (c & p). 11. 1. 3. 48-51
6 St. Antrim, Ireland. Hon. N. C. Rothschild 1. 9. 3. 5.

PLECOTUS 261

2 6 al. Longford. Dr. G. E. Dobson (c & p). 76. 2. 12. 2.

6. Pembroke, Wales. W. E. dc Winton (c & p). 11. 1. 3. 393.

1 St. Winfarthing, Norfolk, E. Markham (c & p). 9G. 7. 27. 3.
England.

5. Diss, Norfolk. (IF. R. C. H. B. Grant (p). 11. 1. 3. 394.

Slwrrin.)
1 al. Woburn Sands, Bed- A. Death (c & p). 7. 9. 2. 1.

fordshire.
9. Boxmore, Hertford- H. Piffard (c & p). 11. 1. 3. 55.

shire.
1 St. Hoddesdon, Hertford- A. Chittenden (c & p). 80. 9. 27. 1.

shire.

6. Tring, Hertfordshire. Dr. E. Hartert (c & p). 11. 1. 3. 50.
9 al. London. No history.

i. London. Sergt. Brown (c & p). 11. 1. 3. 57.

6. S.Kensington, Middle- 11.1.3.54.

sex.
i. Godalraing, Surrey. W. T. Blanford (c & p). 11. 1. 3. 58.

1. Bonchurch, Isle of Rev. C. A.Biuy (c & p). 41.3.29.0.

Wight.
S,9. Honiton, Devonshire, G. C. Shortridge (c & p). 11.1.3.52-53.
300 ft.
4 (5, 4 9 al. Devonshire. No history.

2 .5, 2 9. Oosterbeek, Guelder- O. Thomas (c & p). 98. 2. 1. 1-4.

land, Holland.
9. Wernigerode, Harz, W. Wolterstorff (c & p). 0.2.8.1.

Germany.
6, 9. Wernigerode, Harz. Lord Lilford (p). 11. 1. 1. 45-46.

{Wolterstorff.)

5. Magdeburg, Saxony. Lord Lilford (p). 11. 1. 1. 47.

(Wolterstorff.)
<J, 9. Ummerstadt, Thiirin- Lord Lilford (p). 11.1.1.43-44.

gen. (Schuchardt.)
i. Niesky, Silesia. (IF. Lord Lilford (p). 99. 1. 9. 1.

Baer.)
(< al. Damsdorf, Silesia. Dr. A. Giinther (p). GO. 8. 1. 10.

7. C sail 6 k 6 z-Somorja, Budapest ^Museum (e). 94.3.1.5-11.

Pressburg, Austria-
Hungary.

6. Thurgau, St. Gallen, 0. Thomas (p). 4. 4. 5. 14.

Switzerland. {Zolli-
kofer.)
1. Switzerland. Leon O. Galliard (p). 75. 9. 20. 2.

9 al. Rimini, Italy. Florence IMuseum (e). 85. 7. 6. 2.

6. Rome. (C. Coli.) G. Barrett-Hamilton (p). 11. 1. 2. 15.

Skeleton Italy. {Prince Bona- Tomes Collection. 7. 1. 1. 728.

(without parte.)
skull).

9 al. Sicily. Purchased (Parzudaki). 52. 2. 26. 18.

1 al. La Granja, Segovia, jM. de la Escalera (c). 8. 7. 30. 5.
Spain.
1. Europe. Leyden ]\Iuseum (e). 37. 4. 28. 25.

262

CHIROPTERA

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BARBASTELLA

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Genus BARBASTELLA Gray.

1821. Barbastella Gray, London Med. Repos., xv, p. .300, April 1, 1821.

1825. Barbastdlns Gray, Zool. Journ., ii, p. 243, July, 1825.

1839. Synotus Keyserling and Blasius, Wiegmann's Archiv fiir Naturgescb.,

1889, I, p. -305.
1857. Synofiis Blasius, Saugethiere Deutschlands, p. 42.
1878. Synotus Dobson, Catal. Cbiropt. Brit. Mus., p. 175.
1897. Barbastella Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 375,

October, 1897.
1907. Barbastella Miller, Families and Genera of Bats, p. 223, June 29,

1907.

Type species. — VespertiUo harhasteUiis Bchreber.

Geographical distribution. — ^Northern Africa ; central and
southern Europe ; west-central Asia to the Himalayas.

Characters. — Dental formula : i '^J, c ^~, pm '—, m || = 34.
Skull with rather long, rounded brain-case and weak rostrum, the
upper surface of which is occupied by a shallow, flattened-concave
area extending from nares to faintly developed supraorbital
ridges ; auditory bullaj not specially enlarged. Ears broad and
short (laid forward they reach slightly beyond tip of muzzle),
joined across forehead ; nostrils opening upward and outward
on a flat median space between two high lateral swellings and
behind a prominent median pad.

Bemarhs. — The short broad ears joined together across fore-
head distinguish this genus from the other European membei-s of
the family Vespertilionidse. Two species are known, one peculiar
to the Himalayan region, the other occurring in Europe.

BARBASTELLA BARBASTELLUS Schreber.

1774. VespertiUo barbastellus Schreber, Siiugtbiere, I, pi. lv (description, i,

p. 168, under name: Das Kurzmaul). Based primarily on "La

Barbastelle " of Daubenton, Hist. Acad. Royale des Sci., Paris,

1759, p. 377. 1765.
1776. VespertiUo barbastelle P. L. S. Miiller, Natursyst. Suppl. u. Regist.-

Band, p. 17 (Burgundy).
18.36. Barbastellus daubentonii Bell, Hist. Brit. Quadr., pt. i, p. 63

(Burgundy ; based primarily on Daubenton).
1838. Barbastellus communis Gray, j\Iag. Zool. and Bot., ii, p. 495,

February, 1838 (Renaming of barbastellus).
1857. Synotus barbastellus Blasius, Saugethiere Deutschlands, p. 43.
1878. Synotus barbastellus Dobson, Catal. Cbiropt. Brit. Mus., p. 176.
1897. Barbastella barbastellus Miller, Ann. and IMag. Nat. Hist., Gth sor.,

XX, p. 385, October, 1897.
1910. Barbastella barbastellus Trouessart, Faune Mamm. d'Europe, p. 11.

Type locality. — Burgundy, France.

Geographical distribution. — Central and soutliern Europe,

264

CHIROPTERA

west to England, north to southern Norway and Sweden, east
into Asia.

Diagnosis. — Characters as in the genus ; foi'earm about 38
to 40 mm.

External cliaraders. — General form slender and delicate, the
legs long, the tail about equal to head and body. Muzzle short
and broad, its width considerably greater than distance from tip
of snout to joining membrane of ears, each side with a very large
glandular mass rising above level of flat median dorsal surface
and extending downward to involve most of iipper lip behind
nosti'il pad ; glandular masses densely hairy in noticeable contrast
with the finely pubescent nostril pad and essentially naked
median region between nostrils and inner bases of ears ; nostrils
crescentic in outline, opening upward, somewhat crowded between
glandular lateral masses ; nostril pad well defined, rounded off"
above, continued downward into median portion of upper lip, the
edge of which, between glandular masses and separated from
them by evident grooves, is distinctly convex, fitting like a
valve, when mouth is closed, over bare median callosity of lower
lip. Ear large and broad, though not specially elongated, the
tijj extending about 5 mm. beyond nostril when laid forward ;
antei'ior basal lobe very small, appearing like a rudimentary
Molossine keel, the portion of anterior border usually reflexed in
Vespertilionine bats thrown forward so that its base joins its fellow
of opposite ear, the region of juncture low but sufficiently well
developed to form a distinct pocket on side next forehead ; out-
line of anterior border strongly convex excejit for an evident
flattening just below the rather abruptly rounded off" and some-
what recui'ved tip ; posterior border abruptly concave above, then
nearly straight except foi' an abruptly projecting lobe near
middle (lobe usually about 1 ' 5 mm. long "by 1 mm. wide, its uppei-
border convex, its lower border concave) ; lower border of ear full
and rounded, but with no dift"erentiated antiti'agus ; tragus large,
somewhat ti-iangular in outline, its greatest width (slightly above
level of anterior base) equal to about two-thirds length of anterior
border, its up^ier jjortion I'apidly narrowing to an attenuate tip ;
meatus with well developed keel-like ridge ; outer surface of ear
densely furred except at extreme tip and along posterior border
to a little below level of projecting lobe ; inner surface with a
hairy line marking juncture of anterior border of conch with
portion usually folded backward, elsewhere irregularly sprinkled
with fine hairs. Membranes thin and delicate, the wing broad,
inserted at base of outer toe, the antebrachial membrane con-
tinued outward as a very narrow fold to base of thumb ; third
metacarpal slightly shorter than forearm and a little exceeding
the sub-equal fourth and fifth ; fur soft and loose, the hairs on
middle of back about 10 mm. in length ; on both ui:)per and
under surface of wing the fur extends to a line joining middle of
forearm with knee ; foot less than half as long as tibia ; calcar

BAKBASTELLA

265

about as long as tibia, with small but evident terminal lobe and
posterior keel ; tail with extreme tip projecting beyond inter-
i'emoral membrane.

Coluur. — General coloui' a very dai'k brown between the seal-
brown and vandyke-brown of Ridgway, the hairs everywhere
dark brown to extreme base, those of upper parts tipped with
light glossy wood-brown producing a sharply contrasted wash
throughout I'egion behind shoulders, those of undei'parts tipped
with a paler, more drabby brown, but without producing so
decided a contrast as on back, though the wash is usually evident
along middle of belly ; hairs on l)ase of under side of interfemoral
membrane pale ecru-drab or whitish smoke-grey nearly or quite
to base. Ears and membranes in dry specimens br-own, hardly
so dark as ground colour of body.

Skull. — The skull, though lightly-built and small, scarcely
exceeding that of Myotis mystacinus in length, is rather deep and
robust, with unusually lai'ge bi'ain-case relatively to the short
rostrum. Dorsal profile abruptly convex over anterior half of
brain-case, then nearly horizontal to somewhat overhanging
occiput ; ventral profile nearly horizontal, the floor of brain-case
scarcely elevated above level of tooth-i'ow. Brain-case high and
inflated anteriorly, relatively low posteriorly, its lateral outlines
when viewed from above essentially as in the
species of Myotis, but less rounded and globular
posteriorly ; floor of brain-case with no special
peculiarities ; auditory bulke scarcely more in-
flated than in the small species of Myotis.
Interorbital region slightly constricted, its sur-
face flattened and not well defined from that of
rosti'um ; lachrymal ridges rather prominent,
especially at their lower extremities. Zygoma
nearly straight, scarcely bowed outward beyond
level of outer surface of brain-case. Ante-
orbital foramen relatively large, over anterior narbasteiia harha-
margin of m^. Rostrum short, its upper sur- stdlus. Nat. size,
face flattened-concave, deeply emarginate in
front by the unusual backward extension of the uppei- margin of
anterior nares, the posterior border of which is at level of ante-
orbital foramina, exposing anterior extremity of vomer. Palate
.short, noticeably concave both longitudinally and laterally ; its
anterior emargination small, extending slightly behind level of
middle of canine ; its posterior border slightly behind level of vi^,
double emarginate with short median projection ; mesopterygoid
space slightly wider than long. Mandible slender, with no special
peculiarities.

Teeth. — In general aspect the teeth resemble those of Plecotus
(iHvitus, though they are throughout smaller and more slender.
1 ncisois botii above and below essentially as in Plecotus auritus
except that crown of outer upper incisor is nearly terete, and

266

CHIEOPTERA

space between it and eingulum of canine is veiy narrow ; lower
incisors forming a less broadly (J -shaped row. Canines as in
P. auritus. Anterior upper premolar very minute, crowded
inward completely from tooth-row, the diameter of its crown
barely one-quarter that of outer incisor, its cusp and ciugulum
distinct ; large upper premolar much as in P. auritus but less
narrowed on inner side and with a narrow concave crushing
surface ; no antero-internal basal cusp. Lower premolars almost
exactly similar to middle and posterior lower premolar of Plecoius
auritus, but crown area of larger tooth slightly exceeding that of
canine. Molars not appreciably different from those of Plecotus
auritus except that they are smaller and the concave median
region of the upper teeth is larger relatively to area of crowns.

Measurements. — Adult male from Cheshire, England : head
and body, 48; tail, 49; tibia, 19; foot, 7- 2; forearm, 38-4;
thumb, 6 ; third finger, 69 ; fifth finger, 52 ; ear from meatus,
15 ; width of ear, 14. Adult female from Epping, Essex,
England: head and body, 49*6 ; tail, 46 ; tibia, 18*2; foot, 7 ;
forearm, 36; thumb, 5-8; third finger, 69 ; fifth finger, 51 ; ear
from meatus, 16 ; width of ear, 14. Average and extremes of
five adults from Damsdorf , Silesia, Germany : head and body,
50(47-52); tail, 47-1 (45-49); tibia, 18-5 (18-19); foot, 6-6
(6-7); forearm, 38 (37-4-39); thumb, 5-8 (5 -4-6 -2); third
finger, 70 (68-72) ; fifth finger, 52-6 (51-54) ; ear from meatus,
15-5 (15-16); width of ear, 14-1 (13-6-15). For cranial
measurements see Table opposite.

Specimens examined. — Sixty-one, from fche following localities : —

England: Elton, Huntingdonshire, 1; Ellingham, Norfolk, 1 ; Arrow
Lodge, Warwickshire, 1 ; Alcester, Warwickshire, 1 ; Kugby, Warwick-
shire, 1; Welford, Rugby, 1 ; Pilton, Oundle, Northamptonshire, 1; Epping,
Essex, 1 ; Llanelwedd, Radnorshire, 1 ; Swindon, Wiltshire, 1 ; Milton,
Hampshire, 1.

France: Caterille, Haute-Cxaronne, 1.

Germany: Biichenberg, Harz Mountains, 1; Niesky, Silesia, 4;
Bavaria, no exact locality, 3 (U.S.N.IM.) ; near Wernigerode, Saxony, 1.

Switzerland : Grotte de Vallorbe, Vaud, 40 (Mottaz).

Remarks. — This species is immediately recognizable among
European bats by its short, joined ears, and liy the peculiar form
of the muzzle.

?. Llanelwedd, Radnorshire, H. E. Forrest (c ct p). 4. 7. 3. 1.

Wales.

9. Elton, Huntingdonshire, Lord Lilford (p). 94. 9. 5. 1.

England.

i. Ellingham, Norfolk. E. R. Alston (p). 81. 6. 9. 1.

(Dr. W. M. Crowfuot.)

6. Arrow Lodge, Warwickshire. Tomes Collection. 7.1.1.348.

1. Alcester, Warwickshire. Tomes Collection. 7. 1. 1. 348.

i St. Rugby, Warwickshire. G. Dalgleish (c & p). 4. 10. 13. 2.

i. Oundle, Northamptonshire. Lord Lilford (c & p). 94. 4. IG. 1.

? al. Epping, Essex. H. Doubleday (c & p). 44. 5. 1. 1.

i. Milton, Hampshire. Rev. J. E. Kelsall (p). 11. 1. 3. 59.

BARBASTELLA

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6. Swindon, Wiltshire. Rev. E. A. Goddard 8. 5. 12. 1.

(c & p).
6. Caterille, Haute-Garonne, 0. Thomas (p). G. 4. 1. 7.

1000 ra. France.

{A. Robert.)
6, 9. Niesky, Silesia, 181 m. Ger- Dr. E. Hamilton (p). 97. 12. 4. 2-3.

many. (W. Bacr.)
2 9. Niesky, Silesia. [W. Bacr.) Lord Lilford (p). 99.1.9.2,4.

i. We mi ger ode, Saxony. Lord Lilford (p). 0.2.8.2.

(WoUerstorff.)

Sub-Family MINIOPTERINtE.

1878. Minioptcri Dobson, Catal. Chiropt. Brit. Mus., p. 170 (part).
1907. Miniopterinx Miller, Families and Genera of Bats, p. 227, June 29,
1907.

Geograj^hiral distrihution. — Africa, southern Europe and
southern Asia, eastward to the Malay region, Japan and
Australia.

Characters. — Like the Vespertilioninae, but presternum with
median lobe enormously developed and forming the greater part
of the bone ; scapula with coracoid straight, directed con-
spicuously inward.

BemarJcs. — The sub-family MiniopterhiBP, though widely dis-
tributed in the warmer portions of the Old World, is at present
known to contain the genus Minioidcrus only.

Uenus MINIOPTERUS Bonaparte.

1837. Miniopterus Bonaparte, Iconogr. Faun. Ital., i, fasc. xx, under

Vcspcrtilio cmarginatns (Sub-genus of Ve^pertilio).
1857. Miniopterus Blasius, Siiugethiere Deutschlands, p. 45.
1860. Miniopteris Gray, Ann. and Mag. Nat. Hist., 3rd ser., xvii, p. 91,

February, 1866.
1878. Miniopterus Dohson, Catal. Chiropt. Brit. I\Ius., p. 347.
1892. Minyoptcrus Winge, Jordfundne og nulevende Flagermus (Chiroptera)

fra Lagoa Santa, Minas Geraes, Brasilien, p. 36.
1900. Minncoptcrus Lampe, Jahrb. Nassau Ver. Naturkundo, Jahrg. 53,

Catal. Saugeth.-Samml., p. 12.
1907. Miniopterus Miller, Families and Genera of Bats, p. 227, June 29,

1907.

Tijjje sperU's. — Vcspertilio urshii Bonaparte = V. srlireihersii
Kuhi

Geographical distribution. — Same as that of the sub-family.

Characters. — Dental formula : i i^'f, c ^J, pm ^~, m t;' = 34.
Skull with unusually high brain-case, and low, flattened rostrum.
Second phalanx of third finger nearly three times as long as fii'st.
Ears short, separate, the upjjer margin (in European species)
appearing almost artificially truncate.

Memarhs. — The peculiar shortening of the ears together with
the remarkable elongation of second phalan.x of thiid linger readily

MINIOPTERUS

l'G9

di,stini,'uisli tliis <((!nus among tlie inenil)ers of the Eui'opefiii fauna.
About a dozen forms have been described, one of which occurs
in soutliern Europe.

MINIOPTERUS SCHREIBERSII Kulil.

1819. Vcspcrtilio schrcibcisii Kuhl, Ann. Wetterau. Gesollsch. Naturk., iv
(= Neue Ann., i), pt. 2, p. 185 (Hungary).

1837. Vcspcrtilio ursinii Bonaparte, Iconogr. Faun. Ital., i, fasc. xxi
(Monto Corno, Ascoli, Italy). Typo in British Museum.

18-11. Vesjxriilio orsinii Temminck, Monogr. do Mamm., ii, p. 179 (modi-
fication of ursinii).

1857. Minioptcrus schrcibcrsii Blasius, Saugethiere Deutschlands, p. 4G.

1878. Miniopterus schrcibcrsii Dobson, Catal. Chiropt. Brit. Mus., p. 348.

1910. Minioptcrus schreibcrsi Trouessarfc, Fauiio Mamm. d'Europe, p. 34.

Type locnlifij. — Kuhnbazer Cave, mountains of soutliern
Bannat, Hungary.

(h'0(jraphic(d distn'hutlon. — Soutliern Europe from the Iberian
Peninsula eastward, north to Switzerland and Hungary. Limits
of range not known.

Dia/jnosis. — Charaters as in the genus ; forearm about
43 mm.

Ejctcrnal characters. — General form rather slender, with long
tail and legs, wing broad at base but conspicuously tapering at
tip, and short ears with a peculiar truncate aspect. Muzzle
rather broad, though without conspicuous glandular swellings,
its greatest width about ecjual to distance from eve to nostril ;
muzzle pad narrow, with slight median emargination, bounded
below by a low horizontal ridge which is continuous with projecting
inner margin of nostril. Eyelids noticeably glandular-swollen ;
a deep horizontal groove in cheek below eye. Ear short, extend-
ing about half way from eye to nostril when laid forward, its
general aspect difierent from that of any other European bat,
owing to the length of the anterior basal lobe, the short, straight
anterior border, and the broadly, evenly convex posterior border
which joins anterior border in such a manner that there is
l)ractically no " tip," the whole anterior border appearing like an
oblicjuely, almost artificially truncate extremity ; antitragus low,
obscurely marked ofli' from posterior border of conch, practically
continuous with lower lip anteriorly ; inner surface of conch
slightly rugose, without evident cross ridges ; tragus about half
as high as conch, a little curved forward owing to .slight con-
cavity of anterior border, the blunt tip and upper half of e.xterior
border forming a uniform, rather noticeable convexity, the
posterior margin straight below to rudimentary basal lobe ;
greatest width of tragus about half anterior border. Wing rather
wide basally, the tifth finger exceeding forearm by about one-tifth
length of latter, the tip unusually slender and elongate owing to
the great length of last bone of third tinger ; third and fourth

"270 CHIROPTERA

metacarpals sub-e(|iial, about 3 mm. shorter tlian forearm ; tifth
metacarpal about 4 mm. shorter than third ; membrane joining
leg at or a little above ankle. Leg rather slender, the foot
scarcely half as long as tibia ; calcar about as long as tibia and
nearly equal to free border of uropatagium, its distal termination
very obscure, its posterior border without trace of keel. Tail
about as long as head and bod\', included in the membrane to
extreme tip.

Fur and colour. — The fur is of a Aery soft, silky texture,
though rather short, the hairs at middle of back only about
7 mm. in length, those of head abruptly nuich shorter (about
4 mm.) in rather noticeable contrast. Above it scarcely extends
on membranes except for a sparse pubescence on basal half of
median portion of uropatagium ; below it reaches line joining-
middle of forearm with knee and continues across extreme base
of uropatagium. General colour of upper parts drab, faintly
lighter and more nearly hair-brown anteriorly ; underparts ecru-
drab. The hairs of back show three evident though not strongly
contrasted colour-bands : (o) at base (3 nnii.), a slaty mouse-grey ;
(h) at middle (tips of shorter hairs, 2 mm.), ecru-drab ; and (c-) at
tip (extremities of longer hairs, 2 mm.), drab. Muzzle, ears and
membranes brown, scarcely darker than body.

Skull. — The skull differs from that of all other European bats
in the great inflation of the anterior portion of brain-case and
the consequent very abrupt angle at which the forehead rises
above the low, flat rostrum. In other respects the general aspect
of skull, especially when viewed from above, is rather slender
and lightly built. Dorsal profile rising gradually from front
of nares to inter-lachrymal region, then
abruptly at an angle of nearly 45 to middle
of anterior portion of brain-case which is
nearly or quite as high as lambda, the region
£<^^ between these two highest points occupied

by a .shallow concavity corresponding to
anterior edge of interparietal ; below and
behind lambda the supraoccipital bulges
noticeably outward, but not sufficiently to
conceal condyles when skull is viewed from
directly above ; ventral profile nearly flat,
*'"'• *^- slightly elevated posteriorly. Brain-case

iliidopterus schreibersU. in i. ii i.T iT i t i

Nat. size. broadly ovate, the outline broken at each

side posteriorly by the slightly projecting
mastoid region ; sagittal crest low Ijut evident anteriorly,
obsolete posteriorly ; lambdoid crest moderately developed at
sides, barely indicated at middle ; general outline of skull when
viewed from behind squarish, the depth through bulla about
equal to breadth of bi'ain-case ; floor of brain-case with no special
peculiarities ; between cochlea and median line a shallow groove
with abrupt pit-like anterior termination a noticeable ridge

MINIOPTERDS

271

at outer side of each pit ; auditory bulla moderately large, the
transverse diameter about equal to distance between bullte.
Interorbital region moderately constricted, slightly hour-glass
shaped, the lachrymal region scarcely more than half as wide
as brain-case, smoothly rounded at sides ; rostrum tapering
gradually, rounded oft" at sides, ftattened concave along median
line, the narial emargination small, squarish, extending backward
about one-third of the distance to front of forehead ; rostral
depth at fi'ont of orbit about equal to distance from orbit to
outer incisor ; anteorbital foramen rather lai'ge, directly above
small pi-emolar, the anteorbital canal half as long as rostrum,
thus much longer than in any other European bat ; lachrymal
foramen just outside of orbit, on level with uppei- border of
anteorbital foramen. Palate long and wide, distinctly concave
both laterally and longitudinally, terminating rather abruptly a
little behind level of last molar, a small but evident foramen at
each side near posterior edge ; median sjiine large ; mesoptery-
goid space squarish, slightly wider posteriorly than anteriorly,
the short hamulars bent inward. Mandible slender, a little
deeper at symphysis than behind tooth-i'ow, the posterioi' seg-
ment vmusually small, with nearly horizontal, slightly concave
upper border ; angular process relatively long, expanded at
outer end.

Teeth. — Relatively to size of skull the teeth are small and weak.
Inner upper incisor low, the crown very oblique, with postero-
external concavity and small posterointernal cusp ; outer upper
incisor considerably larger than inner, the crown flattened in
axis of tooth-row, the width of its flattened-conca^•e posterior
surface about half its height, that of its outer border scarcely
one-fifth height, cingulum obsolete, but forming a minute postero-
external cusp ; the two teeth lie in curve of anterior portion of
tooth-row, and the outer is separated from canine by a space
about equal to its greatest diameter. Lower incisors closely
crowded but not imbricated, the crown-area increasing regularly
from first to third, the outline of the row as a whole broadly
V-shaped ; i^ and i., with crown very low, the cutting edge
t>bscurely tritid, the crown of each tooth wider posteriorly (in line
of tooth-row) than anteriorly, but this more evident in second
tliau in first ; /.j nearly terete, the middle and posterior cusps
eidarged and separated by a deep groove, the anterior cusp
reduced to a mere rudiment. Upper canine slender and weak,
sub-terete, but distinctly flattened on inner side, the shaft with
distinct anterior and posterior longitudinal grooves but without
well developed cutting edge, the cingulum narrow, complete,
but without cusps ; lower canine scarcely higher than main
cusps of molars, its inner and posterior surfaces flattened, its
antei'o-external surface smoothly rounded, cingulum forming a
low but evident antero-internal cusp. Anterior upper premolar
with crown area about equal to that of canine, its general form

272 CUIKOPTERA

essentially as in large premolar except that the antero-internal
basal cusp is barely indicated, the cutting edges of the main cusp
are less developed, and the crown is narrower extei-nally than
internally ; large upper premolar with crown area about equal
to that of first molar and double that of small premolar, its
anterioi' and postei'ior borders very slightly concave, the antei'O-
internal basal cusp small but evident, the main cusp with well
developed anterior and posterior cutting edges, and antero-internal
and postero-external longitudinal groove ; lower premolars
essentially alike in form, the crown area of the posterior tooth
about equal to that of canine, that of the two others successively a
little less, the outline S(iuarish, with antero-external angle tending
to become I'ounded off", jiarticulai'ly in the posteiior tooth ; cusp of
first about half as high as canine, that of second slightly higher
than tii'st, that of third nearly as high as main cusps of molars ;
cingulum well developed, forming in each tooth a small but
evident antero-internal basal cusp. First and second upper
molars with posterior border- deeply and almost angularly
emarginate near iniier edge of tooth, the anterior border nearly
straight ; protocone low but rather broad ; hypocone clearly
indicated but not distinct from posterior commissure of jirotocone ;
pai'acone and metacone nearer outer edge of crown tlian usual,
leaving a wide, pit-like median concavity, metacone slightly the
higher of the two cusps ; styles and commissures well developed,
Wpattern normal ; third upper molar with crown area about
half that of first, its width through metacone slightly less than
half length of anterioi- border, its three cusps and three
commissures normally developed. Lower molars with no special
peculiarities, the second triangle oi vi.^ narrower than first, but
of about the same area.

Measurements. — Adult male and female from Neuchatel,
Switzerland : head and body, 59 and 59 ; tail, 58 and 60 ;
til)ia, 20 and 19 ; foot, 10 and 9-4 ; forearm, 45 and 44-6 ;
thumb, 7 and 7 ; third finger, 92 and 92 ; fifth finger, 53 and 54 ;
ear from meatus, 12-6 and 12 ; width of ear, 12 and 12. Adult
male and female from Corinth, Greece : head and body, 57 and
56; tail, 57 and 55; tibia, 18-6 and 19; foot, 10 and 9-6;
forearm, 44 and 45 ; thumb, 6 • 6 and 7 • 2 ; third finger, 92 and
90 ; fifth finger, 54 and 53 ; ear from meatus, 12 and 12 ; width
of ear, 12 and 12. Two adult males from Silos, Burgos, Spain :
head and body, 60 and 60 ; tail, 60 and 57 ; tibia, 19 and 19-4 ;
foot, 11 and 9*6 ; forearm, 44 and 44 ; thumb, 7-4 and 7 4;
third finger, 93 and 89 ; fifth finger, 55 and 56 ; ear from meatus,
11-4 and 11-6; width of ear^ 13-4 and 12-6. Two adult
females from the same locality : head and bod}^ 59 and 59 ; tail,
60 and 59 ; tibia, 19-6 and 20 ; foot, 9-8 and 10 ; forearm,
45-4 and 46 ; thumb, 8 and 7-4 ; third finger, 93 and 9:! ; fifth
finger, 55 and 55 ; ear from meatus, 12 and 11*6 ; width of ear,
12 '6 and 12-6. For cranial measurements see Table, p. 274.

MINIOPTERUS

273

Siiecimcns examined. — Three huudred and fifty-one, from the following
localities : —

Spain : Silos, Burgos, 9 (B.M. and U.S.N.M.) ; Seville, 1 ; Minorca,
Balearic Islands, 3 ; llajorca, Balearic Islands, 2.

France: Troubate, Hautes-Pjreuces, 2; Dions, Gard, 9; Marseilles, 1
(U.S.N.M.).

Switzerland: Geneva, 235 (Mottaz) ; Neuchatel, 2 (U.S.N.M.).

Austria-Hungary : Ofener Mountains, Hungary, 1 ; Hungary, no exact
locality, 3 (U.S.N.M.).

Montenegro : Beri, 1 ; Velgi, Czolo, 1.

Italy : Western Liguria, 14 (Genoa) ; Finalborgo, Liguria, 5 (Genoa) ;
Maremma, Tuscany, 1 (U.S.N.M.); Pisa, 1; Florence, 1 (U.S.N.M.);
Livorno, 6 (U.S.N.^NI.) ; Spezia, 5 (U.S.N.M.); Monte Corno, Ascoli, 1
(tyipe oi ur. si nii Bonaparte); Rome, 10; Velletri, Rome, 2 (U.S.N.M.) ; no
exact locality, 1 ; Elba Island, 5 (U.S.N. Til.) ; Marsala, Sicily, 2.

Sardinia : Sassari, 3 (U.S.N.IM.) ; Cagliari, 4 (U.S.N.M.) ; Mount
Gennargentu, 2 (U.S.N.M.).

Greece : Corinth, 17 (U.S.N.M.) ; Labyrinth, Crete, 1.

MemnrTcs. — The lengthened second phalanx of the third finger,
the peculiar cropped appearance of the ears, and the short, dense,
velvety fur of the head are the most obvious external character-
istics of Minio])terus schreibersii as compared with other European
bats.

2.

S,9.

3S.

<?, 9.

8 <5, 1 9
1 al.

9.

9.
7 <J, 3 9.

Silos, Burgos, Spain. Rev. S. Gonzalez (c).
Inca, Majorca, Balearic O. Thomas and R. I.

Islands. Pocock (c & p).

San Cristobal, Minorca. 0. Thomas and R. I.

Pocock (c & p).
Troubate, Hautes- 0. Thomas (p).

Pyrenees, 400 m.

Prance. (A. Robert.)
Dions, Gard, 70 m. 0. Thomas (p).

France. (C. Mottaz.)
Ofener Mountains, Budapest Museum (e).

Budapest.

Beri, Montenegro.
(L. Fllhrer.)
Velgi, Czolo.

(L. Fiihrer.)
Rome. (C. Coli.)
Skeleton Monte Corno, Ascoli,
(without Italy,
skull).
2 6. Marsala, Sicily.

[A. Hubert.)
6. Labyrinth, 200 ft.

Crete.

0. Thomas (p).

0. Thomas (p).

Lord Lilford (p).
Tomes Collection.

88. 1. 5. 46-47.
0. 7. 1. 4-5.

0. 7. 1. 32-34.

6. 4. 1. 8-9.

8. 8. 10. 1-9.
94. 7. 18. 10.
5. 8. 4. G.
5. 8. 4. 5.

11. 1. 2. 5-14.

7. 1. 1. 735.

(Type of Yespertilio ursinii Bonaparte.)
0. Thomas (p). 6. 8. 4. 24-25.

Miss D. Bate (c). 5. 12. 2. 9.

274

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Family MOLOSSID^.

1865. Mulossi Peters, Monatsber. k. preuss. Akad. Wissensch., Berlin, p. 258.
1872. Molossidx Gill, Arrangement of the Families of Mammals, p. 17.
1878. Emhallonuridx (part ; Molossime, part, Molossi) Dobson, Catal.

Chiropt. Brit. Mus., p. 402.
1907. Molossidx Miller, Families and Genera of Bats, p. 241, June 29, 1907.

Geograjjlncal clistrihufion.- — Warmer parts of both hemispheres ;
in the Old World north to the Mediterranean region and
southern Asia.

Characters. — Essentially like the VespertHiotiidse except that
the secondary articulation of humerus with scapula is more
perfectly developed and the fibula is robust, adding appreciably
to strength of leg. Auditory bulla noticeably emarginate on inner
side. Tail projecting very consj^icuously beyond membrane, a
character by which the only European member of the group may
be immediately I'ecognized.

BemarJcH. — The family Molossidse is widely distributed in the
warmer parts of both hemispheres. Eleven genera are now
known, one of which is represented in the Mediterranean region
of Eurojae.

Genus NYCTINOMUS Geoftroy.

1818. Nyctinomus Geoffrey, Descr. de I'Egypte, ii, p. 114 ({egyptiacus).
1821. Nyctinoma Bowdich, Anal. Nat. Class. Mamm., p. 28 (Modification
of Nyciincmiiis).

1821. Nyctinomes Gray, London Med. Eepos., xv, p. 299, April 1, 1821

(Modification of Nyctinomus).

1822. Nyctinomia Fleming, Philos. of ZooL, ii, p. 178 (Modification of

Nyctinomus).
1842. Mops Lesson, Nouv. Tabl. Regno Auim., p. IS (Mops indicus Lesson

= Dysopcs mops F. Cuvier).
1878. Nyctinomus Dobson, Catal. Chiropt. Brit. Mus., p. 420 (part).
1902. Nyctinomops Miller, Proc. Acad. Nat. Sci. Philadelphia, p. 393,

September 12, 1902 [fcmorosaccxis).
1907. Nyctinomus Miller, Families and Genera of Bats, p. 251, June 29, 1907.

Type species. — Nyctinomus segypiiacus Geoffroy.

Geographical distribution. — Warmer portions of both hemi-
spheres, north to the southern United States and to the Mediter-
ranean coast of Euroj^e, east to the Pliilipjsines and Norfolk
Island.

Characters. — Dental formula: i 'J; or y.,* c '^, jmi j;-^, m :: ;' = 30
or 32 ; brain-case not unusually flattened, its dej^th at least half
its width ; bony palate with a small median anterior emargination
extending to behind level of roots of incisors.

Reynarhs. — As thus defined the genus Nyctinomus contains

* In the species occurring in Europe.

NYCTINOMUS

277

fibout forty species, two-thirds of wliich are peculiar to the Old
World, one of them occurring in the Mediterranean region of
Europe.

NYCTINOMUS TENIOTIS llalinesque.

1814. Ccphalotes teniotis Bafinesquo, Pr6c. des D6couv. Somiol., p. 12 (Sicily).
1825. Dino2)s ccstoni Savi, N. Giom. de' Letterati, Pisa, x, p. 235 (Pisa, Italy).
1810. Dysopcs savii Schinz, Europ. Fauna, I, p. 5 (Substitute for ccstoni).
1871. [Dysopcs cestonii] var. nifjrogrisciis Schneider, Neue Denkschr.

Schweiz. Gesellsch. Naturwiss., xxiv, p. 5 (articles separately

paged). Basel, Switzerland.
1877. Nyctinomus cesionii Dobson, Catal. Chiropt. Brit. Mus., p. 423.
1891. Nyctinomus ttenioiis Thomas, Proc. Zool. Soc., London, p. 182.
1897. [Dysopcs} viidas Schulze, Abh. Ges. Nat. iv, No. 10, p. 23 (Substitute

for cestoni). Not of Sundevall, 1842.
1910. Nyctmomus txniotis Trouessart, Paunc Mamm. d'Europe, p. 36.

Type locality. — Sicily.

Gcog7-aphical flistrihuiion. — Mediterranean region of Europe
and noi'thern Africa. Accidental ? at Basel, Switzerland.

Diagnosis. — Like the African Nyctinomus segyptiacus but
lai'ger, condylobasal length of skull about 23 mm. instead of
about 20 mm. ; lower incisor 3-3 instead of 2-2 ; small upper
premolar with crown area more than one-half instead of less
than one-fifth that of upper incisor. Distinguishable among
European bats by the generic characters ; forearm about 60 mm.

External characters. — Form heavy and robust, the legs short, the
feet large, the wings long and narrow, the membranes thick and
leathery, the ears very large, sub-orbicular, joined at their anterior
bases ; these characters in connection with the thick muscular tail,
projecting by at least one- third of its length be3^ond interfemoral
membrane, immediately distinguish the animal from all other
European bats. Muzzle projecting rather noticeably beyond
upper lip, oblicpiely truncate ; nostril pad well defined, wider
than high, the sub-circular nostrils opening forward and slightly
outward at its outer margin, its surface with very fine reticula-
tions and a few coarse wrinkles, glabrous except below, where it
is sprinkled with fine hairs, this hairy area continuous with the
l)rush-like fringe of i-ecurved blunt hairs along middle of upper
lip ; upper margin of pad broadly concave at middle, convex
laterally over the nostrils, its edge thickly set with small, laterally
compressed horny excrescences, about thirty-five in number,
the outermost lying a little above level of middle of nostril ; a
row of about ten similar excrescences crosses middle of pad
vertically. Upper lip very large and full, marked by manj^ deep
ol)li([ue wrinkles, its surface both abo^■e and below rather densely
hairy. A small wart on chin just behind level of symphysis.
Ear very large, sub-orbicular, the margin with no very decided
irregulaiities, though the anterior border usually shows some
slight flattening and the posterior border is faintly concave above

278

CHIROPTERA

and neai' base ; anterior basal margin of ear with a narrow
outward-folded hem, the exposed side of which is densely
pubescent except at extreme anterior margin, where it is glabrous
and set with about six terete wart-like projections, widely and
irregularly spaced ; jiosterior basal margin with a shorter, deeper,
inwardly-folded lobe just above concavity limiting posterior
base of antitragus ; keel well developed, not thickened at edge,
its height posteriorly ecjual to about one-third its length, the
margin of keel hairy, the line thus begun curving upward and
backward under upper margin of conch ; antitragus well defined,
its height slightly greater than that of keel, its length about
twice height, its anterior border with ridge-like continuation
forward to corner of mouth ; tragus squarish in outline, its
posterior border longest and with distinct angle below middle, its
anterior border shortest, its upper border almost horizontally
truncate ; antei'ior and upper margin of tragus fringed with long
loose hairs. Wings longer and nai'rower than in any other
European bat, the membrane inserted on side of tibia just above
ankle ; antebrachial membrane extending as a narrow fold along
forearm to base of thumb ; the fur of body extends on both
surfaces of wing to line joining middle of humerus with knee,
and on to extreme base of uropatagium ; otherwise the membranes
are essentially naked except for some fine dense pubescence on
upper side of propatagium. Thumb short and robust, with
moderately developed pad at distal end of metacarpal. Foot
broad and robust, more than half as long as tibia, the sole with
a low and rounded but evident pad at middle, four small elongated
pads in a row at bases of toes, and some smaller, less definite
callosities in space between this row and the large median pad.
Outer and inner toe thickened, their outer surfaces densely
covered with short stifiened hairs with recurved points ; sprinkled
among these shorter hairs and also at the ends of the other toes
are a few much longer bristles. Calcar about as long as tibia,
its point ill-defined ; no indication of keel or of terminal lobe.
Tail about half as long as head and body, robust and muscular,
the terminal third or half pi'ojecting beyond membrane.

Fur and colour. — The fur is everywhere dense and velvety
in texture, the hairs at middle of back about 7 mm. in length,
those on throat longer and looser. Colour a uniform light drab,
with faint darker shading in certain lights, the hairs pale ecru-
drab at extreme base. Ears and membranes in dry sjaecimens
blackish. Fringes on feet and hairy lines on ears, drab like
body.

SJcull. — The skull is large, but rather slender, about equal to
that of Myotis myotis in length. In general form it is distinguish-
able among those of the bats of Europe by the depressed brain-
case, the high, somewhat tubular rostro-interorbital portion, and
the conspicuously emarginate inner side of auditory liulhe. Dorsal
profile essentially straight from nares to lambda, though with a

NYCTINOMDS

279

slight convexity over anterior two-thirds of brain-case, and a
more aV)rupt though not very conspicuous swelling posteriorly,
the two convexities separated by a narrow concavity ; general
direction of dorsal profile more nearly horizontal than in any
other European bat, since the depth of occiput through condyles
is scarcely greater than that of rostrum through anterior portion
of tirst molar. Brain-case low and wide, its depth at middle
about half mastoid breadth, its surface smooth and evenly
rounded, with faintly indicated sagittal crest posteriorly and
median groove between latei-al swellings anteriorly ; lambdal
crest evident though not high ; outline of brain-case when
viewed from above a somewhat triangular ovate owing to the
rather squarely truncate posterior border and the strong contrast
between the wide mastoid region and narrow interorbital con-
striction ; floor of brain-case with median ridge and lateral
depressions posteriorly, the basisphenoid
with two shallow but rather well-delined
pits about as large as glenoid surface ;
auditory bulhe rather large, deeply emar-
ginate on inner side so that cochlea is
conspicuously exposed, the region in
front of middle of meatus reduced to a
narrow ring barely more than one-third
as wide as meatus, the extreme anterior
border with a flange-like inward-curved
projection. Interorbital region long, sub-
cylindrical, a little wider at lachrymal
level than posteriorly, but scarcely enough
so to impart a distinctly hour-glass general
form ; least interorbital breadth slightly
greater than breadth of tubular narial
region and slightly less than that across
roots of canines. Zygoma simple, not
bent upward, its margin with a barely indicated expansion behind
middle. Rostrum proper short and deep, not w^ell difierentiated
from interorbital region, the distance from orbit to front of pre-
maxillary about equal to depth at front of in} ; lachrymal ridge
short but well defined, the small, inconspicuous anteorbital fora-
men and minute lacluymal foramen opening forward under its
anterior edge, at level of middle of large premolar ; nares with
distinctly tubular latei'al margins separated from roots of canines
by evident grooves, the dorsal emargination extending about half
way to lachrymal level. Palate moderately wide, noticealjly
concave laterally, the anterior emargination small, about as deep
as wide, its posterior border scarcely extending behind level
of front of canine ; posterior border of palate double emargi-
nate with well developed median projection, the emarginations
extending forward to level of posterior margin .of m^ ; mesop-
terygoid- space large, its width anteriorly equal to that of

yycHnomus teniotU.
Nat. size.

280

CHIROPTERA

temporal fossa at sauie level, its length about twice width ;
hamulars small and inconspicuous. Mandible long and straight,
its axis scarcely bent upward posteriorly ; symphysis deep, scarcely
or not subtended by a concavity in lower border of ramus ;
posterior portion of mandible low, the depth through coronoid
process barely equal to distance from front of coronoid to back
of condyle ; angular process large, straight, directed backward,
outward and downward.

Teeth. — Relatively to size of skidl the teeth are rather large,
the lower molars in particular. The most obvious peculiarities
of the dentition as compared with that of other European bats are
the single, well developed incisor in each side of upper jaw, and
the presence oi a large hypocone, quite distinct from commissure
of jarotocone, in i«' and wi". Upper incisor simple, a little moi'o

than half as high as canine, the
shaft set at an angle so that the
points of the two teeth are much
nearer together than their bases ;
cingulum faintly indicated in front,
better developed and forming an
incipient cusp postero-externally ;
posterior surface of shaft somewhat
flattened ; space between incisor
and canine aljout equal to greatest
diameter of smaller tooth. Lower
incisors three on each side, much
imbricated and closely crowded in
narrow space at front of canines,
their cutting edge barely rising to
level ■ of canine cingulum. The
inner and middle tooth sub-equal,
rather deeplj' bifid, the outer
barely equal to outer lobe of the
others, its apex faintly notched.
Upper canine moderately large, its crown area somewhat more
than double that of upper incisor, the general outline of its
base an irregular triangle with its longest side directed
inward, and with the postero-external side slightly concave ;
cingulum narrow but well developed throughout, not tending to
develop small cusps ; lower canine with no special peculiarities,
its cingulum tending to form a slight anterior cusp. Small
upper premolar in the tooth-row, usually in contact with canine
and separated from large premolar by a narrow space, its crown
area somewhat less than that of upper incisor, its cingulum and
cusp well developed, the cusp about one-fourth as high as shaft
of canine, triangular in outline when viewed from the outside,
with well developed posterior cutting ridge ; lower premolars
similar in form but larger, their crown areas sub-equal, the shaft of
the second higher than that of first and about equal to larger cusps

FIG. 49.
Xyctiiiomus Icnlotis. Teetli X 5.

NYCTINOMDS

281

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JOJ CHIROPTERA

of lower molars ; lai'ge upper j^remolar with crown area about
two-thirds that of m}, its antero-internal cu.sp high and distinct.
First and second upper molars alike in sti'ucture, the first slightly
larger than second, the cusps all well developed and distinct,
showing no special peculiarities aside from the presence of a
conspicuous terete hypocone nearly as large and distinct as antero-
internal cusp of large premolar, and separated from commissure
of protocone by a deep notch ; third upper molar with crown
area not much less than that of second, except for the absence of
postero-internal heel and hypocone ; third commissure as long as
in the other teeth, but metacone smaller than paracone. Lower
molars with no special peculiarities, the cusps all well developed
and distinct.

Meaattroncnts. — Young adult male from Italy : head and
body, 87; tail, 56; free portion of tail, 20; tibia, 19; foot,
10' 4; forearm, GO* 4; third linger, 115; lifth linger, 65; ear
from meatus, 30' 4 ; width of ear, 32. Young adult female from
Greece : head and body, 84 ; tail, 57 ; free portion of tail, 27 ;
tibia, 20; foot, 11 "4; forearm, 61 ; third finger, 117; fifth
finger, 61 ; ear from meatus, 31 • 4 ; width of ear, 31. Two adult
females from Cintra, Portugal : forearm, 60 and 61 ; third finger,
110 and 115 ; fifth finger, 59 and 63. For cranial measurements
see Table, p. 281.

Specimens examined. — Six, from the followiug localities : —
Portugal : Cintra, 2.

Italy : No exact locality (probably from Pisa), 2. (B.M. and U.S.N.M.) ;
Sicily, 1 (U.S.N.M.).

Greece : No exact locality, 1.

2 9. Cintra, Portugal. O. Thomas (c & p). 98. 2. 2. 6-7.

i. Italy (probably Pisa). Zoological Society (r). 10. 5. 24. 1.

(jSVui.)

?. Greece. Parreys. 46.0.15.121.

CAKNIVORA

283

Order CARNIVORA.

1827. Carnivora Gray, Griffith's Cuvior. Auim. KirigcL, v, p. 111.

Geoi/rnplrlcal dinfrihutinn. — Continents and larger islands of
the entire world, Australia,"' New Zealand, and the Antarctic
region excepted.

CJiaracters. — -Tei'restrial (rarely aquatic or semi-aquatic), non-
volant, placental mammals with rather high development of
brain, the cerebral hemispheres with distinct convolutions ; feet
unguiculate, never modified as fins or flippers ; dentition of a
modified tuberculosectorial tyjie, the posterior upper premolar
and antei'ior lower molar usually develojjed as special carnassial
or flesh-cutting teeth.

BcmarJcx. — The mammals of this order present much diversity
of form and structure, though less than in the case of the
Insectivora. Most of the living members of the group are
carnivorous in habits, and immediately recognizable among
placental mammals by the presence of a specially modified flesh-
tooth in each jaw. In cei'tain groups, however, as in the Ursidse
among the European representatives of the order, both habits
and dentition are of a more generalized type. The order contains
seven 7'ecent families, five of which occur in Europe.

KEY TO THE EUROPEAN FAMILIES AND SUB-FAMILIES
OP CARNIVORA.

Larger cheek-teeth with crowns of a crushing type, the
cusps sub-equal, low, subterete, without noticeable
cutting edges ; upper carnassial 2-rooted, in front of
anteorbital foramen, its inner lobe posterior ; size
very large; form heavy ; feet plantigrade (Bears) ... LVsida;, p. 284.
Larger cheek-teeth with crown at least partly trenchant,
the outer cusps of one or more in each jaw narrow
and with well developed cutting edge, the inner
cusps reduced or absent ; upper carnassial 3-rooted,
behind anteorbital foramen, its inner lobe median
or anterior.
Cheek-teeth without crushing surfaces ; upper molar
scarcely larger than outer incisor ; claws com-
pletely retractile (Cats) Fclidnj, p. i55.

Cheek-teeth, at least the hindermost, with evident

crushing surface ; upper molar (or first when more

than one are present) much larger than outer

incisor ; claws partly or not retractile.

Tooth-row relatively long (more than half condylo-

basal length of skull) ; number of teeth in

European members of family 42 (Dogs) Canidx, p. 303.

* Represented in Australia by a species of Canis, probably introduced.

284

CARNIVOEA

Tooth-row relatively short (less than hal£ condylo-

basal length of skull) ; number of teeth in

European members of family not more than 40.

Auditory l)ulla divided into two chambers, the

boundary between which is marked externally

by an oblique constriction ; upper molars

usually (always in European genera) 2-2,

the crown of the first wider externally than

internally (Genets and Mongoose) Vivcrridn', p. HO.

Auditory bulla simple ; upper molars 1-1, the

crown wider internally than externally Miistclida' , p. 340.

Upper carnassial with evident crushing surface

on inner side, the crown triangular or

rhombic in outline ; upper molar large, the

length of its outer portion usually equal to

or greater than that of carnassial.

Skull normal, the rostrum longer than broad ;

external form not modified for aquatic

life, the toes long-clawed, not webbed,

the tail not conspicuously muscular ; fur

loose and coarse (Badgers) Mdlnas, p. 341.

Skull much flattened ; rostrum broader than
long ; external form modified for aquatic
life, the toes short-clawed, webbed, the
tail conspicuously muscular ; fur dense

and fine (Otters) Lulrinx, p. 354.

Upper carnassial without crushing surface on

inner side other than a small concave area

between small inner lobe and main cusp,

the crown not triangular or rhombic in

outline ; upper molar much reduced, the

length of its outer portion one-third to

one-half that of carnassial.

Dentition highly trenchant; small premolars

not opposite, at least one x^air capable of

shearing action ; upper carnassial with

posterior cusp narrow and trenchant ;

auditory bulla longer than broad; form

slender; feet digitigrade (Martens and

Weasels) Mufteliiuc, p. 3G4.

Dentition not highly trenchant ; small pre-
molars opposite, not capaljle of shearing
action, the points of all but pni^ and pni^
widely separated when jaws are closed ;
upper carnassial with posterior cusp
broad, almost flat-topped; auditory bulla
broader than long ; external form heavy ;
feet sub-plantigrade (Glutton) Guloniitie, p. 432.

Family URSIDiE.

1825. Ursid;i- Gray, Thomson's Annals of Philosophy, xxvi, p. 339,
November, 1825.

(jeot/rapliicnl distribution. — Northern hemisphere, south in the
Old World to the Atlas Mountains and the Malay Archipelago,
and in America to the Andes.

Characters. — Larger cheek-teeth of a strictly crushing ty23e,
the crowns wide and flattened, with large terete cusps, the last

DRsus 285

upper premolar and first lower molar scarcely differentiated as
carnassials, the former 2-rooted, its inner lobe at posterior border
of crown, its position so far anterior to level of anteorbital
foramen as not to he at point of greatest mechanical efficiency ;
auditory bulla flattened, without septum ; form heavy ; size
large ; feet strictly plantigrade ; digits, 5-5.

Memarlcs. — The family Ursidse, containing the bears, is at
jiresent represented by five or six genera, though the fossil
remains of others are known. The members of the group are so
easily recognizable by the peculiarities of the cheek-teeth that
they require no special comparisons with other carnivora. Two
genera occur in Europe.

KEY TO THE EUROPEAN GENERA OP UBSIDM.

Cheek-teeth relatively large ; incisors and canines not

specially enlarged and prehensive (Ordinary Bears) Ursus, ]). 285.

Cheek-teeth relatively small ; incisors and canines
enlarged and unusually prehensive in character
(Polar Bears) Thalarctos, p. 297.

Genus URSUS Linnwus.

1758. Ursus Linnaeus, Syst. Nat., i, 10th ed., p. 47 (arctos, by tau'touymy).
1857. Ursus Blasius, Saugethiere Deutschlands, p. 196.
1864. Euarctos Gray, Proc. Zool. Soc. London, p. 692 [americanns).
1864. Myrmarctos Gray, Proc. Zool. Soc. Loudon, p. 694 (eversmanni = arctos).
1898. Ursarctos Heude, Mdm. Hist. Nat. Emp. Chinois, iv, pt. i, p. 18
(yesoc7isis).

Tijpe species. — Ursus arctos Linnteus.

Geographical distribution. — Northern hemisphere from northern
limits of the great continental areas south to the Atlas Moun-
tains, the Himalayas and Mexico.

Characters. — Dental formula : i ^\ c J"*, pm '^-*, m P^ = 42 ;
inner upper incisor well developed, permanent ; first, second and
third premolars in both jaws small, single-rooted, readily deciduous,
especially jj?(i'-, j»/i._, and jjvHj ; molars large and robust, the length
of the two upjier teeth together eqiial to width of palate.

I{e7nar]cs. — The genus TJrsus as thus restricted is a very
homogeneous gi'oup practically confined to the north temperate
region. The species are at present so impei-fectly knowm that no
fair estimate can be made of their number. Recently about
thirty forms have been recognized, only one of which is definitely
known to occur in Europe.

URSUS ARCTOS Linnreus.

1758. [LVsifs] arctos Linnreus, Syst. Nat., i, 10th ed., p. 47 (Sweden).
1772. [JJrsus'] ursus Boddaert, Kortbegrip van het zamenstel der Natuur,

I, p. 4C (Renaming of arctos).
1788. [Ursus arctos'\ a nigcr Gmelin, Syst. Nat., i, 13th ed., p. 100

(Northern Europe).

286 CAENIVORA

1788. [Ursus ardos'] fi fiiscus Gmelin, Sj-st. Nat., i, 13th ed., p. 100 (Alps).

1778. [UrsHS arctos] y albus Gmelin, Syst. Nat., i, 13th od., p. 100
(Unknown ; based on the " ours blano terrestre " of Buff on).

1792. U'[rs;(s] arctos griscus Kerr, Anim. Kingd., p. 184 (Germany ; also in
northern North America).

1797. Ursjis arctos rufus Borkhauseu, Deutsche Fauna, i, p. 40 (Swiss and

Tirolean Alps).

1798. Ursus baclius Schrauk, Fauna Boica, I, p. 55 (Forests on the

Bohemian boundary).
1808. Ursus fuscus Ticdemann, Zoologie, i, p. 374 (Substitute for arctos).

1814. Ursus alpinus Fischer, Zoognosia, iii, p. 161 (Alps? Based on an

individual seen alive in Paris).
1820. Ursus arctos ma jar Nilsson, Skand. Fauna, i, p. 112 (Wooded portions

of southern Scandinavia).
1820. Ursus arctos minor Nilsson, Skand. Fauna, i, p. 123 (Northernmost

Scandinavia).
1827. [Ursus arctos] & brunncus Billberg, Synopsis FauniB Scandinavia?,

p. 15 (Northern Scandinavia).
1827. [Ursus arctos] y annulatus Billberg, Synopsis Faunre Scandinavia?,

p. 15 (Northern Scandinavia).
1827. [Ursus arctos] 5 argenteus Billberg, Synopsis Faunae Scandinavite,

p. 15 (Northern Scandinavia).

1827. [Ursus] otii/rmcphagus Billberg, Synopsis Faunae Scandinavise, p. IG

(Northern Scandinavia).

1828. [Ursus] formicarius Billberg, Synopsis Faunse Scandinavise, 2nd ed.,

p. 16 (Renaming of myrmephagus).

1829. U[rsus] pyrcnaicns Fischer, Synopsis Mamm., p. 142. Latinization

of "Ours des Pyrenees" F. Cuvier, Hist. Nat. des Mammif., v,

fasc. 44, 1824 (Asturias, Spain).
1829. U[rsus] norvcgicus Fischer, Synopsis Mamm., p. 142. Latinization

of "Ours de Norwege " F. Cuvier, Hist. Nat. des Mammif., ii,

fasc. 7, 1819 (Norway).
1836. ? ZJ;-s?(s/aZci5(C)- Reichenbach, Regn. Anim. Icon., i, p. 82 ("Pyrenees" ;

afterwards supposed to be an individual of " 17. ferox." See Natur-

gesch. des In- und Auslands, Raubsiiugeth., p. 299, 1852).
1842. Ursus pyrenxus F. Cuvier, Hist. Nat. des Mamm., Tabl. gen., p. 3

(Described in fasc. 44, 1824) (Asturias, Spain).
1847. ? Ursus curyrhinus Nilsson, Skand. Fauna, i, 2nd ed., p. 212 (Sweden ?

Type an individual raised in captivity).
1857. Ursus arctos Blasius, Siiugethiere Deutschlands, p. 196.
1801. Ursus arctos aureus Fitzinger, Wissensch.-pop. Naturgesch. dcr

Siiugeth., I, p. 372 (Norway).
1864. [Ursus arctos] var. 1. normalis Gray, Proc. Zool. Soc, London, p. 082

(Renaming of arctos).
1864. [Ursus arctos] sub-var. a. scandinavicus Gray, Proc. Zool. Soc,

London, p. 082 (Based on Nilsson, Ilium. Fig. Skand. Fauna,

pi. 23).
1864. ? [Ursus arctos] sub-var. c. rossicus Gray, Proc. Zool. Soc, London,

p. 682 {nomcn mulum).
1864. [Ursus arctos] sub-var. f. polonicus Gray, Proc. Zool. Soc, London,

p. 682' (Poland ; based on Cuvier, Oss. Fossiles, iv, p. 332, pi. xxii,

fig. 3).
1864. [Ursus arctos] var. 2. grandis Gray, Proc. Zool. Soc, London, p. 684

("North of Europe " ; based on " a male purchased at Hull, living

iu the Zoological Gardens from 1852 to 1863").

uRsus 287

1804. [Urstts arctos] var. 4. stenorostris Gray, Proc. Zool. Soc, London,

p. G85 (Poland ; based on Cuvier, Oss. Fossiles, iv, p. 332, 2ad var.,

pi. xxii, fig. 4).
1864. Myrmarctos evcrsmanni Gray, Proc. Zool. Soc, London, p. C95

(Norway).
1910. Ursus arctos, U. arctos formicarius, U. arctos alpinus, and JJ. arctos

pyrenaicus Trouessart, Faune Mamm. d'Europe, pp. 67-08.

Type locality. — Sweden.

Geographical distribution. — Entire continent of Europe wher-
ever sufficiently extensive forests remain ; east into Asia ; west
formerly to Great Britain, where it became extinct about the
eleventh century ; not certainly known to have occurred in
Ireland.

Diagnosis. — Size moderate, condylobasal length of skull
ranging from about 260 to 350 mm. ; interorbital region notice-
ably elevated, the frontal profile strongly convex ; mesopterygoid
region not specially shortened and broadened, the width between
pterygoids decidedly less than half distance from hamular to
level of last molar ; colour brown or buffy, varying much in exact
shade, the legs usually darker than body, and feet darker than
legs.

External characters. — General form short and heavy, this
made more apparent by the long rather loose fur. Head
moderately pointed, rather broad posteriorly ; ear short, narrowly
rounded off above, nearly concealed in the fur, its tip not extend-
ing to eye when laid forward ; muzzle squarely truncate, its piad
naked, the upper border somewhat projecting backward, its lower
border separated from upper lip by a broad hairy area crossed
at middle by a nearly bare perpendicular line. Fore foot with
digits robust, inconspicuously graduated, the third and fourth
sub-equal and loiagest, the fifth and second sub-equal and slightly
shorter, the first with anterior edge of ball extending about to
middle of that of second, this interval greater than in the case
of the other digits ; claws strongly curved, blunt, without evident
cutting edges, their length at least twice that of those on hind
foot ; balls of digits large, pad-like, their surface, like that of
pads, coarsely rugose ; main pad wider than long, covering more
than half surface of palm, its outer border about twice as long as
inner, its porterior border slightly concave, its inner portion, at
base of thumb, marked off from rest of pad by a slight furrow ;
region between main pad and balls of digits densely furred ;
wrist pad about as large as ball of digits, near outer margin of
palm, its long diameter transverse ; region between wrist-pad and
main pad densely furred ; hairs along edge of palm standing out
stiffly, especially on outer side. Hind foot longer than fore foot,
the second and third digits sub-equal and longest, the first and
fourth slightly shorter, the fifth with anterior edge of ball at
middle of that of fourth ; pad like that of fore foot, but with a
broad backward extension passing along inner side nearly or

288

CARNIVORA

quite to heel ; region between pad and balls of toes, and at outer
side of backward extension densely furred ; fringe along edge of
foot conspicuous. Tail very short, concealed in the fur.

Colour. — The colour of body is usually a light brown or dull
buff, the head not essentially different, but feet and outer surface
of legs darker. Many individual difierences in colour have been
described, some of which are probably characteristic of geographical
races.

Skull. — Genera] form of skull rather robust, the rostrum

Via. 50.
i^r.^us arctos.

moderately long (distance from orbit to front of premaxillary
contained about 2^ times in condylobasal length), the brain-case

289

deep (depth to level of under side of poistglenoid process con-
siderably more than distance from tip of postorbital process to
middle of intei-parietal) but not unusually wide (mastoid breadtli
slightly exceeding depth to under side of postglenoid process).
Dorsal profile usually with an evident concavity in interorbital
region, but this character showing much variation ; highest point
at bregma or slightly further forward, the profile nearly straight
and sloping away at an angle of about 20° behind tliis j^oint to
slightly overhanging lambdal region ; ventral profile faintly and

Fig. 51.
Uistis arcton. x J.

evenly concave throughout. Brain-case broadly ovate in outline, its
greatest breadth about equal to distance from bregma to lambda,"
its depth at middle, exclusive of sagittal crest, slightly less than
greatest breadth ; lambdoid and sagittal crests well developed in
adults, tlie lambda noticeably projecting so that occiptal condyles
are scarcely visible when skull is viewed from above, and region
between crest and foramen magnum is deeply concave ; sagittal
crest dividing in region of bregma into two ridges, one of which

* Except in very old individuals in which the lambda is greatly
produced backward.

U

-i90 CAlRNn'ORA

runs to extremity of each jaostorbital process, the hinder margin
of which it forms ; occiput when viewed from behind moderately
broad, the depth from lambda to lower lip of foramen magnum
contained If to 1| times in mastoid breadth ; mastoid processes,
paroceipital processes, and condyles extending to about the same

level. Floor of brain case nearly flat, the basioccipital with
raised edges applied to inner surface of bullw, and in some
specimens marked by an evident concavity on each side of median
line ; auditory bulla flat, not rising above edge of basioccipital, the
greatest longitudinal diameter less than transverse diameter, the

URSUS 291

meatus distinctly tubular, usually longer than wide ; postglenoid
process heavy, rising to level of hamulars. Interorbital region
broad, the width across robust, triangular, postoi-bital processes
about equal to that of brain-case, the region immediately between
orbits always a little concave and sometimes conspicuously so,
that at base of each postorbital pi'ocess usually somewhat swollen.
Zygomata moderately expanded, the greatest zygomatic breadth
opposite anterior glenoid edge ; orbital process well developed,
liostrum equal to less than half condyloljasal length of skull, the
width across alveoli of canines equal to or less than depth at
front of orbit, the depth at front of nasal equal to about half
distance from orbit to front of premaxillary ; nares rather large,
their lateral margins slightly everted ; nasal bones elongate wedge-
shaped, squarely truncate anteriorly, their posterior extremity on
level with or extending slightly behind nasal branch of maxillary ;
anteorbital foramen over metacone of m^ or paracone of nr ;
palate nari-ow, its width between posterioi' molars contained about
3^ times in median length ; extension behind molars nearly
parallel-sided, its length equal to about three-quarters bi-eadth ;
mesopterygoid space 1 A- times to twice as long as wide, its anterior
border squarish or I'ounded, its lateral borders nearly parallel ;
hamulars small but distinct, slightly hooked outward. Man-
dible robust, the depth of ramus behind large premolar con-
tained about five times in length, the height of jjosterior portion
(measured to level of lower border) a little less than half length ;
coronoid process broad, its width at level of alveolus slightly
greater than height, the anterior border at first straight then
evenly convex to overhanging tip, the posterior border concave ;
angular process short, extending slightly if at all behind level of
articular process, its inner border nearly straight, its outer border
convex.

Teeth. — The teeth are moderately large relatively to size of
skull. Upper incisors forming a continuous row, separated at
each side from canine by a diastema about as wide as inner
incisor ; i^ and P sub-equal, the former slightly the smaller, the
anterior face smoothly rounded, a little more than half as wide
as high, the posterior face abruptly concave, with shelf-like
posterior extension, the cingulum slightly de^ eloped and forming
a rudimentary nodule on inner and outer side of /- and on outer
side of i^ near level, of middle of anterior surface ; P with crown
area nearly double that of r, and height nearly half that of
canine, its anterior surface smoothly rounded but with pronounced
nodule on inner side, its posterior sui-face gradually concave
and without shelf-like extension ; a thickened ridge along its inner
border and a low but somewhat trenchant ridge slightly outside
of middle. Lower incisors forming continuous row between
canines, their crown area increasing regularly from first to third,
their height approximately equal ; each has a high inner cusp and
a low outer tubercle best developed in ('3 ; posterior border slightly

u 2

292

CARNIVOEA

Fig. &3.
Wrsus arctos. Teetli nat. size.

URsus 293

concave, with inner, middle and outer ridges, the inner and middle
low and confluent in i\. Canines large and strong, oval or
slightly ovate in cross section, the longest diameter at level of
alveolus equal to or slightly greater than distance from alveolus
to median line of palate ; lower canine shorter and a little more
curved than upper ; a slightly de^■eloped posterior and antero-
internal longitudinal ridge, most evident in upper tooth ; no
cingulum. First and third upper premolars small, flat topped,
with slightly indicated cusp and posterior and antero-internal
ridge, the crown area slightly less than that of smallest incisor,
the first close to canine, the third close to large premolar ; third
near middle of space between first and second, much smaller than
the others and frequently deciduous, its crown indefinitely
rounded ; fourth upper premolar with crown area about half that
of first molar, its general outline triangular with apex directed
forward, the outer side longest, the posterior border shortest, the
contrast between them sometimes noticeable, in other instances
slight, the three cusps lying near respective angles, the anterior
highest (reaching level of main cusps of molars) and most robust,
the posterointernal and jiostero-external abruptly smaller, sub-
equal, the inner usually lower than the outer ; a small accessory
tubercle usually present at posterior base of postero-outer cusp ;
cingulum obsolete but usually visible along inner base of anterior
cusp and outer base of postero-external cusp ; first and second
lower premolars approximately like corresponding upper teeth in
both size, form and position ; third very early deciduous, usually
if not always absent in adult individuals ; fourth in contact with
first molar and with from one-third to nearly one-half its crown
area, the outline irregularly quadrilateral with well develoj^ed
antero-external cusp nearly as high as main cusps of molars, a
rudimentary antero-internal cingulum cusp, and a tuberculated
I'idge extending along outer side of crown from antero-external
cusp to posterior border ; occasionally a similar ridge is present
on inner side of crown, its anterior tubercle forming an evident
cusp at inner posterior base of main cusp. First upper molar
with crown much less than twice as long as broad, its outer side
bi-convex, its inner side evenly rounded, its two outer cusps
sub-equal in both height and diameter, the two inner cusps
decidedly lower than outer, and less well defined, owing to the
presence of a low, ridge-like tubercle between them ; both outer
and inner cusps when unworn have distinct though low anterior
and posterior trenchant ridge ; space between outer and inner
cusps occupied by a rather well defined longitudinal groove, the
surface of which is marked by irregular low ridges and furrows \
cingulum obsolete, but indicated in the regions between the
cusps ; second upper niolar nearly twice as long as broad, the
anterior two-thirds approximately like first molar, with the same
four cusps and intermediate longitudinal groove, the main axis
of which is, however, in axis of tooth-row instead of slightly

294

CARNIVORA

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296 CARNIVOEA

oblique to it ; postero-external cusp slightly smaller than antero-
external, and contrast between outer and inner cusps less marked
than in in^ ; posterior third of crown occupied by a flattened heel,
variable in form and size, but usually narrowing off, chiefly by
slanting inward of outer border, to about half anterior width of
tooth, and in some specimens bearing a low but evident third
inner tubercle ; surface of heel sculptured by irregular small
tubercles and furrows ; cingulum obsolete but usually evident
along anterior half of inner border. First lower molar about as
long as second but noticeably narrower, its crown showing more
traces of the jsrimitive trituberculate form than any of the other
teeth ; protoconid and hypoconid wide apart, separated by a deep
groove, the protoconid the highest cusp in the tooth, and with
evident antero-external commissure ; paraconid forming narrow
anterior extremity of crown and provided with a distinct com-
missure, similar to and joining that of protoconid ; metaconid
subterete, without commissure, near to and slightly behind inner
base of protoconid, a minute though evident accessory tubercle
just in front of it ; entoconid like hypoconid, at extreme posterior
edge of crown, the deep wide groove between it and metaconid
with small accessory tubercle at its lowest point. Second lower
molar about 1^ times as long as wide, its outline an irregular
parallelogram with rounded-off corners, the siirface of the crown
occupied principally by a flattened, irregularly sculptured crushing
surface, the cusps near border ; Ave cusps are usually well
developed : a rather large antero-internal and antero-external
opposite each other, and joined by a low transverse ridge ; a
small postero-external and two smaller postero-internal cusps ; a
small but evident accessory tubercle at anterior base of large
antero-internal cusp, this tubercle not infrequently dividing into
two. Third lower molar varying from rounded-triangular to
ovate in outline, its area about two-thirds that of riin, its surface
entirely flat excejit for a slightly raised rim which forms a
small antero-internal cusp.

Measurements. — Adult male from Sweden (mounted) : head
and body, 1900 ; tail, SO; hind foot, 195 ; ear, 90. For cranial
measurements see Table, p. 294.

Specimens examined. — Eight, from the following localities : —
Sweden : Kvickjock, Norbotten, 1 ; bo exact locality, 3 (B.M. and
U.S.N.M.).

Switzerland : Engadine, 1.
Adsteia-Hungaey : Near Hatszeg, Hunyad, 3.

Bcmarhs. — As regards the existence of geographical races of
the large European bear it is impossible to form any opinion on
the basis of the few sjiecimens seen. Ur.'ius nrctos is related to
the grizzly bear of North America, U. horribilis and its local
forms, but is readily distinguishable by the relatively greater
height of the frontal region and the consequently more abrupt
slope of posterior half of dorsal profile, a character suggesting

TIIALARCTOS

297

Umus richarfhnni ; forehead in most specimens rising tabriiptly
al)Ove level of rostrum so as to produce a noticeable concavity in
dorsal profile, but this character subject to marked variations,
the exact nature of which is not fully understood. The skulls
of the two species are of approximately the same size, those of
U. horrihilis perhaps averaging somewhat the larger. The teeth
of the two animals are also much alike.

9. Kvickjock, Norbotten, Stockholm Museum 90. 8. 1. 3.

Sweden. (e).

2 S. Sweden. (Lloyd.) Purchased (Stevens). 02. 3. 29. 7-8.

9 St. Engadine, Switzerland. H. Justen (p). 86. 1. 23. 1.

3 skeletons. Hatszeg, Transylvania, C. G. Danford (c). 78. 1. 16. 1-3.
Hungary.

Note on the Ursus formicarius of Bieler.

A peculiar small bear supposed to have been taken in the
Canton of Grisons, Switzerland, has been described by Professor
Bieler of the Lausanne Agricultural College as Ursus formicarius
Eversmann.* Through the author's kindness I have had the
opportunity to examine this skull. It is that of a rather young
individual, apparently a female, with basal suture open, but with
teeth showing slight indications of wear. In size it is smaller
than in a female of the same age or slightly younger from
Sweden (see Table of cranial measurements, p. 294), the inter-
orbital region is much depressed, so that the frontal profile is
nearly flat, and the mesopterygoid fossa is unusually broad and
short, the width between pterygoids equal to a little more than
half distance from hamular to level of last molar. The teeth on
the other hand are slightly larger than usual in females (see
Table, p. 295) ; inform they show no special peculiarities. Small
bears, presumably of this type, have been reported from Spain,
northern Italy, Russia and Scandinavia, and have formed
the basis of such names as Ursus formicarius, U. arctos minor
and Myrmarctos eversmanni. Until more is known of them, as
well as of the normal variation in ordinacy Ursus arctos^ their
status must remain in doubt.

Genus THALARCTOS Gray.

1825. Tlialarctos Gray, Ann. of Philosophy, N.S., x, p. 62, July, 1825.
1825. Thalassarctos Gray, Ann. of Philosophy, N.S., x, p. 339, November,

1825.
1896. TJialassiarcJius Kobclt, Boricht Senckenberg. naturforsch. Gesellsch.

Frankfurt am JIain, p. 93 (Substitute for Tlialarctos).

Type species. — Tlialarctos polaris Gray = Ursus viariiimns
Phipps.

* Compto-Rendu des Stances du Sixi^me Congrus Internationale do
Zoologie, Berne, 1904, p. 248. 1905.

298

CARNIVORA

G(jo(jraiiltical distribution. — North Polar region, south to
northernmost continental coasts.

Characters. — Similar to Ursus, but cheek-teeth much less
robust, the combined length of the two upjaer molars not equal
to width of palate, and canines and incisors enlarged and more
prehensive in general form.

Remarks. — The genus Thalarcios, though not very strongly
dift'erentiated from Ursus, is a well defined and perfectly natural
group.

THALARCIOS MARITIMUS Phipps.

1774. Ursics maritimns Phipps, Voyage toward North Polo, p. 185 (Spitz-

bcrgen).
1776. Ursus mariiius Pallas, Eeise durch verschiedeue Provinzen des

russischen Beichs, iii, p. 691 (Arctic Ocean).
1792. Ursus folaris Shaw, ^Museum Levorianum, i, p. 7 (P^enaming of

marinus).
1862. Thalarctos maritimus Gray, Catal. Bones Mamm. Brit. JIus., p. 105.
1908. ? Thalassarctos jenacnsis Knottnerus-Meyer, Sitz.-Ber. Gesellsch.

Naturforsch. Freunde, Berlin, p. 184, July, 1908 (Jena Island,

Spitzbergen).
1908. ? Thalassarctos spitzbergensis Knottnerus-Meyer, Sitz.-Ber. Gesellsch.

Naturforsch. Freunde, Berlin, p. 184, July, 1908 (Seven Island,

Spitzbergen).
1910. Ursus [Thalassarctos) maritimus Trouessart, Faune Mamm. d'Europe,

p. 66.

Tijpe locality. — Spitzbergen.

Geographical distrihution. — Arctic Ocean, south on floating ice
occasionally to the northern coast of Norway. Details of
distribution unknown.

Diagnosis. — General characters as in the genus ; size very
lai'ge ; colour uniform whitish or buffy.

External characters. — Form lijnger and less heavy than in
Ursus arcios, the neck noticeably longer and head longer and
more pointed ; ear actually as well as relatively shorter ; fore
foot with palmar tubercles and balls of toes essentially as in
U. arctos but smaller ; pad on hind foot without backward
continuation along inner portion of sole ; claws much less
elongated than in U. arctos, not strongly curved, but with acute
jjoints and well developed cutting edges. Fur very dense, its
texture almost seal-like in the short summer coat.

Colour. — Entire animal a uniform whitish or buffy, the winter
pelage tending to be a creamy-white, the summer coat yellowish
buff.

Skull. — The skull is considerably larger than that of Ursus
arcios, with relatively longer brain-case, deeper, wider rostrum,
and less elevated frontal region ; lambdal region less produced
backward than in Ursus arctos, the condyles usually visible when
skull is viewed from above. Base of brain-case essentially as in
U. arctos, but portion at base of condyles more narrowed and

THALARC'TOS

299

elongate. Palate iioticea1)ly broadei' than in Uisus arctos, a
character made more conspicuous by the relative weakness of the
teeth. Mandible with no sj)ecial [leculiarities except that the

FIO. 54.
Thalnrclos maritimiis.

lower margin is nearly straight thi'oughout, the posterior concavity
being very slightl}' indicated, and lower border of angular process
only a little elevated above general outline.

Teeth. — While the general character of the dentition differs

300

CAKNIVOEA

notably from that of Ursus arctos in the reduction of the molars
and increased size and prehensiveness of the canines and incisors,
the details of the individual teeth present little that is specially

FIO. 55.
Thalarctos maritimus. x ;.

noteworthy. General form of upper incisors as in Ursus arctos,
but points of i^ and i'^ narrower and more hooked backward ; P
with cusp more slender and ridges nearly obsolete ; lower incisors

THALARCTOS

301

with lobes more sharply defined. Canines both above and below
essentially similar to those of Ursas arcfos, except for their greater
size. Owing to the greater width of palate the proportion of

FlO. 5C.
Thalarctos maritimus.

diameter of upper canine to palatal width is about as in the
smaller-toothed animal. Small premolars showing no special
peculiarities. Large upper premolar with relati\ely higher

302

CAP.NIVORA

Thalardos marUimux. Teetli iiat. size.

THALAKCTOS

303

anterioi' cusp than in Z7. nrctos, its inner side more flattened,
giving tlie tooth a more carnassial appearance ; postero-internal
cusp relatively less developed. Large lower premolar essentially
as in U. arctos but with somewhat more slender cusp. Molars
differing from those of Ursus arctos in their smoother, less sculp-
tured crushing surface, and slightly more trenchant cusps. Form
of viy not peculiar, though outer cusps are higher and narrower
and inner cusps relatively lower ; «!- with inner cusjjs obsolete
and heel relatively narrower and less developed. Anterior lower
molar with metaconid and its accessory tubercle reduced to a low
irregularly tuberculate ridge ; commissure of protoconid and
paraconid obsolete ; hypoconid and entoconid smaller and much
nearer together than in Urms arctos, though separated from
anterior cusps by a normally wide interval, in which, however,
there are no definitely formed accessory tubercles. Second lower
molar with the same elements as in U. arctos except for the
absence of all trace of an intermediary tubercle on inner side of
crown. Third lower molar with crown nearly flat, its margin
showing only the faintest trace of antero-internal and antero-
external elevations.

Measurements. — Adult male from Behi'ing Strait (mounted) :
head and body, 2670 ; tail, 90 ; hind foot, 370 ; ear, 80. For
cranial measurements see Table, p. 294.

Specijiiens examined. — Nine, from the following localities: — Spitzbergen,
1 (U.S.N.M.); Griffin Bay, Wellington Channel, 1; Melville Island, 1;
Arctic Ocean, 1 ; no history, 5.

[The Museum specimens appear all to have come from the
American side of the Atlantic]

Family CANID^.

1821. Canidx Gray, London Medical Repository, xv, p. 301, April 1, 1S21.

Oeographico I distrihution. — Essentially cosmopolitan ; in E urope
west to Ireland.

Characters. — Larger cheek-teeth of a combined trenchant and
crushing type, the last upper premolar and first lower molar
strongly differentiated as carnassials, the former 3-rooted, its
inner lobe in front of middle of crown, its position, somewhat
posterior to level of anteorbital foramen, at point of greatest
mechanical efficiency ; auditory bulla moderate!)' or considerably
inflated, without septum ; form rather light, the legs long ; size
moderate ; feet digitigrade ; toes, 5-4 or 4-4.

licinarlcs. — Notwithstanding its wide disti'ibution the family
Caniiiue is not rich in genera. About a dozen are now recognized,
three of which occur in Europe.

304

CAENIVOEA

KEY TO THE EUROPEAN GENERA OF CANID.E.

Interorbital region distinctly elevated ; postorbital processes

convex above ; pupil of eye round Ca«is, p. 304.

Interorbital region not elevated ; postorbital processes not
convex above ; pupil of eye elliptical.
Postorbital processes flat or very slightly concave above ;
forehead rising abruptly above level of rostrum ; ear

rounded Alopex, p. 318.

Postorbital processes distinctly concave above ; forehead

rising gradually above level of rostrum ; ear pointed VuIjm's, p. 325.

Genus CANIS Linnasus.

1758. Ganis Linnaeus, Syst. Nat., i, 10th ed., p. 38 (type by tautouymy

C. familiaris).
1810. Lupus Oken, Lenrb. d. Naturgesch., iii, pt. 2, p. 1039 [Canis lupus,

by tautouymy).
1837. Vulp)icanis Blainville, Ann. Sci. Nat., Paris, 2nd ser., Zool., viii,

p. 279, November, 1837 (Canis auretts Linngeus).
1839. Lyciscus H. Smith, Jardine'a Naturalists' Library, Mammals, ix,

p. 160 {Canis latrans Say).
1839. Thous H. Smith, Jardine's Naturalists' Library, Mammals, ix,

p. 193 (Canis antlms F. Cuvier).
1839. Sacalius H. Smith, Jardine's Naturalists' Library, Mammals, ix,

X3. 213 (Canis aureus Linnaeus).
1841. Oxygous Hodgson, Calcutta Journ. Nat. Hist., ii, p. 213 (Canis

aureus Linnaeus).
1857. Canis Blasius, Saugethiere Deutschlands, p. 177.
1868. Neocyon Gray, Proc. Zool. Soc, London, p. 506 (Canis latrans Say).
18G9. Dieba Gray, Catal. Cam. Pachyd. and Edentate Mamm. Brit. Mus.,

p. 180 (Canis anthus F. Cuvier).

Type species. — Canis famiUaris Liniifeus.

Geographical distribution. — Nearly as in the family, but
absent from the Malay Archipelago and South America ; in
Europe west within historic times to Great Britain, but now
restricted to the continent.

Characters. — Skull heavy and deep (depth of brain-case more
than one-third condylobasal length) ; interoi-bital region thickened
and elevated, the frontal sinuses i-ather large, the postorbital
processes thick, convex above, their edges rounded off ; dorsal
profile of forehead rising rather abruptly and noticeably above
level of rostrum ; dental formula : i ||, c j^J, ^jwt t^, in |^ = 42 ;
teeth heavy and large, the length of carnassial and upper molars
together contained about 21 times in palatal length ; canines
robust and not specially elongated, the point of ujjper tooth
extending scarcely beyond middle of mandibular ramus when
jaws are closed (fig. 65).

Bemarks. — Much uncertainty exists at present with regard
to the limits of the genus Canis. As here defined the group
includes the domestic dogs, the true wolves, the American prairie

CANIS

305

wolves, .-u.d the Old World jackals. Two species are known

%o:::;l^ :;::::''' - ^^ -^^-^^ ^« -p— ^^d ,y seve.ai

KEY TO THE EUROPEAN FORMS OP CANIS.

Condylobasal length of skull less than 200 mm • teeth
not so large as in the largest domestic dogs ('len"th
of upper carnassial 17 to 18 mm.); cingulum on
outer margm of m' broad and conspicuous (South-
eastern Europe ; Jackal) ^ r n„.

Condylobasal length of skull more " than Iw' mm"- ^- «"'^'««' P' 315.
teeth larger than in the largest domestic dogs'
(length of upper carnassial 25 to 27 mm.) • cingu-
lum on outer border of m' narrow, tending to bo
mcompleto at middle (Distribution general • true
Wolves) r 1

Size rather small "(exact' "dViension's'^unknowni '^' '^"^'

(Southern Spam n , , ■,

Size large, C. l. dettanus, ]). 615.

White of throat not extending uninterruptedly
WI,ih?nfn.^^?*''v^'''^"°'"'^°™ Europe) ... C.Llu,,„.,r, 313
White of throat extending uninterruptedly on to

cheek (Spain except extreme south) c. I. signatus, p. 314.

CANIS LUPUS Linnc-eus.
(Synonymy under subspecies.)
G«.i,r«y./caZ d/.<n-J«fion.-Originally throughout Europe from

SrfttwT r'in°'^,^"Tx.^^'''^' °«^^' exterminated in the
British Islands, Holland and Denmark

/09,f/"T'''-""^°°'ly^^^''''^ l^'^gth of skull more than 200 mm
(220 to 250 mm.) ; cheek-teeth larger than in the largest racS
o domestic dogs the upper carnassial 25 to 27 mm. In length
but structure not peculiar, the upper molars with narrow, incon!
spicuous cingulum on outer side (licr 61)

d^„fH'7,^;!/-^";™tr-^^e^«ral Form essentially as in doitestic
Sed if '? X'V ^^'' ^«derately long, erect, somewhat

Z h ' ^■^^"'^"^g '^^^""V" 'y' ''^'''^ ^^''^ ^«^^^^"'^^- Muzzle pad
completely bare. Fore foot with third and fourth digits s^ib-
e.]ual and longest, second and fifth shorter, the large Vd-like

tdtlvtHfol rl^^f^''" ?^"^^ "^ *^"'-^^ -"d fourth and the

fkof If ' hfi't-shaped main pad, the combined area of

u o digits greater than that of pad; thumb much shorter

than other digits, the nail smaller, but not peculiar in form its

extremity not reaching level of posterior boiler of ma n pa i

1 scarcely indicated, no pad at its base; wrist pad single, near
Hini w' ^y^'^^fTf^^h^t more than half that of bait of toes.
•Uion/ rf''"*'''"7 ^'^ ^? ^"^^' ^^"t ^^'^""-^ '-^"d posterior pad

Padsnndfir' '"^"f' 'l^'^^l^^ '''''''''^' .s"b-e.,ual throughout,
lads and balls narrowly edged with short hair

of legs yellowish brown or buff, darker along median region of

X

306

CARNIVORA

back, on posterior portion of head and outer surface of ears,
lighter and more incHned toward greyish at sides of shoulders
and between ears and eyes ; longer hairs of back and sides black-
tipped, producing an evident dark shading over middle of back.

I'IG. 5S.
Canis lujms. x i.

especially behind middle and at base of tail ; pencil narrowly
clear black, rest of tail essentially like back. Underparts and
inner surface of legs pale buft" or buffy white, not strongly
contrasted with sides, the chin and inturraiuia usually grizzled.

CANIS 307

frequently margined with blackish ; upper lip to muzzle pad and
including lower half of cheek dull whitish, usually not very
different from throat ; inner surface of ear light buff.

Skull . — In general aspect the skull diHers slightly if at all

I'lG. 59.
Canis liquis. x *.

from that of some of the larger races of domestic dogs, though
often attaining a greater size. The rostrum, however, appears
to bo relatively less rtibust than in dog skulls of approximately
the same length. Dorsal profile rising gradually from nares to

308

CAIINIVORA

just in front of orbit, then abruptly to a little in front of ])regma,
behind which it is nearly flat to strongly overhanging lambdal
region. Depth of brain-case through auditory bulla about 2h

Fig. CO.
Canis lupus.

times that of rostrum behind canine, and about equal to mastoid
breadth. Brain-case rather elongate ovate in outline when
viewed from above, its breadth above roots of zygomata about
l.", times that of rostrum over canines and apj^roximately equal

CANIS 309

to distance from bregma to most posterior point of occiput.
Posterior portion of occiput strongly concave when viewed from
the side, tlio condyles nearly hidden beneath the projecting
lambdal region. Floor of brain-case with no specially noteworthy
features, the auditory bullte sub-circular in outline, with short but
evident meatal tube, their surface evenly inflated (more so than
usual in domestic dogs) except for an evident flattening on antero-
external aspect. Sagittal and lambdoid crests well developed,
the former dividing just in front of bregma into two ridges
curving outward to form posterior border of postorbital processes.
Interorbital region modei'ately elevated, well defined, with distinct
longitudinal concavity between raised and thickened postorbital
processes. Zygomata widely spreading, the greatest zygomatic
breadth (at level of anterior glenoid edge) a little more than half
greatest length of skull ; orbital process well developed, the
orbit surrounded by bone through about four-fifths of its
circumference. Rostral breadth at canine about equal to depth
at front of carnassial ; premaxillary extending posteriorly to
about middle of nasal ; mnxillary extending back nearly to middle
of orbit, slightly exceeded by nasal ; anteorbital foramen about
9x5 mm. in diameter, over posterior root of third premolar.
Palate moderately wide, nearly flat, not extending posteriorly
beyond level of last molar, terminating in an obscure median
spine ; incisive foramina between canines, 11 to 17 mm. in length,
their combined breadth usually a little less ; mesopterygoid fossa
rather more than one-third as long as palate, considerably
narrower posterioi'ly than anteriorly. Mandible strong, but not
remarkably robust, the depth at posterior edge of carnassial
contained about six times in length ; symphysis rather long ;
coronoid process high, the depth of mandible through its middle
noticeably greater than distance from last molar to back of
condyle ; angular process heavy, nearly horizontal, distinctly
raised above level of under margin of ramus.

Teeth. — The teeth are relatively larger than in any of the
races of domestic dogs, though in form they show no tangible
features by which they may be distinguished. Upper incisors form-
ing a continuous, slightly convex row, the outer tooth separated
from canine by a distinct space ; size, when viewed from in front,
increasing regularly from first to third, but third abruptly much
larger than the others in cross-section and nearly half the size of
canine ; anterior surface of i^ and /- slightly more than half as
wide as high, smoothl}' rounded ofl", the cutting edge narrow but
not acute ; a small but distinct secondary lobe at each side of
fi'ont aspect, that of inner side a little below middle, that of
outer side about ecjually above ; posterior sui'face of crown con-
ca\e longitudinally though without backward-projecting basal
shelf ; a well developed median longitudinal rib, and a low but
noticeable cingulum, the latter terminating abruptly and forming
the lobes seen in front view ; outer incisor with no secondary

310

CARNIVORA

lobes, its general form intermediate between that of canine and
of inner incisors. Lower incisors forming continuous row between
canines, the three teeth essentially alike in form, but increasing

no. Gi.

Canis lupua. Tectli nat. size.

regularly in size from iirst to third though less conspicuously
than in the case of the upper incisors ; crowns (viewed from in
front) about twice as high as wide, distinctly bilobed, the outer

CANIS 311

lobe scarcely half as wide as inner ; on inner tooth the outer lobe
is nearly level with cutting edge, on second it lies slightly above
middle of crown, and on third slightly below middle of crown ;
posterior surface oblique, slightly concave, with noticeable longi-
tudinal furrow extending back from notch between lobes. Canines
large, usually 15 mm. or more in diameter at alveolus and about
twice as long, a size rarely if ever attained in domestic dogs,
their surface smooth except for a low antero-internal and
posterior-median logitudinal ridge, the upper teeth slightly longer
and less recurved than the lower. Premolars Jiioderately spaced
except that j-im-* is nearly or quite in contact with the
carnassial ; first, second and third teeth essentially alike in the
two jaws, those of the mandible, however, slightly the less robust ;
first premolar both above and below single-rooted, the crown
simple, that of pm^ subterete, that of j»wi nearly twice as long as
broad, the height in both slightly less than length, the crown area
approximately the same as tbat of corresponding inner incisor, the
small cusp a little in front of middle and with slightly developed
anterior and posterior ridge. Second and third premolar's two-
rooted, the crown about twice as long as wide, sub-elliptical in
outline, the inner margin sometimes (especially in 2J'»h ^'^^ jwu'-)
slightly concave, the long axis nearly parallel with sagittal plane
except in pm^, which is obliquely set ; main cusp a little in front
of middle of crown, its height distinctly more than half length of
crown, its anterior and posterior cutting ridge well developed, the
posterior bearing a distinct secondary cusp situated over middle
of posterior root and relatively larger in lower than in upper
teeth ; a slight shelf -like projection behind secondary cusp ; pm^
similar to jjJHg but considerably larger, its secondary cusp
better develoi^ed and succeeded by a small but evident postero-
basal cusp sprin.ging from the posterior edge of crown ; cingulum
of all the smaller premolars complete though low and incon-
spicuous. Upper carnassial large and robust, the length of crown
along middle slightly more than twice greatest breadth exclusive
of antero-internal lobe, the main axis of the tooth extending
evidently through middle of crown, so that the small, cuspless
inner lobe stands as an offset, slightly breaking the symmetry of
the outline ; main cusp slightly behind middle of crown, its
height more than half length of tooth, its axis slanting distinctly
backward, its anterior and outer surfaces evenly convex except
for the rudimentary longitudinal ridge on basal two-thirds of
fi-ont, its inner surface, together with that of posterior cusp,
flattened ; posterior cusp low and robust, obscured by its very
high nearly horizontal commissure which meets the somewhat
shorter but eciually trenchant conuiiissure of main cusp at an
angle of about 75" ; cingulum complete, though low and incon-
spicuous. Lower carnassial narrower than upper but with eiiually
high crown, the most elevated portion in front of middle instead
of behind it ; protoconid large and robust, resembling main cusp

312

CARNIVORA

of upper carnassial, but with well developed cutting edge both
in front and behind ; paraconid near middle of anterior portion
of crown ; its general form like posterior cusp of upper carnassial,
its connnissure bearing essentially the same relation to that of
protoconid as in the case of the two cusps of the upper tooth,
except that tlie relative lengths of the cutting edges is reversed ;
metaconid small but evident, at postero-internal base of proto-
conid ; liypoconid and entoconid low, occupying the posterior edge
of a well developed though relatively small heel (area of heel
scarcely more than one-third that of anterior portion of tooth)
separated from the cusjjs of the main triangle by a wide transverse
groove ; crown area of entoconid equal to about half that of
hypoconid, its cusp apjaroximately the same size as that of meta-
conid. Second lower molar essentially like heel of carnassial
but larger, its two anterior cusps corresponding in size and form
with liypoconid and entoconid of the large tooth, the posterior
edge of its crown with a small outer cusp resembling the antero-
inner, and sometimes with a slightly developed inner ridge or
rudimentary fourth cusp. Third lower molar single-rooted, the
crown subterete, about as large as that of first premolar, with low
central cusp and rudimentary longitudinal ridge. First upper
molar large, with high outer two-cusped sectorial portion and low
inner crushing portion, the two areas sliarjjly differentiated, the
antero-posterior diameter of the outer decidedly greater than that
of inner ; 23aracone and metacone conical, terete, with slightly
developed anterior and 2:)osterior cutting ridges, the area and
height of metacone about two-thirds those of paracone, the width
of base of which is at least equal to width of inner portion of
tooth ; protocone very low, with low but distinct anterior and
posterior commissures, each of which joins cingulum at base of
corresponding large outer cusps, and each of which bears an
intermediate cusp soon disappearing with wear, the posterior
intermediate cusp larger and more definite in form than anterior
cusp ; hypocone ridge-like, at postero-inner border of crown,
separated from protocone and its posterior commissure by a deep
groove. Second upper molar with about half the crown area of
first, its elements essentially the same, though so reduced that
tlie paracone is scarcely larger than protocime of large tooth,
intermediate cusps on commissures of protocone are barely
indicated, and hypocone is not distinguishable as a cusp distinct
from the cingulum. In both molars the cingulum on outer border
is narrow and inconspicuous relatively to the broad cusps ; in
region between paracone and metacone of m^ it is usually
obsolete (compare figs. 61 and 62).

Remarks. — The material availal^Ie for study has been so poor
tliat I have found it impossible to come to any conclusion with
regard to the existence of local forms of the European Wolf.
The following races have been distinguished by INlr. Caljrera.
There seems to be no good reason to doubt their validity.

CANIS 313

Tlu! only known characters by which the skull of Canifi lupus
can be distiuguishcd from that of the larger domestic dogs is the
greater average general size and the relatively larger teeth. In
a dog's skull with condylobasal length of 2,30 mm. the length of
upper and lower carnassials is respectively 21 "6 and 25 "0 mm.
In ten skulls with condylobasal length of more than 200 mm.
the average and extremes for these teeth are : upper, 20 • 5
(19-22) ; lower, 24 • 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 Linnajus.

1758. [Cams'] lupus Linniseus, Syst. Nat., i, 10th ecL, p. 39 (Sweden).
1792. C[anis] lupus flavus Kerr, Anim. Kingd., p. 137 (France and

Germany).
1804. Canis lupus niger Hermann, Observ. Zool., p. 32. Not of Kerr,

1792 (Forest of Hagenau, Alsace, Germany).
1839. ? [Canis hqms'] var. canus do Selys-Longchamps, Etudes de Micro-

mamm., p. 144 (nomen nudum).
1839. ? [Canis lupus'] var. fulvus de S61ys-Longchamps, Etudes de Micro-

mamm., p. 144 (nomen nudum).
1841. Lupus oricntalis Wagner, Schreber's Saugthiere, Suppl., ii, p. 367

(Europe).
1857. Canis lupus Blasius, Saugethiere Deutschlands, p. 180.
1863. [Canis lupus] var. major Og6rien, Hist. Nat. du Jura, iii, p. 59

(Lower slopes of the Jura).
1863. [Canis lupus] var. viinor Og^rien, Hist. Nat. du Jura, in, p. 69

(Higher portions of the Jura).
1897. Canis lujnis minor Mojsisovics von Mojsvar, Thierlebeu der osterr.-

hung. Tiefebonen, p. 241 (Southern Hungary). Based on the

" Rohrwolf ," an animal supposed to be smaller and greyer than

true lupus.
1910. Canis lupus and C. lupus hjcaonf Trouessart, Faune ilamm.

d'Europe, p. 90.

Ti/pc locdlitij. — Sweden.

GccHjrupliiral distribution. — Northern and central Eui'upe,
exact limits of range unknown ; formex'ly west to Ireland.

Characters. — Size maximum for the species ; general colour

* Wingo (Danmarks Fauna, Pattedyr, p. 123, 1908) states that in the
skull of a dog from a prehistoric grave (Iron Age) in Denmark, the length
is 209, and that of the two carnassials 20 and 22-5 respectively, while in a
rather large modern "great Dane " the corresponding measurements arc
255, 22 and 28. This author (p. 124) regards the domestic dogs as derived
from Canis aureus.

t -Vpplicd to the wolf of the Pyrenees ; but Schreber's plate lxxxix,
the basis of the name, is a copy of BufEon's plate xli, representing an
animal brought alive to Paris from Canada.

314

CARNIVORA

not markedly tawny ; white of throat not extending to cheeks.
The few skulls examined agree with Asiatic specimens in having
the outer cusps of m^ moderately large, the paracone with trans-
verse diameter of base about equal to width of large flattened
jiortion of crown.

Measurements. — For cranial measurements see Table, j). 316.

Specimens examined. — Pour skulls, from the following localities : —
Sweden: No exact locality, 2 (U.S.N.M.).
Russia: No exact locality, 1 (U.S.N.M.).
Italy : Near SasscUo, Liguria, 1 (Genoa).

Canis lupus signatus Cabrera.

1907. Canis lupus signatus Cabrera, Bol. Real Soc. Espaii. Hist. Nat.,

Madrid, vii, p. 195.
1910. Canis htpus signatus Trouessart, Fauue Mamm. d'Europe, p. 91.

Ti/2)e localitjj. — Escorial, Madrid, Spain.

Geogrnjjhical distrihution. — Central Spain.

Characters. — Size and general appearance as in Canis Inpns
lupus ; colour a more tawny brown than in the northern animal,
particularly on muzzle ; white of throat extending unintei'iuptedly
to cheeks.*

Measurements. — Type (adult male), from Cabrera : head and
body, 1230 ; tail, 400 ; hind foot, 265 ; ear, 125. Young adult
male and female from Province of Burgos, Spain : head and
body, 1130 and 1180 ; tail, 350 and 380 ; hind foot, 225 and 220 ;
ear 120 and 115. For cranial measurements see Table, p. 316.

Specimens examined. — One from Seville, Spain, and two from Province
of Burgos, Spain.

liemarhs. — Tn dentition the Seville specimen differs from all
the other Old World wolves with which I have compared it in
the unusual development of the outer cusps of the upper molars.
The transverse diameter of paracone in m} conspicuously exceeds
width of the small inner 2:)ortion of tooth. Mr. Cabrera informs
me that the tyj^e shows much the same peculiarities. This
character is also present, though less pronounced, in the two
skulls from Burgos, which further differ from northern specimens
in the smaller size and more glol)ular form of the auditory
bulL-e.

i. Seville, Spain. (.1. Rjiiz.) Lord Lilford (p). 95. .3. 3. G.

6,9. Riocabado, Bm-gos. Hon. N. C. Roths- 11. 10. 5. 1-2.

(Rev. S. Gonzalez.) child (p).

* In the Burgos specimens the colour is not unusually tawny : back
and sides a coarse inixture of black, whitish, ochraceous-buff, and drab grey
(underfur), the black and whitish most conspicuous along back, the
ochraceous-buff ou legs and feet (clear and unmixed on latter) ; ear
ochraceous-rufous on outer side (darker and duller at tip), pallid
ochraceous-buff on inner surface ; throat and lower half of cheeks the same
pallid ochraceous-buff ; chin and interramia blackish.

CANIS . 315

Canis lupus deitanus Cabrera.

1907. Canis lupus deitanus Cabrera, Bol. Real Soc. Espafi. Hist Nat

Madrid, vii, p. 107. ''

1910. Canis lupus deitanus Troues.sart, Fauue Mamm. d'Europe, p. 91.

%>e Zorr</%.— Moratalla, Murcia, Spain.

Genf/mphical (h'.strihnt!on.~-l<^o\v known from the type locality
only. -^ •'

D!a<jnosis.~^miiUev than Canis hipm lupus and brighter in
colour, the general ai)pearance much as in C. aureus.
Measurements. — Unknown.

Specimens examined.~l have seen the two living examples in :\radrid
on which the form was based.

Iteniarh'i.^Tn general appearance the two Moratalla wolve.s
are strikingly different from Cam's lupm. Unfortunately their
ti-ue characters are not yet known.

CANIS AUREUS Linnteus.

1758. [Cams'] atireus Liouaeus, Syst. Nat., i, 10th ed., p. 40.
1835. Canis aureus var. moreotica I. Geoffroy, Exp6d.'sci. de Mov&e Zool
pi. I (Morea, Greece). ''

1841. C[anis] dalmatinus Wagner, Schreber's Saugthiere, Suppl ii n 383
(Dalmatia). '-'■ ' ' ^"

1841. C[anis-\ gr^'cus Wagner, Schreber's Saugthiere, Suppl ir p 383
(Peloponesus). ' '■'

1892. Canis aureus halcanicus Brusina, Glasnik Hrvatskoga Naravoslovno^a
Drustva, Zagreb, vii, p. 317 (Drava River, Croatia). "

Type locaUhj. — Province of Lar, Persia.

(ic(>[iraphical distribution.—Indin and westward through Asia
Min.)r to the Balkan Peninsula, north to Heves Comitat,
Hungary.

D/ag-no-s/s.— Smaller than Canis lupus (condylobasal len<^th of
skull less than 200 mm.) ; teeth not equal to those of the larger
domestic tlogs m size, the upper molars with wide, conspicuous

Icmgulum on outer .side (fig. 62).
_ CoZowr.— Upper parts buffy cinnamon, clouded by black hair
tips along dorsal region, nearly clear on sides, and becoming clear
bright cinnamon on outer surface of legs and ear and on area
around and behind base of ear ; muzzle more heavily washed
with black ; from middle of back to base of tail the cinnamon is
replaced by whitish, causing a rather noticeable contrast between
this region and the surrounding parts when fur is disarranged ■
tail essentially like l^ack, the basal half above greyish,°the
terminal half and underside bully much overlaid with black •
pencil blackish ; underparts and niner side of legs dull buffy

316

CARNIVOEA

o

1

Basal suture closed.
„ closed.

„ open.
„ closed.

OOOO O^^O ^CMCNOOCO
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255-0
232-0

235-0

221-0
234-0
226-0

158-0
157-0
151-4
148-0
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1035 U.S.N.M.
1037
910

Genoa

95. 3. 3. 6
11. 10. 5. 1.
11. 10. 5. 2.

8. 10. 2. 50
6. 5. 1. 31
6. 5. 1. 33
6. 5. 1. 34
6. 5. 1. 35

3

C. lupus lupus.

Sweden .

Northern Sweden .
Russia .

Italy: near Sassello,|
Liguria . j

C. lupus signatus.

Spain : near Seville
Prov. Burgos

C. aureus.

Greece : Pirreus
Asia ]Minor ; Khotz

317

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318

CARNIVORA

Skull. — The skull is much smaller than that of Cam's lupus.
In form it differs slightly in the less elevated frontal region and
somewhat more inflated auditory bulla?.

Teeth. — While agreeing with those of Cain's lupus in general

form and in the position of the cusps, the teeth

are on the whole more trenchant in character, as

shown by the general tendency toward narrowness

of crown and prominence of ridges. This is

particularly noticeable in the upper molars, in

which the large cusps are relatively higher, more

slender, and less terete than in Canis lupus, their

cutting ridges much more developed ; transverse

diameter of metacone in m^ noticeably less than

width of inner portion of crown ; cingulum on

outer border of both upper molars wide and

cons2:)icuous in contrast with narrow cusps,

KiG. 62. showing no tendency to become obsolete in

Canis aureus, regioia between paracone and metacone. Lower

^''cheek-teeth"^ carnassial with metaconid actually as well as

Nat. size. relatively larger than in Canis lupus, and posterior

heel with area equal to nearly half that of anterior

portion of tooth, its cus2:)s strongly developed.

Measurements. — For cranial measurements see Table, p. 316.

Siiccimens examined^ — -One from Greece (Piraeus) ; numerous others
from Asia Minor and India.

Bcmarlcs. — The single specimen from Greece agrees sufficiently
with a series of five from Khotz, near Trebizond, Asia Minor,
to make it appear unwise, in the absence of more satisfactory
material, to use one of the Balkan names.

1. Pirfcus, Greece. (C. Mottaz.) Hon. N. C. Roths- 8. 10. 2. 49-50.

child (p).

Genus ALOPEX Kaup.

1829. Alopcx Kaup, Eutw.-Gesch. und Natiirl. Syst. Europ. Thicrw., i,

p. 83.
1857. Lcucocyon Gray, Proc. Zool. Soc, London, p. 512.

Tape species. — Canis lagopus Linnteus.

Gcograpliieal distribution. — Arctic region of l)oth Old and New
"Worlds ; in Europe south to southern Norway and Sweden.

Characters. — Skull intermediate in general form between that
of Canis and Vulpes ; occipital depth about one-third condylo-
basal length ; interorbital region more elevated than in Vulpes
owing to greater inflation of the frontal sinuses ; postorbital
processes thin, flat or slightly concave above, with liead-like,
overhanging edges ; dorsal profile of forehead rising abruptly

ALOPEX 319

above rostrum as in Conin ; teeth moderately heavy and large,
the length of cariiassial and upper molars together contained
about 2j times in palatal length ; canines and incisors inter-
mediate between those of Cams and Vulpes (see fig. 65) ; external
form fox-like, but ear short and rounded, not conspicuously
overtopping the surrounding fur.

liemdrks. — Although in most respects intermediate between
Canis and Vulpes the Arctic foxes form such a natural and
circumscribed group that it seems desirable to set them apart as
a distinct genus.* Half a dozen species have been described, two
of which come within the scope of the present work.

KEY TO THE EUROPEAN SPECIES OP ALOPEX.

Condylobasal length of skull about 130 in males,

124 in females (Scandinavia and Finland)... A. lagopus, p. 319.
Condylobasal length of skull about 120 in males,

114. in females (Spitzbergen) A. sjntzbcrgmcnsis, p. 324.

ALOPEX LAGOPUS Linnaius.

1758. [Canis] lagopus Linnaeus, Syst. Nat., i, 10th ed., p. 40 (Lapland).
1816. V[;ulpes] arctica Oken, Lehrb. d. Naturgesch., iii, pt. 2, p. 1038

(Renaming of Canis lagopus).
1820. C[a7iis'] violpcs cxrulca Nilsson, Skand. Fauna, i, p. 88 (Lapland).
1827. ICanis lagopus] B argcntcus Billberg, Synopsis Faunae Scandinavia3,

p. 14 (Lapland).
1910. Vul2:ies lagopus Trouessart, Faune ]\Iamm. d'Europe, p. 96.

Ty^je locality. — Lapland.

Geo<jrapMcal distribtition. — Arctic portions of the mainland
of Europe and Asia ; in Europe south along the mountains of
Scandinavia to south-western Norway, and as an occasional
visitant as far as southern Sweden.

Dia(iuosis. — General characters as in the genus ; condylobasal
length of skull about 130 mm. in males, about 12 i mm. in
females.

External characters. — In general external charactei's Alopex
lagopus resembles Vulpes ruljX'S, though the muzzle is less
elongated, and the low, rounded ears (not extending to eye when
laid forward) impart a somewhat lui-fox-like appearance to the
head. Fur very dense, the underfur in summer about 12 mm.
deej) on back, nearly twice as deep and somewhat looser in
texture on sides (in winter longer throughout) ; longer hairs
rather sparse, not concealing underfur. Tail bushy, with
abuiulant underfur. Feet as in Cauis, but claws longer and
more .slender, and entire palm and sole covered with a dense
woolly growth of hair, 10-17 mm. deep in winter, shorter in

* See Collett, Norges Pattedyr, p. 275, 1911, for account of apparently
complete sterility of Arctic fox male with Red fox female, a fact which
indicates a fundamental physiological diflcrencc between the two animals.

320

CARNIVORA

suininer when it sometimes wears away sufficiently to expose
lialls of toes, and parts of pads.

Colour. — Summer pelage : ground colour of back, shoulders,
and outer side of legs drab, darkening to about prout-brown or

Fig. G3.
Alopcx laijoinia.

dark bister on feet, head, chin, interramial region and outer
surface of ears, the face thickly sprinkled with whitish hairs,
especially on cheeks and between eyes, the inteiramial region
tinged with slaty grey ; each of the longer hairs of back with

321

one cream-buff sub-terminal aiinulation, producing a noticeably-
speckled appearance throughout dark area ; flanks with a few
long, entirely butt' hairs ; sides of body and of neck light cream-
buff tinged with clay-colour, in striking contrast with dark areas,
the buff lateral area divided at shoulder by band about 60 mm.
wide where drab of back crosses to leg ; anteriorly the buff
lateral area extends to about level of ears where it abruptly gives
place to dark brown of head ; inner surface of ear light bufiy
grey ; underparts and inner surface of legs buffy greyish, slightly
contrasted with sides ; under surface of tail like sides of body,
upper surface essentially like back at base (where line of
demarcation is well defined), but becoming tinged with buffy
toward tip. Winter pelage : entirely white, tinged with yellowish
on throat, neck and face ; underfur and posterior surface of ear
light drabby grey. In the " blue " phase the entire animal is at
all seasons a bluish drab, usually washed with sepia on head and
feet, and sprinkled with pure white hairs on face, chin and throat.

Skull. — The skull is shorter and narrower than that of Vulpen
vulpes, but of nearly equal depth, a difference in form due in part
to the greater elevation of the interorbital region and in part to
the relatively greater depth of brain-case (depth equal to one-
third condylobasal length instead of noticeably less as in
V. milpes) ; muzzle less produced
than in the common fox and rela-
tively wider proximally ; zygomata
less abruptly spreading anteriorly,
so that the region of greatest
zygomatic breadth is noticeably at
glenoid level. In other respects
there is essential agreement with
the skull of V. vulpes. Anteorbital
foramen over space between 'pm^
and pm* ; auditory bulla relatively
as large as in the larger animal ;
depth of brain-case through bulla
equal to greatest breadth above
roots of zygomata ; postorbital pro-
cesses slightly less concave on
underside and somewhat more
flattened above ; mandible with
ramus relatively a little deeper
and more compressed than in the
common fox.

Teeth. — In general the teeth
closely resemble those of Vidpes
vulpes except for their slightly
smaller size. Incisors both above
and below with relatively wider crowns than in the common fox,
the cingulum of i^ and i? much more developed and forming a

Alopex lagopu.

322

CARNIVORA

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324 CARNIVORA

noticeable inner and outer secondary cusp (the former soon
wearing away in /') much as in Canis ; along posterior border of
crown the cingulum shows a strong tendency to develop irregular
tubercles, a condition not observed in Vulpes vulpes. Canines
and premolars with no special peculiarities, the form of the upper
carnassial quite as in V. vulpes. Lower carnassial with posterior
heel narrower than main portion of tooth but not otherwise
peculiar. First upper molar with metaconule relatively less
developed than in Vulpes vulpes.

Measurements. — For cranial measurements see Table, p. 322.

Specimens examined. — Ten, from the following localities : —
Norway: Roros, Trondhjem, 1; Tolgen, Hedemarken, 2; Dovre, 2;
Egersund, Stavanger, 1.

Sweden : No exact locality, 1 skull (U.S.N.M.).

Lapland : No exact locality, 3 skulls (B.M. and U.S.N.M.).

?. Roros, Trondhjem, Nor- Christiania Museum 95. 11. 14. 1.
way. (e).

skull. Egersund, Stavanger. K.H.Schaauning(c). 11. G. 3. 12.

2 9. Dovre. Christiania Museum 95. 11. 14. 2-3.

(E).

2 imm. Tolgen, Hedemarken. Christiania Museum 93. 3. 1. 1.

(e). 0. 5. 2. 1.

2 skuUs. Lapland. Wheelwright (c). 64.3.8.3-4.

ALOPEX SPITZBERGENENSIS Barrett-Hamilton and Bonhote.

1799. ? Canis fuliginosus Bechstein, Thomas Pennant's allgem. Uebersicht
d. vierfiiss. Thiere, i, p. 270 (Iceland).

1799. ? Canis grocnlandicus Bechstein, Thomas Pennant's allgem. Ueber-
sicht d. vierfiiss. Thiere, i, p. 270 (Greenland).

1898. Canis lagopus spitzhergcncnsis Barrett-Hamilton and Bonhote, Ann.
and Mag. Nat. Hist'., 7th ser., i, p. 287, April, 1898 (Spitzbergen).
T}'pe in British Museum.

1910. Vulpes lagopus spitzbcrgensis and ? V. lagopus fuliginosxis Trouessart,
Faime Mamm. d'Europe, p. 97.

Type locality. — Spitzbergen.

GeograpMcal distribution. — Spitzbergen ; also Iceland and
Greenland 1

Diagnosis. — Like Alopex lagopus but smaller, condylobasal
length of skull about 120 in males, about 114 in females.

Colour. — Type (summer pelage) : colour pattern well defined
and exactly as in A. lagopus, but dark areas wood-brown against
which the cream-bufi" annulations of longer hairs make no marked
contrast. Another skin, also in summer pelage, is a uniform
dark slaty drab throughout, the hind feet darker and more
brown ; .sides and underparts with a few long white hairs
(50 mm.) ; lijis with slight grizzling due to presence of short
white hairs.

Measurements. — For cranial and dental measurements see
Tables, pp. 322, 323.

VULPES

325

Specimens examined. — Six, all from Spitzbergen(B.M.and U.S.N.M.): —

6 skull. Spitzbergen. Stockholm Museum (e). 90. 8. 1. 2.

i, 9. Spitzbergen. Dr. J. W. Gregory (c & p). 96. 9. 23. 2-3.

(96. 9. 2.3. 3. Type of species.)
$, 9 skulls. Spitzbergen. Dr. J. W. Gregory (c & p). 96. 9. 23. 4-5.

Genus VULPES Oken.

1816. V[ulpcs] Oken, Lehrb. d. Naturgesch., iii, pt. 2, p. 1033, in full on
p. 1034 and in index, p. 1268 (Vulpcs communis Oken = Canis
vulpcs Linnifius).

1821. Vulpes Bowdich, Anal. Nat. Classif. Mamm., p. 40 (Canis viilpes).

1857. Vulpes Blasius, Siiugethiere Deutschlauds, p. 178 (Sub-genus of
Canis).

Type species. — Canis vulpes Linnajus.

Geographical distrihution. — Northern portion of the northern
hemisphere from about the hmit of tree growth south to Morocco,
India and Mexico ; in Europe west to Ireland.

Characters. — Skull slender and low (depth of brain-case less
than one-third condylobasal length) ; interorbital region nearly
flat, the frontal sinuses scarcely inflated, the postorbital proces.ses
thin, slightly concave above, their edges overhanging and bead-
like ; dorsal profile of forehead rising very slightly and gradually
above level of rostrum ; dental formula as in Canis ; teeth
relatively light and small, the length of upper carnassial and
molars together contained about 2| to 3 times in palatal length,
the general character of cheek-teeth somewhat more trenchant
than in Catiis, the canines slender and elongated, the point of
upper tooth extending to about level of lower margin of mandi-
bular ramus when jaws are closed (fig. 65).

MemarJcs. — As thus restricted the genus Vulpes contains about
thirty-five forms, all jjeculiar to the northern hemisphere. Five
of these occur in Europe.

KEY TO THE EUROPEAN FORMS OP VULPES.

Size small, hind foot in adult male about 125 mm.,
condylobasal length of skull in both sexes less

than 130 mm. (Sardinia and Corsica) T'. ichnus,T, p. 336.

Size large, hind foot in adult male 135 to 165 mm.,
condylobasal length of skull in adult male 136 to

165 mm., in adult female 127 to 155 ram V. vulpcs, p. 326.

Teeth larger and more robust, the premolars tend-
ing to be slightly spaced or in contact, their
secondary cusps well developed (Scandinavia)... V. v. vulpes, p. 330.
Teeth smaller and less robust, the premolars tend-
ing to be widely spaced, their secondary cusps
usually obsolete or absent.
YeUowish and reddish tints bright ; posterior
half of back seldom much frosted with whitish ;

tail never clear grey (Central Europe) r. v. crucigem, p. 331.

Yellowish and reddish tints pale and dull ; pos-
terior half of back usually much frosted with
whitish ; tail often clear grey (Iberian Penin-
sula) V. V. yilacea, p. 333.

326

CARNIVORA

FtG. 65.
Incisors and canines from finnt, of fVon'-v (a), Alopex (i), and Vulpes (c).

VULPES VULPES Linnjeus.

(Synonymy under subspecies.)

Ty2je loralHii. — TTpsala, Sweden.

GeograpMcul distrihufion. — Europe from the Arctic coast to
the Mediterranean, and from Ireland eastward into Asia.

Diar/nosis. — Size large: hind foot, in adult male 135 to 165
mm.; condylobasal length of skull in adult male 135 to 165 mm.,
in adult female 127 to 155 mm.

External characters. — Form more slender and legs relatively
shorter than in Canis ; muzzle long and pointed ; ear high,
pointed, rising conspicuously above surrounding fur ; tail long,

VULPES 327

thick anfl bushy, with abundant underfur ; longer hairs of back
normally concealing the underfur ; feet as in Cauls, but with
the soles hairj' between the pads, the pads themselves sometimes
furred.

Colour. — General colour a yellowish brown brighter and more
inclined toward reddish along median dorsal region and on face,
duller and more yellowish or greyish on sides of body to shoulder
and on sides of neck to base of ear, the flanks and usually the
sides sprinkled with white hairs which may produce a decided
effect of frosting ; underparts very variable, ranging from
whitish to slaty black, rarely almost concolor with sides ; feet
dusky or blackish ; ear tawny or buff at base and on inner
surface, the terminal half of outer side black or very dark brown
in strong contrast with surrounding parts ; upper lip dull white.
Blackish and greyish variations not uncommon, especially at the
north.

Skull. — General form of skull slender and somewhat flattened,
with widely spreading, nearly parallel zygomata. Dorsal profile
almost flat from nares to slightly beyond midfUe of nasal bones,
then rising at a slight angle (10° or less) to or a little beyond
bregma, behind which it slopes away by an evenly convex curve
(distorted in old individuals by the development of the sagittal
crest) to lambda, which lies a little above level of middle of orbit ;
ventral profile essentially straight. Brain-case distinctly broader
than high, its outline ovate when viewed from above, the
lambdoid and sagittal crests well developed, the latter extending
forward about to bregma, where it di^'ides, sending a branch to
form posterior border of each postorbital process. Occiput
obliquely truncate, so that condyles are not visible from above,
but region between lambda and foramen magnum slightly if at
all concave. Floor of brain-case with no special peculiarities ;
auditory bulhe moderately and evenly inflated, slightly flattened
antero-externally, meatal tube short but distinctly indicated,
especially its hinder wall. Interorbital region flattened, with
median longitudinal groove, the postorbital processes prominent,
flattened, triangular, much shorter along posterior edge than
along antero-external edge, the margin slightly raised so that
the upper surface is some\\hat concave, the under surface so
abruptly concave that the process is much less thick than in
Alojjex and Cani.s. Rostrum moderately long (the distance from
orbit to gnathion about etjual to that from postorbital process to
lambda), rather abruptly narrowed proximally, so that the sides
are nearly parallel through a noticeable portion of their extent
(occasionally the sides diverge from region of greatest narrowing
to bases of canines) ; nasal slender, narrowing gradually back-
ward and extending nearly to level of middle of orbit ; nasal
branch of premaxillary extending to about middle of nasal and
usually not in contact with frontal ; posterior extremity of
maxillary extending slightly behind that of nasal ; anteorbital

328

CAENIVORA

foramen over posterior root of pm? or over space between pw/^
and ptu*. Palate relatively narrower than in the European

Fia. CG.
Viilpes vuljies. x i.

species of Alopex and Canis, its median posterior termination
about at level of middle of last mola.r ; mesopterygoid fossa

329

I

about twice as long as wide, the sides gradually converging
posteriorly ; haiiuilars slender, straight ; incisive foramina about
four times as long as wide, the median septum usually wider
than the foramen, the posterior margin a little in front of
posterior margin of alveolus of canine. Mandible slender, but
with no special peculiarities of form.

Teeth. — In general form and structure the teeth do not differ
very widely from those of Canis lupus. The canines are, however,
relatively much longer and more slender (fig. 65), the incisors are
weaker and narrower, and the premolars show a more decided
tendency to develop secondary
cusps. Upper incisors slender,
not closely crowded, their
crowns relatively narrower than
in either Canis lupus or Alopex
liujopus, the secondary cusps
obsolete, early disappearing
with wear, the cingulum barely
indicated ; lower incisors with
crowns less simplified than in
the upper teeth, the usual lon-
gitudinal groove present on
posterior surface, and i^ with
well developed outer basal lobe.
Upper canine slender, its dia-
meter at alveolus about 7 mm.,
its height when unworn usually
about three times as great ;
lower canine with diameter
contained about 2;V times in
height. Premolars with no
special peculiarities, their crowns
relatively narrower than in
Canis lupus, and secondary cusps
tending to be more developed,
the latter character varying in
different geographical forms.
Upper carnassial with inner
lobe better developed than in
Canis, and bearing a distinct terete cusp, its position a little
more forward and outward than in the dogs, so that it appears
to lie in or nearly in the main axis of the tooth. Lower carnassial
with 2)Osterior lieel essentially as broad as main portion of tooth,
the cu.sps es.sentially as in Canis, but general aspect of tooth more
trenchant. Ujiper molars differing from those of Canis in the
relatively smaller size of paracone and metacone, these cusps
apjiearing to stand in from border of crown, leaving a noticeable
cingulum beyond them ; paraconule obsolete. Second and third
lower molars with no marked peculiarities.

FIG. 67.
Vulpes vulpes. Teeth.

330

CARNIVORA

RcmarJiS. — With the material at hand it has been impossible
to reach wholly satisfactory conclusions with regard to the
number of local forms represented by the common foxes of various
parts of Europe. The existence of three such races, one in the
Scandinavian Peninsula, a second in Central Europe, and a third
in the Iberian Peninsula, seems well established ; but the status
of the forms inhabiting Italy and Greece is still in doubt.

VuLPES vtiLPKs VULPES Linuieus.

1758. [Canis] vulpes Liiiuffius, Syst. Nat., i, 10th ed., p. 40 (Sweden).
1758. [C'auis] alopex Liniiieus, Syst. Nat., i, 10th ed., p. 40 (Sweden).
1798. C[anis] vulpus Thunberg, Beskrifning pA Svenske Djur, Mamm., p. 7

(Variant of vulpcs).
1816. Vl^ulpes] vulgaris Oken, Lehrb. d. Naturgesch., iii, pt. 2, p. 1034

(Renaming of vulpes).
1820. Canis nigro-argentcus Nilsson, Skand. Fauna, i, p. 91 (Lofoten

Islands, Norway).
1827. [Canis vulpus] y oiigrocaudatus Billberg, Synopsis Faunae Scandi-

navise, p. 12 (Uppland, Sweden).
1827. [Canis vul2ms] ( variegatus Billberg, Synopsis Faunas Scandinavise,

p. 1.3 (Uppland, Sweden).
1827. [Canis vulpus] i) Uncatus Billberg, Synopsis Faunae Scandinaviae,

p. 13 (Skane, Sweden).
1830. ? [Vulpes] communis Burnett, Quart. Jouru. Sei. Lit. Art, sxviii,

1829, p. 349 (Substitute for vulpes). Nomen nudum.
1898. Vulpes vulpes Thomas, The Zoologist, 4th ser., ii, p. 100, jMarch,

1898 (part).
1910. Vulpes vulpes Trouessart, Fauna Manun. d'Europe, p. 93 (part).

Type locality. — Upsala, Sweden.

Geographical distribution. — Scandinavian Peninsula.

Diagnosis. — Teeth larger and more robust than in the central
and southern races, the premolars tending to be slightly spaced
or in contact ; skull attaining maximum size for Eiu'opean
foxes.

Measurements. — For cranial and dental measurements see
Tables, pp. 334, 335.

Specimens examined. — Seven skulls from Sweden (for exact localities
see Table of cranial measurements), and eleven from Egersund, Stavanger,
Norway.

Remarks. — While I have seen no skins of this fox, the
distinctness of the race from those occurring in central and
southern Europe seems well established by the characters of
the teeth.

skull. Sweden. Wheelwright (c). 04. 3. 8. 2.

11 skulls. Egersund, Stavanger, K. H. Schaanning (c). 11. G. 3. 1-11.
Norway.

VULPES 331

VuLPEs VULPES CRUCiGERA Bechsteiii.

1789. [Canis] crucigera Bechsteio, Gemeinn. Naturgescb. Deutschlands,
I, p. 250 (Thiiringen, Germany).

1792. C[anis] Vulpes cilopex europaeus Kerr, Anim. Kingd., p. 142 (Bur-
gundy, France).

1797. Canis vulpes alba Borkhausen, Deutsche Fauna, i, p. 33 (Vogelsberg,
near Rudigshain, Hessen, Germany).

1797. Canis vulpes nigra Borkhausen, Deutsche Fauna, i, p. 33 (Hessen
and Thiiringen, Germany).

1801. Clanis] vl^idpes] lutea Bechsteiu, Gemeinn. Naturgesch. Deutsch-
lands, I, 2nd ed., p. G28 (Thiiringen, Germany).

1801. C[a7iis] v[ulpes] cinerca Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2nd ed., p. 628 (Thiiringen, Germany)..

1832. Canis melanogaster Bonaparte, Iconogr. Fauna Ital., i, fasc. 1
(Neighbourhood of Rome, Italy).

1841. Vulpes hyponielas Wagner, Schreber's Saugthiere, Suppl., ii, p. 405
(Oberbayern, Germany).

1857. Canis vulpes Blasius, Siiugethiere Deutschlands, p. 191.

1861. ? V[2dpes] vulgaris meridionalis Fitzinger, Wisseusch.-pop. Natur-
gesch. der Saugeth., i, p. 194 (Dalmatia).

1910. Vulpes vulpes (part) and V. vulpes melanogaster Trouessart, Fauue
Mamm. d'Europe, pp. 93-94.

Type locality. — Thiiringen, Germany.

Geographical distribution. — Central and .southern Europe from
Ireland eastward and from the coast of the Baltic to the
Pyrenees, Italy and Greece.

Diagnosis. — Maximum size I'ather less than in V. v. vulpes,
and teeth distinctly smaller, the premolars rather widely spaced
and seldom if ever in contact ; general colour a bright yellowish
or reddish brown, the posterior half of back not conspicuously
frosted with whitish, and tail never clear greyish.

Colour. — In seventeen skins the general colour ranges from
nearly cinnamon-rufous to a light ochraceous-rufous, the sides
of neck and region immediately behind shoulder lighter than
median dorsal area (^in extreme in.stances clear buff with a
decided rufous tinge) ; posterior half of back with evident
white frosting in some specimens, scarcely any in others, but
this character never so pronounced as in average Spanish skins ;
underparts dull slaty overlaid with white, the slaty nearly always
predominating, except on throat, and not infrequently giving the
effect of an almost blackish tinge throughout, this apparently
not in the least dependent on regional or local climatic con-
ditions. In a flat skin from Cephalonia, Greece, the characteristic
slaty and white is confined to the throat and chin, all the rest
of the ventral region being a dull tawny-ochraceous like sides.
Two specimens from Tatoi, near Athens, taken in July, have
shed all the longer hairs of the back, leaving only the velvety
underfur. This is of the usual colour, a dull umber brown, in
one specimen with a slaty cast.

Sltill and teeth. — The skull and teeth do not attain so srreat

332

CARNIVORA

size as in true Vuljies ruJpes, and the small premolars are seldom
in contact, the spaces between them usually conspicuous.

Measurements. — Adult female from Cappagh House, Water-
ford, Ireland : head and body, 613 ; tail, 340 ; hind foot, 146 ;
ear from meatus, 90. Adult male and female from Tunbridge
Wells, Sussex, England : head and body, 690 and 630 ; tail,
343 and 370; hind foot, 160 and 141 ; ear from meatus, 95 and
82. Adult male from Ingelheim, Rheinhessen, Germany : head
and body, 578 ; tail, 440 ; hind foot, IGO ; ear from meatus, 98.
Adult male and female from Haute-Garonne, France : head and
body, 610 and 610; tail, 370 and 350 ; hind foot, 150 and 135 ;
ear from meatus, 94 and 89. Two adult males from Porlezza,
Como, Italy (Ghidini) : hind foot, 150 and 148-6. Adult male
from Borghetto S. Spirito, Italy : head and body, 745 ; tail, 380 ;
hind foot, 157. Adult male from Zinnigas, Siliqua, Sardinia
(measured from mounted specimen, Genoa) : head and body, 700 ;
tail, 300; hind foot, 144± ; ear from meatus, 83.

Specimens examined. — Fifty, from the following localities : —

Ireland : Kilmanock, Wexford, 2; Cappagh House, Waterford, 1.

Scotland : Ben Nie, Sutherland, 1 ; Inversanda, Ardgour, 1.

England : Northumberland, 1 ; Thame, Oxford, 2 ; Hassocks, Sussex, 1 ;
Tunbridge Wells, Sussex, 3 ; Ditchling, Sussex, 1 ; Mayfield, Sussex, 1
(Grant).

France : As-les-Thermes, Ari^ge, 3 ; Caterille, Haute-Garonne, 1 ; Pic
Sessire, Haute-Garonne, 1 ; St. Aventin, Haute-Garonne, 1.

Germany : Ingelheim, Rheinhessen, 1 ; Nuremberg, Bavaria, 1
(U.S.N.]\I.) ; Grossgraben, Silesia, 1 (Breslau) ; Rieseugiberge, Silesia, 1
(Breslau) ; southern Germany, 3.

Austria-Hungary : Trentino, 1 (Genoa) ; Gazza, Treutino, 1 (Genoa) ;
Vigolo Vattaro, Trentino, 1 (Genoa).

Switzerland: Geneva, 1 (Ghidini); Valais, 1 (Ghidini).

Italy : Porlezza, Como, 4 (Ghidini) ; Garbagna, Piedmont, 1 (Genoa) ;
Borghetto S. Spirito, 1 (Genoa) ; Vargo, Liguria, 1 (Genoa) ; Torriglia,
Liguria, 1 (Genoa) ; Cornigliano, Liguria, 2 (Genoa) ; near Genoa, 1 ;
Molasana, 1 (Genoa) ; Tuscany, 1 ; Pisa, 1.

Sardinia: Zinnigas, Siliqua, 1 (Genoa).

Greece : Cephalonia, 1 ; Tatoi, near Athens, 2.

Remarlcs. — The fox of Italy appears to be the same as that of
Central Europe, though further material from the southern
portion of the peninsula may show that it should be distinguished.
The three specimens from Greece are in such unsatisfactory
condition of pelage that their status cannot be determined with
any degree of certainty. If they represent a peculiar local race
this should probably take the name vteridioiiaUs Fitzinger.

9. Ben Nie, Sutherlandshire, E. R. Alston (p). 79. 9. 2.5. 80.

Scotland. {H. Brown.)

6. Inversanda, Ardgour. H. Leigh (c & p). 1. 16. 12. 1.

2. Kilmanock, Wexford, G. Barrett-Hamilton 6. 6. 20. 1.

Ireland. (c & p). 9. 12. 15. 4.

?. Cappagh House, Waterford. R. J. Ussher (c & p). 96. 12. 28. 1.

1. Northumberland, England. Rev. H. H. Slater 0.2.24.5.

(c & p).

vuLPES 333

2 6. Thame, Oxfordshire. Hon. N. C. Roths- 0. 10. 31. 1-2.

child (p).
2 (J. Ditchling, Sussex. Guy Dollman /9. 11. 3. 1.

(c & p). \10. 9. 13. 1.

3. Tunbridge Wells, Sussex. G. H. B. Grant (c). 1. 2. 1.5. 1-3.

2(5,9. Ax - las - Thermes, Arifege, V. Builles (c & p). 8.3.27.6-7.
Franco. 8. 3. 27. 14.

9. Caterille, Haute-Garonne, O. Thomas (p). 8. 7. 15. 2.

900 m. [A. Robert.)
9. PicSessiro,Haute-Garonno. 0. Thomas (p). 8. 7. 15. 3.

[A. Robert.)
6. St. Aventin, Haute-Ga- O. Thomas (p). 8. 7. 15. 4.

ronne, 900 m. (A. Robert.)
i. Ingelheim, Rheinhessen, C. Hilgert (c). 8. 11. 2. 15.

Germany.
3 skulls. South Germany. Dr. A. Giinther (c). 59. 9. 6. 83, 88.

175. k.
S. Genoa, Liguria, Italy. MarquisG. Doria (p). 88. 12. 1. 3.

skull. Pisa. Zool. Soc. Mus. 68. 5. 4. 126.

lyg. S. Italy. (Prof. Savi.) Zool. Soc. Mus. 55.12.24.240.

9. Tuscany. Purchased (Dr. 45. 7. 22. 15.

RiippeU).
6, 9. Tatoi, Athens, Greece. Hon. N. C. Roths- 8. 10. 2. 22-23.
(C. Mottaz.) child (p).

VuLPES VULPES siLACEA Miller.

1907. Vulpcs vidpes silaceus Miller, Ann. and ^Mag. Nat. Hist., 7th ser.,

XX, p. 393, November, 1907. Type in British Museum.
1910. Vulpes vulpes silaceus Trouessart, Faune Mamm. d'Europe, p. 45.

Type locality. — Near Silos, Province of Buri^os, Spain.

Geographical distribution — Iberian Peninsula.

Characters. — Size about equal to that of V. v. vulpes, but
teeth noticeably less enlarged, the wider spacing of the premolars
especially evident ; general colour a dull buff without bright
yellowish or reddish tints, the posterior half of back conspicuously
frosted with whitish (except in abraded pelage) the tail often a
clear greyish.

Measurements. — Type (adult male) : head and body, 750 ;
tail, 370; hind foot, 150. Adult female from near the type
locality: head and body, 670; tail, 330; hind foot, 125. Old
male from Elche, Alicante : head and body, 770 ; tail, 480 ;
hind foot, 160; ear from meatus, 113.

Specimens examined. — Eighteen, from the following localities in
Spain : Olot, Gerona, 1 (probably not typical) ; vicinity of Silos, Burgos, 5 ;
near Burgos, 1 ; Arrechavaleta, Vitoria, 1 ; Torres del Alio, Coruna, 1 ;
Madrid, 1 ; near Seville, 3 ; Goto Dofiana, Huelva, 3 (B.M. and U.S.N.M.) ;
Elche, Alicante, 1 ; Barracas, Gastellon, 1.

Bemarks. — The Spanish fox is well characterized by its light
colour, large size and rather small, widely-spaced premolars as
compared with those of the Scandinavian form.

2 6. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 46-47.

(8. 8. 4. 46. Type of subspecies.)
(J, 2 9, 1. Burgos. Rev. Saturio Gon- 8.7.7.12-15.

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9. Arrechavaleta, Vitoria. 0. Thomas (p). S. 2. 9. 49.

(y. Gon.'alez.)

6. Torres del Alio, Comua. Dr. Y. L. Seoane 94. 5. 29. 1.

(c & p).

6. :Madrid. M. de la Escalera (c). S. 9. 24. 7.

6, 9. SeviUe. {A. Ruiz.) Lord Lilford (p). 95. 9. 4. 7-8.

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VULPES ICHNUS.E Miller.

1907. Yulpes icliniisx Miller. Aun. and Mag. Xat. Hist., 7th ser., xx, p. 391,

November, 1907. Type in British Museum.
1910. Tidpcs vuJpcs ichniisx Trouessart, Faune Mamm. d'Europe, p. 94.

Type locality. — Sarrabus, Sardinia.

Geographical distribution. — Sardinia and Corsica.

Diagnosis. — Smaller than any of the races of Vidpes vulpes ;
both hind foot and condylobasal length of skull in adult male
less than 130 mm., ear from crown 60 to 70 mm.

Colour. — Face and head dark rufous becoming Hghter and
more dull on base of ears and on neck, and fading to ockraceous-
rufous on shoulders and back ; sides of neck, outer surface of
upper arm and region just behind axilla still lighter, a ta^^Tiy
butf, hairs of underfur on back drab grey at base, tawny clay
colour at tip ; longer hairs of head, sides, and back (behind
shoulders) much speckled by huffy white subtermmal areas about
5 mm. in length, the extreme tips reddish : feet and legs
ochraceous-rufous, slightly clouded with blackish and a little
speckled with bufiy white : tail like back above, the tawny
gradually fading out through a bufiy grey to the whitish buff"
tip, the longer hairs except in pencil black tipped (30-40 mm.) ;
underparts to front legs huffy wliitish tinged with hair-brown,
this nearly clear on middle of throat ; rest of underparts a
mixture of hair-brown and dull tawny, the latter predominating
along sides.

Measurements. — Type (adult male) : hind foot, 123 ; ear from
meatus, 70. Adult male and female from the type locality :
head and body, 640 and 590 ; tail, 280 and 290 : hind foot,
127 and 125 : ear from meatus, 74 and 71. Adult female from
Siliqua : head and body, 600 : tail, 350 ; hind foot, 127 : ear, 74.
Adult from near St. Florent, Corsica : hind foot, 123 : ear from
meatus, 73.

Specimens examined. — Eight, from the following localities : —
Coksica: Grotto Campu Consule, near St. Florent, N.W. Corsica, 1

(Major).

Saedesia : No exact locality, 1 (Turin) ; Lanusei, 1 ; Sarrabus, 3 (B.M.

and Genoa) ; Zinnigas Siliqua, 2.

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cientzal ^Seaodinavia.

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- J" the Tiiwimimm i^j^Bam.

342

CARNIVORA

Geograplrical distribution. — Northern temperate portions of
Old World ; in Europe from central Scandinavia to the
Mediterranean, and from Ireland eastward.

Characters. — Skull narrow and high (depth of brain-case
much more than half mastoid breadth), the zygomatic arches not
widely expanded, and postorbital region not specially narrowed
(distance from point of greatest narrowing to zygoma less than
breadth of postorbital constriction) ; rostrum elongate, the
distance from orbit to gnathion about three-fourths length of
brain-case ; auditory bulla elevated along inner margin, but else-
where somewhat flattened, the meatal tube distinct ; paroccipital
process robust, standing out conspicuously behind bulla ; dental
formula : i ||, c t], jmi ti,* m ^^ = 38 ; teeth not specially enlarged
as compared with width of rostrum and palate (greatest trans
verse diameter of upper carnassial equal to a little more than
one-third distance between carnassials) ; three small premolars
(one upper and two lower) capable of trenchant action ; upper
carnassial with crown triangular in outline, the cutting portion
consisting of a single large antei'ior cusp with its slightly developed
anterior commissure and more distinct posterior commissure, the
crushing portion represented by the obliquely sloj^ing inner base
of this cusp and ridge ; upper molar rhombic in outline, its crown
area about three times that of carnassial, its greatest diameter
in axis of tooth- row, the four primitive cusps present ; lower
carnassial with the three anterior cusps well developed, sub-equal,
the posterior heel decidedly larger than anterior triangle ;
external form heavy, the head pointed, the ears short but plainly
visible, the body thick, the tail short, not muscular ; feet sub-
plantigrade, the toes with long fossorial claws and without webs ;
fur coarse and loose.

Reiiiarlcs. — The genus Meles contains about half a dozen
named forms, the status of several' of which is at present not
clearly understood. Two species occur in Europe.

KEY TO THE EUROPEAN FORMS OF MELES.

Maxillary tooth-row, exclusive of incisors, about
35 mm. ; auditory bullae rather strongly in-
flated, the inner Isorder not ridge-like (Crete) M. arcalus, p. 352.
Maxillary tooth-row, exclusive of incisors, about
40 mm. ; auditory bullae slightly inflated, the
inner border ridge-like (distribution general) M. mcles, p. 343.
General colour moderately jpale ; teeth averaging
' smaller, less frequently attaining maximum

size (Central and southern Europe) M. m. meles, p. 348.

General colour slightly paler ; teeth averaging
larger, and more frequently attaining maxi-
mum size (Iberian Peninsula) M. m. marianensh! , p. 352.

* In adults usually '-p-" owing to the early disappearance of the small

343

MELES MELES Linnjfius.
(Synonymy under subspecies.)

Type locality. — Upsala, Sweden.

Gi'cxjrapMcal distribution. — I^urope, west to Ireland, south to
the Mediterranean and north to central Scandinavia. Eastei'n
limits of range not known.

Diagnosis. — Size large (upper length of skull in adult more
than 120 mm.; maxillary tooth-row, exclusive of incisors, about
40 mm. or more ; hind foot about 90 to 110 mm.) ; auditory bulla
moderately inflated, the highest region close to inner margin and
forming an evident longitudinal ridge, the region between ridge
and meatus noticeably concave ; teeth large, the two lower molars
together 22 mm. or more in length ; postero-external border of
upper molar usually convex, though occasionally straight or
slightly concave.

External characters. — ^Pelage coarse and loose, practically
without underfur, the hairs at middle of back about 60 mm. in
length, those on sides longer ; underparts scantily haired, the
skin usually visible ; palm bare, usually with a slight bristly
pubescence at middle in area between tubercles ; a large
tubercular mass at base of digits, convex in front, concave behind,
wider on outer border than on inner border, showing no tendency
to trilobation ; a roundish tubercle about half as large at postero-
external border of palm, separated from anterior mass by a wide
space ; a small, ill-delined pad at base of thumb ; sole densely
haired from heel to a little beyond middle, then completely naked ;
plantar tubercles essentially like those on palm except that
small pad at base of hallux is absent, and the two large masses
tend to coalesce, owing to absence of the wide intervening space ;
surface of pads on both palms and sole finely rugose, this
especially noticeable in dried specimens ; muzzle pad entirely
naked, but separated from upper lip by a narrow hairy band.
Mammte : a 2-2, * 1-1 = 6.

Colour. — Back and sides a coarse grizzle of black and buffy
white, the black usually predominating on back, the lighter
colour on sides. Throat, median ventral area and all four legs
and feet black or blackish ; face, chin and entire neck (except
underside) clear whitish except for a broad dark brown or black
band beginning on each side about 15 mm. behind nostril pad
and extending back, including eye and ear, to middle of neck,
where it fades in.sensibly into colour of back ; width of white
median area on face usually greater than that of the dark lateral
stripe, and about ecjual to that of the light area between lateral
stripe and dark ventral area : ear black, its anterior border
white in strong contrast ; eye usually a little below middle of
dark baud ; tail like back at base, soon fading to soiled white.

Skull. — Except for the greatly (.leveloped sagittal crest, the

344 CARNIVORA

height of which in old individuals is equal to nearly half inter-
orbital breadth, the skull of Meles meles is rather smooth and
without marked angularity. Depth of brain-case (without crest)
and depth through posterior portion of orbit nearly equal, and
about two-thirds mastoid breadth ; depth at front of nasal about
two-thirds that at orbit. Dorsal profile (without crest) nearly
flat from middle of brain-case to interorbital region, convex over
posterior half of brain-case ; rostrum sloping downward at angle
of about 30° with surface of brain-case. When crest is fully

FIG. 08.
Meles melex.

developed it reduces this angle to 20° or less. It is slightly
convex anteriorly, rather abruptly convex posteriorly, the hinder-
most portion slightly overhanging. Postorlntal piocesses short
but well developed, sharply outlining the small orbit (greatest
diameter of orbit slightly less than half that of interorbital
region) which, though widely open posteriorly, is more than half
encircled with bone. Rostrum moderately long, the distance
from gnathion to front of zygoma about equal to width of palate
including molars, and continued about 3,'; times in condylobasal
length of skull. Zygomata widel}' spreading posteriorly, narrow

MELES 346

anteriorly, the widest portion opposite front of glenoid fossa, the
arch Aery slightly bowed upward. Anteorbital foramina rather
large, sub-circular in outline, their greatest diameter about half
that of orbit. Ventral prolile of skull slightly concave through-
out. Palate moderately wide, the distance between molars
contained about 2 J times in distance from gnathion to level of
posterior edge of molar. Incisive foramina small, at level of
space between canine and P, elongate pyriform in outline, the
minute median foramen a little behind middle. Posterior exten-
sion of palate about equal to distance between molars, and slightly
more than half that from molar to hamidar. Mesopterygoid

Fig. 69.
Meles meles. x i.

space short and wide, its length only a little more than distance
between tips of hamulars. Auditory bullte slightly inflated,
irregularly triangular in outline, the meatus lying in angle formed
by the large, forward-projecting mastoid process. The surface is
irregular, with evident ridge near inner margin along crest of
most highly inflated region, this ridge often terminating anteriorly
in a bluntly pointed projection. Length of flattened portion
extending inward from meatus about equal to transverse diameter
of inflated portion of bulla, but the two regions not sharply
defined ; least distance between bulla> about \}, times diameter of
inflated portion ; paroccipital process short, triangular (not ridge-
like), its extreme base applied to posterior border of bulla.

346

CARNIVORA

Mandible heavily built, the I'amus nearly straight, the lower
border with the usual upward curve posteriorly, its depth at
middle about one-third that through coronoid process ; angular

riQ. 70.

Meles meles. x J.

process short, thick, and ill-defined, lying close to base of articular
process ; coronoid process squarely truncate above, its height
above articular process about equal to width at articular level.

Teeth. — The teeth are moderately large relatively to size of
skull, the incisors and canines rather short, the crown area of the
upper molar fully double that of any of the other maxillary teeth,
this last peculiarity unique among the European Mitstelidse. Upper
incisors robust, the crowns somewhat higher than wide when
viewed from in front, the teeth closely crowded in a slightly
convex row, the longitudinal diameter of crown a little greater
than transverse diameter ; i^ and r sub-equal, the latter slightly
the larger, their anterior surface convex with two faint longi-
tudinal grooves soon disappearing with wear, the cutting edge
entire, the posterior surface concave with narrow but distinct
heel ; P with crown area about double that of P and nearly half

347

that of canine, its crown higher than in the other incisors,
owing to lower insertion of root, but extremity of cutting edge
not extending beyond level of smaller teeth ; laterally the cutting
edge is continued down outer-posterior margin of shaft where it
functions against lower canine. Lower incisors not so large as
upper, forming a continuous, nearly straight row between
cauiiaes, the shafts straight, sloping obliquely forward, the root
of 1.2 implanted behind the others ; cutting edge irregularly
2-lobed, that of i^ longer than the others. Canines rather short
and weak, the shaft simple, that of upper tooth nearly straight,
that of lower tooth strongly curved backward, the enamel surface
of both essentially smooth, though that of lower canine shows
indications of a rudimentary cingulum in front. Anterior
premolar both above and below a minute or spicular tooth closely
crowded between canine and
second premolar, pm^ early
deciduous, pvii usually more
persistent. Second premolars
similar to each other, the crown
area about equal to that of P,
the outline irregularly oval when
viewed from above, the cusp
about as high as long, its apex
sHghtly in front of middle of
crown ; rootof each tooth single,
that of pm.T, showing a tendency
to become divided longitudi-
nally. Other small premolars
(pm^, pvi^ and j^^'^i) alike in form,
the crowns laterally compressed,
triangular when viewed from
the side, with apex slightly in
front of middle, jj)«^ and pm^
nearly alike in size, prii^ some-
what longer ; cingulum very slightly developed, not forming
secondary cusps. Upper carnassial triangular in outline, the
posterointernal border longest ; outer, trenchant portion of teeth
consisting of a main anterior cusp resembling ^jjh^ in size and form,
joined by a nearly horizontal commissure to a moderately high
posterior cusp ; cingulum low but bearing three distinct secondary
cusps, one at anterior base of main cusp, one (sometimes obsolete)
near middle of antero-internal border, and the third and largest
at middle of posterointernal border. Lower carnassial with
anterior triangle well developed, the three cusps sub-equal in
height, the metaconid subterete, the protoconid more compres.sed
than paraconid, its commissure slightly longer than that of the
anterior cusp ; crushing portion of crown longer and wider than
anterior triangle, its inner portion occupied by a large basin-like
concavity, its border with two large cusps on outer edge (of

Fig. 71.
Meles meles. Teeth.

348

CARNIVORA

which the anterior is the larger), a large cusp similar to postero-
external cusp at middle of inner margin ; posterior margin
occupied by three or four minute cusps soon disappearing with
wear, and a similar minute cusp sometimes in angle behind
protoconid and nietaconid. Second lower molar flat, terete, its
area about half that of crushing portion of carnassial ; middle of
crown with basin-like concavity ; margin with small but evident
antero-external and postero-external cusps, and a smaller
elevation at middle of inner edge. Upper molar rhomboidal
in outline, the inner and outer margins parallel, the former
nearly li times as long as latter; outer border occupied by two
rather large, subterete, conical cusps, the anterior (pai'acone)
slightly larger than posterior (metacone), so that greatest trans-
verse diameter of crown is slightly behind anterior border ;
remainder of crown a shallow basin-like concavity with rugose
surface and raised, irregularly nodulate margin, the middle of
concavity crossed by a ridge, convex internally, extending from
anterior base of paracone to posterior base of metacone and
bearing from three to five small cusps soon obscured by wear,
the three anterior separated from two posterior by a deep angle ;
the anterior group probably representing the protocone, the
posterior group the hypocone.

liemarJcs. — Two moderately well differentiated local races are
represented by the material that I have examined, one confined
to the Iberian Peninsula, the other general in distribution.
British specimens do not differ appreciably from Swedish
examples of the typical form.

Meles meles meles Linnseus.

1758. [TJrsus'] meles Linnaeus, Syst. Nat., i, 10th ed., ]}. 48 (Sweden).
1785. [MelesJ taxiis Boddaert, Eleuchus Animalium, i, p. 80 (Europe).
1789. [Urstts meles] /3 alba Gmelin, Syst. Nat., i, 13th ed., p. 102.
1789. [Ursus meles] y maculata Gmelin, Syst. Nat., i, 13th ed., p. 102.
1808. Taxus vulgaris Tiedemann, Zoologie, i, p. 376 (Renaming of Ursus

meles).
1816. Meles europn'us Desmarest, Nouv. Diet. d'Hist. Nat., iii, p. 465

(Renaming of Ursus meles).
1822. Taxus meles F. Cuvier, Hist. Nat. des ^Nlamm., iii, fasc. 36, January,

1822.
1827. [Meles] communis Billberg, Synopsis Faunae Scandinavise, p. 16

(Renaming of Ursus meles).

1827. [Meles conwiunis] & caninus Billberg, Synopsis Faunae Scandiuavise,

p. 17 (Scandinavia).
1857. Meles taxus Blasius, Saugethiere Deutschlands, p. 204.
1894. Meles meles Dahl, Die Heimat, iv, p. 125, June, 1894.

1899. M{eles] m{eles] typicus Barrett-Hamilton, Ann. and Mag. Nat. Hist.,
7th ser., iv, p. 384, November, 1899.

MELES 349

1906. Meles meles britannicus Satunin, Mittoilungen des Kaukasi.schen
Museums, ii, p. 115 (Based on the cranial measurements of
xf r'4 specimens recorded by Barrett-Hamilton, Ann. and Mac.

iQin ,.f*'-H>stv Tthser., IV p.384). Co-types in British Museum.

1910. Meles meles and M. meles britannicus Trouessart, Faune Mamm
d'Europe, p. 69.

Type locality. — Upsala, Sweden.

Geographical distribution.—Centvul and southern Europe from
southern Sweden to the Pyrenees and Italy, and from Ireland
eastward ; eastern limit of range not known.

Characters.~S>knll and teeth moderately large, the teeth
rarely attaining maximum size for the species ; colour moderately
light, the sides not often conspicuously whitish.

3Ieasuremcnts.— Adult male and female from Woodpark
Ga way, Ireland : head and body, 686 and 618; tail, 153 and
loO; hmd foot, 108 and 102; ear from meatus, 48 and 45
Adult female from Boucoune, Gers, France : head and body 670 •
tail, 170 ; hind foot, 100 ; ear from meatus, 45. Adult female
from Ingelheim, Rheinhessen, Germany : head and body 672 •
tail, 180; hind foot, 104; ear from meatus, .50. For cranial
measurements see Table, p. 350.

Specimens cxnmi7ied.-Thirty-&ve, from the following localities •-
Ireland : Woodpark, Galway, 2. '

England: Burnley, Lancashire, 1; Ross, Hereford, 1; Kentchurch
Herefordshire, 4; Tetworth, Oxfordshire, 1 ; Ibstone, Buckingham hire 1-
NoRw;v''T;'= B'?^°»' Devonshire; Cornwall 1; noexactloca ty,4:
^orway: Holme, Mandal, 7 ; Egersund, Stavanger 2
bWEDEN : Middle Sweden, no exact locality, 1 (Stockholm) ■ TAkern

Phaxce : Poret de Bouconne, Gers, 1
Bavarlafr"'' Ingelheim, Rheinhessen, 1; Strass, near Burgheim,
Italy : Liguria, 1 (Genoa).

6, 9. Woodpark, Galway, Ire- R. p. Hibbert (p) 95 6 ^ i
^^''^- , 95. 12. 3. 1.

o „„ , -- , .\(^o-typesoiM.7n.britannicus,^a.t\xmn.)

V. Burnley, Lancashire, Hon. N. C. Roths- 0 10 31 4

England. child (p). ■ ■ • •

i. Ross, Herefordshire. Hon. N. C. Roths- 0. 10. 31. 3.

o o ■ i T^ , child (p).

■»■, 3 ]uv. St. Kentchurch, Hereford- W. E de Winton
shire. (c & p).

i. Tetworth, Oxfordshire. Hon. H. Parker

6. Ibstone, Buckingham- A. H. Cocks (c & p) 4 1 25 1

shire. ^ ' • • . .

i. Horsham, Sussex. E. C. Hawes (c & p) 7 4 13 1

w it'- ^ ^ictou, Devonshire. J. C.Stagdon(c& p). 97.2 23 1

Sr^ S°T'^11- LordLilford(P). ^ 89.10.28.1.

skull. Sweden. Purchased (Wheel- 64. 3. 8. 1.

If 00 Wright).

4 <5, i 9. Holme, Mandal, Norway. R. J. Cuninghame 8. 8 9 7-12

(c & p) '

2 skulls. Egersund, Stavanger. K. H. Schaanuing /ll. 6. 3. 13

(c)' (11.16.23.2.

350

CAENIVORA

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352 - CARNIVORA

?. Foret de Bouconne, Gers, O. Thomas (p). 8. 7. 15. 5.

260 m. France. (A.

Robert.)
9. Ingelheim, Rheinhessen, C. Hilgert (c). 8. 11. 2. 16.

Germauv.

Meles meles maeianensis Graells.

1897. [Meles taxus'] var. marianensis Graells, Mem. Real Acad. Cien.,

Madrid, xvii, p. 170 (Central Spain).
1899. Meles meles mediterraneus Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7tli ser., iv, p. 384, November, 1899 (Seville, Spain). Type

in British Museum.
1910. Meles meles mecliterraneus Trouessart, Faune Mamm. d'Europe, p. 70.

Tyjje locality. — Central Spain.

Geographical distribution. — Iberian Peninsula.

Diagnosis. — Skull and teeth large, the teeth frequently attain-
ing maximum size for species ; colour light, the sides often
conspicuously whitish, especially in region bordering dark ventral
area.

Measurements. — For cranial and dental measurements see
Table, p. 351.

Specimens examined. — Seven, from the following localities in Spain ;
Arrechavaleta, Vitoria, 1 ; Quintanar de la Sierra, Burgos, 2 ; near
Seville, 2 ; Goto Doiiana, Huelva, 2.

Benmrks. — Although not very strikingly differentiated, the
large-toothed, pallid, Iberian badger seems worthy of recognition
as a geographical race.

6. Arrechavaleta, Vitoria, N. Gonzalez (c). 8.7.7.18.

Spain.
6, 9. Quintanar, Burgos. Rev. Saturio Gou- 8. 7. 7. 19-20.

c?, ?. Seville. (A. Ruiz.) Lord Lilford (p). 95. 3. 3. 7-8.-

(95. 3. 3. 7. Type oi mediterraneus Barrett-Hamilton.)
<?, 9. Goto DoSana, Huelva. A. Chapman (p). 8. 3. 8. 6-7.

MELES ARCALUS Miller.

1899. Meles meles mediterraneus Barrett-Hamilton, Ann. and ^Nlag. Nat.
Hist., 7th ser., iv, p. 131, November, 1899 (part).

1906. Meles meles mediterraiieus Bate, Proc. Zool. Soc, London, 1905, ii,

p. 318, April 5, 1906.

1907. Meles arcalus Miller, Ann. and Mag. Nat. Hist., 7th ser., xx, p. 394,

November, 1907. Type in British Museum.
1910. Meles arcalus Trouessart, Faune Mamm. d'Europe, p. 70.

Tj/jje locality. — Lassethe Plain, Crete.
Geographical distribution. — Island of Crete.
Characters.— Size small (upper length of skull in adult less than
110 mm. ; maxillary tooth-row, exclusive of incisors, about 35 mm. ;

353

DENTAL MEASUREMENTS OE MELES MELES AND M. ARC ALUS.

Locality.

Number.

Sex.

Upper molar.

Lower molar
(first).

1 M. meles meles.

Sweden : no exact locality

64. 3. 8. 1

9

13-0 X 11-4

15-4 X 7

0

Wegeliu ....

Stockholm

6 ?

15-0 X 13-0

16-2 X 7

8

Skane ....

i '!

15-2 X 13-0

18-0 X 8

2

Ulriksdal ....

,,

<5juv.

13-4 X 11-6

15-8 X 7

S

Sm&land ....

V ?

14-2 X 11-8

16-2 X 7

4

Orebro ....

i iuv.

14-2 X -il-S.

15-8 X 7

0

Takern ....

6 ■>

13-G X 11-2

15-6 X 7

2

Norway : Holme, Mandal

152622

6

li-6 X 10-4

14-0 X 6

4

»» »»

8. 8. 9. 9

6

12-8 X 11-8

14-G X 7

0

8. 8. 9. 10

6

12-6 X 11-2

14-6 X 7

0

M '*

8. 8. 9. 11

9

11-2 X 10-8

14-0 X 6

8

8. 8. 9. 12

9

12-2 X 10-4

14-0 X 7

0

Egersund, Stavanger

11. G. 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

<5?

13-4 X 12-0

16-4 X 8

0

., ,, , .

21]f

9?

14-0 X 12-4

16-2 X 8-2

„ . .

—

14-2 X 12-2
13-G X 11-8

—

Burnley, Lancashire

0. 10. 31. 4

9

14-4 X 12-2

16-2 X 8-2

Cornwall

—

15-0 X 12-2

18-0 X 8-2

Ireland : Woodpark, Co. Galway

95. 12. 3. 1

15-6 X 12-8

17-0 X 7-8

,, ....

95. 6. 2. 1

14-8 X 12-2

16-0 X 7-8

Denmark: Zealand.

Andersen

6

14-0 X 13-2

17-0 X 7-8

Copenhagen*

6

15-0 X 12-6

18-5 X 8-2

,,

6

15-0 X 13-7

18-3 X 8-3

,, .

,,

6

15-5 X 13-6

18-0 X 9-0

,, .

9

14-6 X 12-5

17-5 X 7-7

various localities, i ^laxi- 1
13 specimens : \ mum j

16-3 X 13-6

_

) Mini-»
ImumJ

13-0 X 12-0

various localities, (Maxi-i
17 specimens : \ mum 1

18-7 X 8-5

1 Mini- 1
" ^mumf

15-0 X 7-0

[Franco: Bouconne, Gers.

8. 7. 15. 5

9

14-0 X 12-0

lG-2 X 7-2

Germany: Ingelheim, Rheinhessen .

8. 11. 2. 16

9

14-8 X 12-8

l'^-2 X 7-8

Italy : Borghctto S. Spirito, Liguria

33 Genoa

9

13-G X 110

15-6 X 7-2

M. meles marianensis.

Spain : Quintanar, Burgos

8. 7. 7. 19

6

15-2 X 12-6

17-4 X 8-G

; Seville ....

95. 3. 3. 7t

6

16-0 X 12-0

16-6 X 7-8

,,

95. 3. 3. 8

9

14-6 X 12-6

16-8 X 8-2

Goto Dofiana, Huelva .

8. 3. 8. G

6

16-4 X 13-2

17-8 X 8-0

....

8. 3. 8. 7

9

14-8 X 12-0

17-6 X 8-2

1 M. arcalus.

|Orete

5. 12. 2. 17t

9 juv.

13-6 X 11-2

15-4 X 7-2

5. 12. 2. 38

ad.

13-8 X 11-G

• -Measured l.y H. Wlnge.

f Type of iiiediten-aiieiii' Banett-Hamilton.

t Type.

354

CARNIVORA

hind foot about 25 mm.) ; auditory bulla strongly inflated, the
highest region near middle of bulla proper (exclusive of meatal-
tube) and so broadly rounded as not to form a longitudinal
ridge, the region between highest portion and meatus not notice-
ably concave ; teeth smaller than in Meles meles (lower molars
together about 20 mm. in length), with smaller cusps tending to
be more strongly developed ; metacone of m^ relatively larger
than M. meles, so that breadth (jf crown through this cusp is as
great as or greater than that through paracone ; colour as in
Meles meles marianeiisis or slightly ^laler.

Measurements. — For cranial and dental measurements see
Tables, pp. 351, 353.

Specimens examined. — Four, all from Crete.

>5, ?. Lassethe, 2,820 ft. Crete. Miss D. Bate (c). 5.12.2.16-17.

(5. 12. 2. 17. Type of species.)

skull. Katharo, Crete. Miss D. Bate (c). 5. 12. 2. 38.

?. Crete. H. 0. Jones, R.N. (c). 99.6.13.1.

Sub-Family LUTEINS.
1857. Lutiin;v Baird, Mamm. North Amer., p. 148.

Geographical distribution. — Nearly cosmopolitan ; absent only
in the Antarctic and high Arctic regions, Madagascar, Australia
and the Pacific Islands.

Characters. — Teeth of the same general type as in the
Melinse ; skull much flattened, and rostrum so shortened that its
length is less than its width ; external form long and slender,
the fur very dense, the legs unusually short ; toes webbed, the
claws short or absent ; tail long and highly muscular.

BemarJcs. — The family Lutrinse., the members of which appear
to be essentially badgers modified for semi-aquatic life, contains
four genera, one of which occurs throughout Europe from
Ireland eastward.

Genus LUTRA Brisson.

1762. Ltitra Brisson, Eegnum Animale in Classis ix distrib., 2nd ed., p. 13

{Lutra Brisson = Mustcla Intra Linnseus).
1780. Liitra Briinnich, Zoologiae Fundamenta, p. 34. First use of name

by an author following the Linntean system (Mustcla liiira).
1806. Lutris Dumeril, Zoologie Analytique, p. 12 (Modification of Lutra).
1817. Lutrix Rafinesque, Analyse de la Nature, p. 59 (Substitute for Lutra).
1843. Lontra Gray, Ann. and Mag. Nat. Hist., xi, p. 118, February, 1843

{Lutra canadensis Schreber).
1843. Latax Gray, Ann. and Mag. Nat. Hist., xi, p. 119, February, 1843

{Lutra lataxina F. Cuvier). Not Latax Glager, 1827.
1843. Lataxina Gray, List Spec. Mamm. Brit. jNIus., p. 70 {L. mollis Gray

= Lutra lataxina F. Cuvier).
1857. Lutra Blasius, Saugethiere Deutscblands, p. 236.
1865. Barangia Gray, Proc. Zool. Soc. Loudon, p. 123 {B. sumairana Gray

= Lutra barang F. Cuvier).

I

LUTRA 355

1865. Lutrogale Gray, Proc. Zool. Soc. London, p. 127 (Lutra monticola

Hodgson).
1865. Nutria Gray, Proc. Zool. Soc. Loudon, p. 128 {Lutra felina Molina).
1867. Lutronectes Gray, Proc. Zool. Soc. London, p. 180 {L. whiteleyi

Gray = L. lutra Linnseus?).

Type species. — Lutra * Brisson = Mudela lutra Linnaeus.

Geographical distribntion. — Same as that of the sub-family.

Characters. — Skull broad and low (depth of brain-case only
about half mastoid breadth), the entire dorsal profile nearly
straight, the zygomatic arches so widely expanded and post-
orbital region so much narrowed that distance from point of
greatest narrowing to zygoma is greater than postorbital con-
striction ; rostrum so shortened that posterior border of narial
opening approaches level of anterior zygomatic root, and distance
from orbit to gnathion is much less than half length of brain-
case ; auditory bulla flattened, with conspicuous meatal tube ;
paroccipital process low and ridge-like but distinct from bulla ;
dental formula : i ||, c ^ , pm ^ , m ^ = 36 ; teeth large as com-
pared with width of rostrum and palate (greatest transverse
diameter of upper carnassial equal to half distance between
carnassials) ; all of the small premolars opposed and capable of
trenchant action with those of opposite jaw ; upper carnassial
with crown triangular in outline, the outer side occupied by a
moderately high cutting edge formed by two cusps and a con-
necting ridge, the inner side (about half the total crown area)
by a flat crushing surface ; upper molar rhombic in outline, its
crown area about equal to that of carnassial (usually somewhat
smaller), its greatest diameter transvei'so to axis of tooth-row,
the four primitive cusps present ; lower carnassial with three
anterior cUsps well developed, sub-equal, the posterior heel
slightly larger than anterior triangle ; external form highly
modified for aquatic life, the body long and of approximately the
same width as neck and head, the tail long, very muscular, broad
at base, tapering distally, the legs short, feet broad, toes webbed,
short-clawed, the fur soft, dense and impervious to water.

Remarlcs. — Although more widely distributed than any other
living genus of land mammals, Eptesicus and Mi/otis perhaps
excepted, Lutra is not rich in species. A dozen or fifteen forms
are currently recognized, only one of which occurs in Europe.

LUTRA LUTRA Linnaius.

1758. [Mustela] Intra LinniEUS, Syst. Nat., i, 10th ed., p. 45 (Sweden).
1777. [Lutra'] vulgaris Erxleben, Syst. Regui Anim.,i, p. 448 (Renaming of

lutra).
1792. M[ustela] Lutra piscatoria Kerr, Anim. Kingd., p. 172 (Renaming of

lutra).

* Not " Lutra lutra" (see footnote under Meles, p. 841).

•2 A 2

356

CAHNIVOIJA

1816. ? Lutra fluviaiilis Leach, Syst. Catal. Spec. Inclig. Mamni. and

Birds Brit. Mus. , p. G (nomen nudum : '' River Otter ").
1827. [Lutra vulgaris] (8 marinus Billberg, Synopsis Faunas Scandinavia,

p. 28 (Coasts of Scandinavia).
1830. ?[Luti-a] fluviatilis Burnett, Quart. Journ. Sci. Lit. Art. xxviii,

1829, p. 349 (Substitute for lutra), nomen nudum.
1834. \_Lutra] nudipes Melchior, Den Danskc Stats og Norges Pattedyr,

p. 50 (Coast of northern Norway).
1834. Lutra roensis Ogilby, Proc. Zool. Soc. London, p. Ill (Roe Mills,

near Newton Lemavaddy, Londonderry, Ireland). Type in British

Museum.
1857. Lut7'a vulgaris Blasius, Siiugethiere Deutschlands, p. 237.

1884. ILzitra'] lutra Lataste, Actes Soc. Linn, de Bordeaux, xxxviii,p. 34.

1885. [Lutra] angustifrons Lataste, Actes Soc. Linn, de Bordeaux, xxxix,

p. 168, August, 1885 (Bone, Algeria). Perhaps in part only :
specimen from Liguria, Italy, referred to this form on p. 239,
September, 1885. Type in Lataste collection.
1910. Lutra lutra Trouessart, Paune Mamm. d'Europe, p. 86.

Type hiraUty. — Upsala, Sweden.

Geugrapliical distribution. — Europe and northern Africa, east-
ward into Asia ; limits of range not known. In Europe west to
Ireland and north to the Arctic coast.

Diagnosis. — Size medium (head and body in adult male about
700 mm., condylobasal length of skull, 105 to 123); tail about
three-quarters as long as head and body ; naked muzzle pad with
upper border strongly convex at middle ; skull much flattened,
the depth of brain-case not conspicuously more than lialf mastoid
breadth ; interorbital region narrow, its least width less than
distance from front of zygoma to anterior extremity of pre-
maxillary ; teeth not specially enlarged, the greatest diameter of
upper carnassial not greater than width of palate between
carnassials.

External characters. — General form long and slender, the
limbs very short, the feet liroad, with conspicuously webbed toes,
the head short flat and ill-defined from neck, the ears incon-
spicuous, the tail long, broad at base, tapering toward tip,
covered with the same short waterproof fur as body. Head
rounded and flattened, not well defined externally from the
muscular neck ; ear rounded, densely haired on both surfaces,
scarcely rising above level of fur, tlie antitragal lobe valve-like;
a second and third valve-like lobe above and behind meatus ;
muzzle short and wide, the nostril pad entirely naked, its
surface reticulate, its upper margin strongly convex at middle,
the lower slightly so, its lower border separated from mouth by
the densely haired upper lip, the width of which at middle is
about equal to height of pad ; whiskers stiff and bristly, the
longest extending about to ear when laid back. Legs short, feet
broad and rounded, with shoi't toes joined by a naked membrane
extending to base of terminal phalanges ; claws short but strong,
non-retractile, those on fore-feet best developed (about 8 mm. in

LUTRA 357

length) ; psilin bai-e, a Large, hejirt-shaped, obscurely ti'ilobed
tubercular mass behind base of digits, and a round posterior
tubercle separated from the larger pad by a deep groove ; sole
with a bare area and heart-shaped pad essentially as on pahu,
but with posterior tubercle represented by an ill-defined prolonga-
tion of the main pad ; posterior portion of sole densely furred.
Fur very dense and waterpi'oof, alike in texture throughout the
body and tail, the hairs of underfur 10 to 15 mm. in length, the
longer overlying hairs, which almost completely conceal the
underfur, about 25 mm. in length. Mamma? : 6* (probably i 3-3).
Colour. — Winter pelage : upper parts, legs, feet and tail a
rich dark brown (about the prout-brown of Ridgway or some-

FIG. 72.
Lutra Intra. Nat. size.

what darker), with a drabby cast more evident in some lights
than in others, the hairs with a conspicuous metallic gloss ;
underfur light grey, the extreme tips of its hairs changing
abruptly to prout-brown ; on underparts the drab becomes more
coiLspicuous as well as paler, usually assuming a tinge of cream-
l)ufi", the throat and cheeks fading to buffy white ; interramial
region and upper lip with irregulai- white mottlings, the hairs of
which are white to base ; whiskers and claws light horn-colour.

The exact colour is subject to considerable v;iriation, but the
material examined is not sufficient to show whether such
differences as occur are correlated with locality or season.

* Southwell, Field, rii, p. 1043, December 19, 1908.

358

CARNIVORA

Sometimes the brown is darker and richer than usual, or the
drab may be especially pronounced. Occasionally the long hairs
of the back ai'e a light dull buff, imparting to the animal a
peculiar faded appearance.

Skull. — General form of skull broad and flattened, more so
than in any other European carnivore, the depth of brain-case at
middle scarcely more than half mastoid width ; the brain-case exces-
sively narrowed anteriorly, Inroad postei'iorly and with greatly

Fig. 73.
Lufra Intra. Kat. size.

deveKiped lambduid crest, ])Ut low though evident sagittal crest ;
widely spreading zygomata forming a marked contrast with narrow
postorbital region ; r'ostrum short, deep and robust. Dorsal profile
nearly flat throughout, though region in front of middle of post-
orbital constriction slopes gradually downward. Owing to depth
of rostrum and shallowness of brain-case the dorsal and ventral
profiles are approximately parallel. Postorbital processes short
but evident, though not forming any considerable portion of border
of rather large, somewhat upturned orbit, the greatest diameter

LUTRA 359

of which nearly e(]uals width of flattened interorbital region.
Rostrum short, broad and deep, the distance from front of
zygoma to gnathion barelj^ equal to greatest breadth across
canines, about equal to depth at front of orbit, and contained
about 4^ times in condylobasal length of skull. Zygoma
widely spreading throughout, the arch heavy, not much bowed
upward, the anterior root conspicuously perforated by the large
anteorbital foramen the upper margin with low but evident

yiG. 74.

Lutra lutra. Nat. size.

angular projection marking posterior border of orbit. Palate
rather narrow, the distance between molars contained about
three times in that from gnathion to level of posterior border of
molar : incisive foramina moderate, about half as wide as long,
lying entirely between canines ; posterior extension of palate
about ecjual to distance between molars and about one-third that
from molar to hamular ; mesopterygoid space narrow anteriorly,
widei- posteriorly, its length about double the width between
haiiuilars. Auditory bulla small, flattened, though a little

3(i0

CARNIVORA

iiiHated along inner margin ; outline triangular-flask shaped, the
tubular portion of meatus not well defined from rest of bulla ;
least distance between bullse about equal to greatest diameter of
bulla including meatus. Paroccipital process low and ridge-
like, widely removed from inflated portion of bulla. Mandible
very robust, the ramus slightly bowed outward posteriorly, its
lower margin essentially straight except for a slight upward
curve behind level of tooth-row, the depth of ramus at middle
about one-third that through coronoid process ; angular process
very short ; coronoid process narrowly rounded oS" above, its height
above articular process about equal to width at articular level.

Teeth. — The teeth are large and strong, with well developed
cusps and commissures and relatively small crushing ai-eas, the
small premolars of upper and lower mandible titting closely
between each other when jaws are shut ; crown area of upper
molar about equal to that of car-
nassial, sometimes less. Upper
incisors forming a nearly straight
transverse row, the anterior faces
of the smaller teeth exactly in line,
those of the two larger teeth slightly
more posterior ; i^ and i~ small,
sub-equal, the shafts compressed,
the cutting edges rounded, simple ;
P higher than the others, somewhat
resembling the lower canine in form,
its apex curved outward and back-
ward beyond level of smaller teeth,
its posteroexternal surface with two
concavities separated by a trenchant
ridge ; lower incisors subterete, the
crown of v, obscurely and unsym-
metrically bilobed, that of the others
simple ; cross section of i-^ about one-third that of iy Canines
robust though not greatly elongated ; shaft simple, v\ithout
evident cingulum, that of upper tooth slightly curved backward,
that of lower tooth abruptly recurved, its axis set obliquely
outward. Anterior premolar both above and below well
developed and functional, the point of the upper tooth lying
internal to that of lower when jaws are closed ; pm^ small, single
rooted, its crown area about equal to that /^, its form es.sentially
like that of succeeding tooth, though with cusp relatively less
developed ; pm" and ptiv^ successively larger, two-rooted, the
crown area of imi^ fully three-fourths that of canine, each tooth
with a well-developed cusp lying in front of middle of crown,
and a distinct anterior and posterior cutting ridge ; cingulum
moderately developed ; outline of crown of pm'^ elliptical, that of
p)m^ ^with inner margin bulging inward behind middle, though
without developing a deflnite inner lobe ; pm., and pm^ much like

Flc. 75.
IjUtra Intra. Teeth.

LUTRA 361

pin- in l)oth size and form, the three teeth cutting a<j;ainst each
other when jaws are closed ; pm^ larger than the other small
premolars, its cusp l<3ss anterior in position, its crown decidedly
in wider posteriorly than anteriorly. Upper carnassial triangular
general outline, though this is somewhat obscured by the broadly
rounded inner portion and the somewhat projecting postero-
external lobe ; outer and posterior borders sub-equal and longest ;
trenchant portion of tooth with high anterior cusp connected
with well developed posterior cusp by a high, abruptly angled
commissure; crushing portion nearly as wide as long, its area
about as great as that of trenchant portion, its inner border
noticeably raised at middle though without secondary cusps ;
cingulum moderately developed along outer border of crown, and
forming a distinct though small antero-basal cusp. Lower
carnassial with anterior triangle well developed, the protoconid
and paiaconid sub-equal, the metaconid somewhat smaller ;
crushing portit)n of crown wider than antei'ior triangle though
not so long, the areas of the two portions of the tooth
approximately equal ; hypoconid evident though not high, the
outer surface of its base in line with that of protoconid, the
inner surface continuous with the concave though scarcely
basin-shaped main portion of crushing area ; cingulum narrow
but evident throughout, not specially developed in region
bordering outer base of hypoconid. Second lower molar flat,
subterete, slightly wider than long, the inner and outer margins
each with a small cusp somewhat behind middle, the cusps joined
by a low transverse ridge. Upper molar about equal to carnassial
in crown area, but length of its outer border decidedly less than
that of preceding tooth ; crown slightly constricted near middle,
the outer portion bilobed, the anterior lobe bearing a low paracone
and a broad outer jjrojection representing the parastvle, the
posterior lobe bearing a low but robust metacone ; iinier portion
of crown with a large protocone and low hypocone, the former
sending forward a conspicuous commissure to base of paracone,
the latter connected with cingulum that extends around base of
protocone.

Measitrements. — Adult male from Warwickshire, England :
head and body, 712 ; tail, 495 ; hind foot, 134 ; ear, 28. Adult
female from Csehtelek, Hungary : head and body, 640 ; tail, 380 ;
hind foot, 115; ear, 22. For cranial and dental measurements
see Tables, pp. 362, 363.

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

Ireland : Londonderry, 1 (t}'pe of roensis) ; Ahascragh, Co. Galway, 1.

England : Pembrokeshire, 1 ; River Stour, Dorsetshire, 1 ; Rugby,
Warwickshire, 1 ; Norfolk, 1 ; no exact locality, 1.

Norway: Egersund, Stavanger, 5.

Sweden : Jockmock, Lappmark, 1 (Stockholm) ; Gnesta, Soderman-
land, 1 (Stockholm) : Skane, 1 (Stockholm) ; no exact locality, 2.

France : St. Gilles, Gard, 1 (Lataste) ; Etupcs, Doubs, 1 (IMotta/).

Germany : Southern Germany, 2.

362

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364

CARNIVOKA

Austria-Hungary : Karlsbad, Bohemia, 1 ; Csehtelek, Bihar, Hun-
gary. 1-

Italy: Torriglia, Liguria, 1 (Lataste).

Spain : Castrillo de la Eeina, Burgos, 3 (B.M. and U.S.N.M.) ; Galicia, 1 ;
Seville, 5.

Seiuarlcx. — Although the specimens show some rather note-
worthy variations in cohnir, and in cranial and dental measure-
ments, the matei'ial examined is not suthcient to indicate the
existence of geographical races in Europe ; while the north
African animal to which the name augustifrons has been applied
appears to be not separable from true Intra.

1.

Newton Lemavady,
Londonderry, Ireland.
{Miss A. Moody.)
6 st. Ahascragh, Galway.

3 juv. st. Pembrokeshire, Wales.

6. Rugby, Warwickshire,

England.
? St. R. Stour, Dorset.

1. England.

4 (i, 1 9, Egersund, Stavanger,
skulls. Norway.

ad., juv. South Germany.

6. Csehtelek, Bihar, Hun-

gary.
1. Karlsbad, Bohemia.

S, 9. Castrillo de la Reina,

Burgos, Spain.
1 juv. Galicia.

skull. Seville.

9, juv. Seville. {Dr. A. Ruiz.)
skull. Seville.

Zoological Society's 67. 12. 14. 4.

Museum. (Type of L. roensis
Ogilby).
Purchased (Row- 5. 7. 10. 1.

land Ward).
Purchased (Row- 6. 5. 21. 1-4.

land Ward).
Dr.T. S. Townsend 9. 12. 16. 1.

(c & p).
J. C. Mansel Pley- 98. 6. 13. 1.

dell (c & p).

K. H. Schaanning
(c).

Dr. A. Giiuther (c).

Hon. Mrs. N. C.
Rothschild (p).

Lord O. Russell
(c & p).

Rev. Saturio Gon-
zalez (c).

Dr. V. L. Seoaue
(c & p).

Lord Lilford (p).

Lord Lilford (i').
Col. L. H. Irby
(c & p).

99. A.
11.6.3.14-18.

59. 9. 6. 62-68.
10. 9. 14. 1.

57. 2. 14. 1.

8. 7. 7. 16-17.

94. 6. 18. 1.

74. 10. 7. 2.
76. 3. 4. 1.
95.9.4.10-11.
2. 11. 16. 1.

Sub-Family MUSTELINiE.
1835. Mustelimr Swainsou, Nat. Hist, and Classif. Quadr., p. vii.

(jrfO(jr<iphiral (Jidrlhntivn. — Northern hemisphere, south into
northern Africa, the Malay Archipelago, and northern and
western South America ; in Europe west to Ireland.

Character!^. — Dentition highly trenchant, upper carnassial
without crushing surface other than a small concave ai'ea between
inner lobe and main cusp, the crown much longer than broad,
not triangular or rhombic in outline, the posterior cusp compressed,
trenchant, barely half as high as main cusp ; upper molar much
reduced, the length of its outer portion one-third to one-half that
of carnassial ; small premolars alternating when jaws are closed,
at least one pair capable of shearing action ; skull varying in

MARTES 365

form but never greatly flattened and never very robust, the
rostrum always at least as long as broad ; external form slender,
the legs usually rather short ; feet digitigrade ; toes partly
webbed ; tail varying in length, slender or bushy, never con-
spicuously muscular.

Reinarhx. — As here understood the sub-family Mustelinse
contains the three genera, Maries, Miisfcln and VormeJa, all of
w Inch occui' in Europe.

Genus MARTES Pinel.

1792. Murtes Pinol, Actes Soc. d'Hist. Nat., Paris, i, p. 55 (M. domestica

Pinel = M. foina Erxleben).
1820. Martcs Nilsson, Skand. Fauna, i, p. 38 (M. foina and M. sylvatica =

martes).
1829. Zibellina Kaup, Entw.-Gescli. u. Natiirl. Syst. Europ. Tliierw., i,

p. 31 (M. zibellina).
1857. Mustela Blasius, Siiugethiere Deutsohlands, p. 211.
1911. Martes Thomas, Proc. Zool. Soc, London, p. 139, March, 1911.

Type species. — Mustela martes Linnaius.

Geographical distrihution. — Isorthern hemisphere from the
limits of tree growth south to the Mediterranean, the Malay
Archipelago, and the central United States ; in Europe west to
Ireland.

Characters. — Skull narrow, moderately high (depth of brain-
ca.se much more than half mastoid breadth), the dorsal profile
moderately curved, the zygomatic arches not specially wide-
spreading, and postorbital region not unusually narrowed (distance
between region of greatest narrowing and zygoma normally less
than breadth of postorbital constriction) ; rostrum narrow and
somewhat elongate, its width noticeably less than that of inter-
orbital region, the distance from anterior rim of orbit to gnathion
exceeding width of rostrum Ijetween anteorbital foramina; auditory
bulla3 moderately inflated, the meatal tube evident though .short,
the longitudinal diameter of bulla greatest ; paroccipital process
small, slightly projecting, partly distinct from bulla ; dental
formula : / — , c — , pm t*, in -* = 38 ; cutting edges of five small

;i-;r i-i' -^ 4-4' '2-2 ' o o

premolars (2 upper and 3 lower) capable of trenchant action ;
upper carnassial long and narrow, not triangular in outline and
without crushing surface, the small inner lobe standing as an
ofl'set to anterointernal extremity of crown, the sectorial portion
consisting of a high anterior and low posterior cusp with some-
what concave connecting ridge ; upper molar pyriform or pan-
durate in outline, its long axis nearly perpendicular to that of
tooth-row, its crown mainly fiat, but with a small paracone, still
smaller, sometimes obsolete metacone, and crescentic ridge-like
protocone ; lower carnassial wider posteriorly than anteriorly,
the anterior triangle much distorted, the metaconid reduced to

3G6

CARNIVORA

a well detined postero-internal process on base of protoconid, the
posterior crushing heel slightly more than half as large as
trenchant jiortion of tooth ; external form slender, somewhat
catlike or squirrel-like, the head moderately elongated, the
muzzle pointed, the ears high and conspicuous, the tail long,
bushy ; feet digitigrade, the moderately long claws partly
retractile ; fur long, dense and soft.

Hemnrks. — The genus Martes contains about ten species, some
of which are represented by numerous geographical races. Three
are known from westei'n and southern Europe.

KEY TO THE EUROPEAN FORMS OF MABTES.

Third upper premolar with crown strongly
convex on inner side, slightly concave on
outer side ; width of inner lobe of upper
carnassial nearly equal to that of tren-
chant portion of cro-wn ; greatest diameter
of upper molar about equal to length of
outer border of carnassial ; fur usually
finer and softer, and throat-patch more

yellow (Pine Martens) M. martes, p. 366.

Throat-patch cream-buff, general colour

darker (Central and northern Europe)... ,V. m. martes, p. 372.
Throat-patch buff-yellow, general colour

lighter (Mediterranean region) M. m. latinorum, p. 373.

Third upper premolar with crown evenly bicon-
vex ; width of inner lobe of upper car-
nassial barely half that of trenchant
portion of crown ; greatest diameter of
upper molar noticeably less than length
of outer border of carnassial ; fur usually
coarser and less soft, and throat-patch more
whitish (Beech Martens).
Condylobasal length of skull in adult male,
76 to 79 mm. ; pale throat patch always
much encroached on by brown of sur-
rounding parts, occasionally obliterated

(Crete) ! M. bunites, p. 380.

Condylobasal length of skull in adult male,
7'J to 84 mm. ; pale throat-patch seldom
much encroached on by brown of sur-
rounding parts and never obliterated
(Central and southern Continental

Europe) M. foina, p. 374.

General hue of upper parts drab iV. /. foina, p. 375.

General hue of upper ]5arts wood-brown .. M.f. mediterranea, p. 380.

MARTES MARTES Linna>US.

(Sj'uonymy under subspecies.)

Geographical distribution. — Entire wooded region of Europe,
from Ireland eastward into Asia, and from the Mediterranean
coast and islands northward to the limits of tree growth.

Diagnosis. — Third upper jjremolar with outline of crown

MARTES 357

concave on outer side, irregularly convex on inner side, the rec^ion
ot greatest width behind middle ; upper carnassial with inner
projection robust, its diameter in line of tooth-row nearly or
quite equal to greatest width of trenchant portion of tooth behind
middle of crown ; upper molar large, its greatest diameter about
equal to outer length of carnassial, the metacone more than half
as large as paracone, the outer border of the crown usually
notched; general colour yellowish brovvn, the throat patch

Fig. 76.
Martes martes. Nat. size.

always distinctly tinged with yellow ; underfur moderately long

External charactcrs.—Fovm slender and graceful, suggesting
that of both cat and squirrel ; legs moderately long ; tail about half
head and body, densely furred and bushv, the hairs at tip about
one-third as long as vertebr*. Head pointed ; eyes of medium
size ; ear rising distinctly above fur, its height from crown about
^5 mm. the outline rounded but forming a slightly indicated
point above, the entire surface densely furred : muzzle pad

368

CARNIVORA

well defined, completely naked, communicating with upper lip
by narrow median line. Feet densely furred throughout in
winter, the pads bare in summer ; palm with deeply divided,
trilobed, heart-shaped tubercular mass at bases of median digits,
a small round pad at base of thumb, and a larger round pad at
outer posterior border near wrist ; sole with similar trilobed
mass, but elements more distinct, almost forming three tubercles :
small pad at base of hallux like that at base of thumb ; no
posterior pad. Fur very dense and soft, the hairs of undei'fur
about l'5 mm. long at middle of back in winter, shorter in
summer, the longer hairs reaching about 40 mm. Mamma? : 4.

Colour. — The colour varies considerably with season and
climate, but is usually characterized by rich yellowish brown and
blackish tints, the legs, tail and underparts darker than the
back, the head noticeably lighter than the body, the throat and
fore part of chest yellowish, and interramia, chin and muzzle
dusky. Claws light horn-colour.

Skull. — As compared with that of the European species of
Mustela the skull of Marfcs marten is characterized by a general

Fig.
Martes inartex

smoothness, lightness and lack of angularity. Dorsal profile
strongly convex posteriorly, flattish behind orbits, the rostrum
falling away at an angle of about 20' ; a slight concavity usually
present near middle of nasals. Brain-case elongate-ovate in
outline when viewed from above, nearly as deep as wide, the

MARTES 369

sagittal crest moderately developed in old individuals and slightly
overhanging occiput. Floor of brain-case with no special
features ; a slightly developed median ridge. Auditory bulla?
moderately inflated, the surface smooth, the general outline
broadly flask-shaped (longitudinal diameter greate.^t), the meatal
tube well defined, i-ather wider than long ; space between buUie
about equal to diameter of bulla without tubular portion.
Postorbital region rather abruptly constricted, much as in
Mustda, its width about half that of brain-case. Interorbital

I'IG. 7S.
Marten martes. Nat. size.

region wider than rostrum, the discrepancy in breadth greater
than in Mustda crminen. Postorbital processes well developed
though short. Rostrum relatively longer than in 3Iustda
erminca or M. ptitoriits, the distance from orbit to gnathion about
equal to width between outer margins of anteorbital foramina,
the width across canines much less than distance from foramen
to gnathion. Orifice of anteorbital foramen over anterior root of
carnassial, and under anterior border of rather large orbit.
Zygamata rather widely but gradually spreading, strongly bowed

2 b

370 CARNIYOKA

upward behind middle. Palate rather narrow, its width
between molars equal to about 1^ times greatest diameter of
molar, incisive foramina small, ovate, slightly oblique, at level of
front of canine, the minute median foramen slightly behind
middle ; posterior extension of palate broader than long, reaching
about half way from level of molar to hamular ; mesopterygoid
space about a quarter longer than broad, the hamulars short,
sHghtly everted.

Teeth. — In proportion to size of skull the teeth are larger
than in the members of the other genera of MusteHme occurring
in Europe ; transverse diameter of upper molar much more than
half width of palate between molars. Upper incisors forming a
straight row separated at each side from canine by a space about
equal to transverse diameter of i^ and P together ; crowns strongly
compressed, the antero-posterior diameter fully twice transverse
diameter, the anterior face convex, the posterior concave with
faintly indicated cingulum ; ?'- slightly
larger than ?'^ i^ abruptly much larger
than the other two teeth together, its
crown more than one-third as high as
that of canine and with the posterior
concavity extending to outer basal
25ortion. Mandibular incisors smaller
and lower than the upper teeth, their
crowns projecting obliquely forward,
the root of r.^ implanted in jaw behind
level of the other two ; crowns ob-
scurely bitid, the outer lobe smaller
than the inner ; crown area of i^
pj(, -g about one-third that of the com-

Martcs martes. Teeth. Nat. size, pressed 'i.^ ov the subterete /,. Canines
with no special peculiarities ; diameter
of the upper tooth along alveolus equal to one-half width of
jialate between canines, that of lower somewhat more ; shaft
of upper tooth with posterior longitudinal ridge, and an antero-
internal ridge which near base curves backward across inner
side of shaft nearly to base of posterior ridge ; cingulum barely
indicated ; shaft of lower canine directed a little forward at
base, a little backward beyond middle, the anterior profile
strongly convex, the posterior equally concave ; surface of
enamel, e.specially on basal half of tooth, much roughened by
longitudinal wrinkles. Anterior premolar both above and below
single-rooted, small (crown area about equal to that of i^ and i'-'
together), subterete, the height of the single ill-developed cu.sp
scarcely ecjual to diameter of crown in p7)i^, much less in pm.i.
Other premolars, except upper carnassial, two-rooted, compressed,
the outline of crown triangular when viewed from the side, with
apex slightly in front of middle of tooth. Crown area of pw*''^
about three times that oi^mi^ ; outline of crown flattened-elliptical

MAllTES 371

or slightly concavo-convex, tlio concavity, when present, on outer
side; cusp simple, its height a little less than length of crown
along hase, its posterior surface with slightly developed longi-
tudinal ridge. Crown of j)Hi'^ somewhat longer than that of piii^,
but its area much greater owing to widening of strongly convex
inner side ; cusp essentially as in pni^ though somewhat higher.
Second lower premolar about equal in size to jyin-, but axis of
shaft more anterior in position ; pm.-^ essentially like pm^, but
cusp with faintly indicated anterior longitudinal ridge, and
posterior ritlge with a slight nodule or rudimentary basal cusp
(sometimes absent) ; pm^ noticeably larger than pm-^ and with a
well developed secondary cusp at middle of posterior border of
main cusp. Upper carnassial ( pm*^) long and narrow, the width
of crown just behind internal lobe less than half length along
outer border, the internal lobe robust, its diameter in line of
tooth-row nearly or quite equal to width of trenchant portion of
crown, its axis nearly perpendicular to main axis of tooth, its cusp
well developed, about as high as that of pm^, and separated from
main cusi? by deep concavity ; main cusp robust, its height
contained about 1§ times in length of crown, its axis slanting
a little backward, its anterior border with well developed
longitudinal ridge ; posterior cusp about half as high as main
cusp, its outer surface sloping obliquely to well developed
cingulum, the two cusps connected by a high, sharply trenchant
and obtusely angled commissure. Lower carnassial with crown
nearly 2^ times as long as broad, the sectorial portion of the
tooth consisting of a much distorted triangle, the paraconid
forming anterior extremity of tooth, its anterior border nearly
perpendicular, its posterior commissure meeting anterior commis-
sure of the higher protoconid at an abrupt angle, the two together
acting in opposition to the angled commissure of upper carnassial ;
metaconid reduced to a subterete postero-internal process on
base of protoconid, its area scarcely one-sixth that of larger
cusp ; crushing portion of crown slightly broader than trenchant
portion, its outline sub-circular with slightly raised edge, this
edge forming a noticeable though low posteroexternal cusp
separated from base of protoconid by a shallow but distinctly
angled notch. Second lower molar about equal to heel of
carnassial in size and essentially like it in form, the crown flat,
with faintly indicated outer ridge and low postero-internal cusp.
Upper molar large, its crown area at least equal to that of
carnassial, its greatest diameter about equal to outer length
of carnassial (see measurements, p. 378), the diameter of inner
portion of crown usually much greater than that of outer
portion, the median constriction well marked ; outer portion of
crown with two small cusps, probably the ]iaraoone and metacone,
tlie latter more than half as large as former, the outer margin of
tooth often though not invariably notched between them ; inner
portion of crown with a slightly curved, ridge-like antero-

2 B 2

372

CARNIVORA

external cusp (usually notched or partly divided into two), the
surface elsewhere finely wrinkled ; in some specimens a minute
tubercle, perhaps representing the hypocone, occurs near posterior
border of crown in region of constriction between outer and
inner portions ; cingulum low but evident, especially around
inner margin of crown.

Hemarks. — The well - known pine marten is so strongly
characterized as to require no special comparisons with any other
European species except Maries foina (see account of latter).
Two geographical races are currently recognized, though their
status is by no means clear.

Martes martes martk.s Linn;eus.

1758. [Mtistcla'] martes Linnaeus, Syst. Nat., i, lUth ed., p. 46 (Sweden).
1816. Miustda] sylvestris Oken. Lchrb. d. Naturgescli., iii, pt. 2, p. 1029

(Renaming of martes).
1827. M[artcs] vulgaris Griffith, Cuvicr's Anim. Kiugd., v, p. 123

(Renaming of martes).
1847. Martes sylvatica Nilsson, Skand. Fauna, i, 2nd ed., p. 41 (Renaming

of martes).
1857. Mustela martes Blasius, Siiugetliiere Deutschlands, p. 213.
I'JIO. Mustela martes Trouessart, Faune Manim. d'Europe, p. 72.
1911. [Maries'] martes Thomas, Proc. Zool. Soc. London, p. 139, March,

1911.

Tijpe locality. — Vicinity of Upsala, Sweden.

Oeogrn-pliical distribution. — Europe north of the Mediterranean
region, from Ireland eastward into Asia.

Diagnosis. — Throat patch cream-bufF or slightly more yellow.

Culoar. — Upper parts a rich dark brown, usually rather near
the bister of Ridgway, the tips of the longer hairs blackish ;
underfur light grey, the tips of the hairs tinged with drab or
with wood-brown ; face essentially like back or not so dark, the
muzzle and chin usually not darker than forehead ; ear edged
with bufly drab ; tail very dark brown (nearly the seal-brown of
Ridgway), its underfur raw-umber or somewliat darker ; feet
and lower portion of legs blackish ; throat-patch varying
considerably, but as a rule rather pale, nearly cream-buif or
slightly more yellow.

Measurements. — For cranial and dental measurements see
Tables, pp. 376, 378.

Specimens examined. — Twenty, from the following localities : —

Ireland : Kenmare, Kerry, 1 ; Go. Kerry, no exact locality, 1.

Scotland : No exact locality, 1.

England: Keswick, Cmnberland, 1; Cockerniouth, Cumberland, 1;
Pontrilas, Herefordshire, 1 ; Ludford Park, Herefordshire, 1 ; no exact
locality, 3 (B.M. and U.S.N.M.).

Norway : Egersund, Stavanger, 3.

Sweden : No exact locality, 3 (U.S.N.M.).

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

Germany : Southern Germany, 2.

Austria-Hungary : Sohemia, 1.

MARTES '"^TS

9. Kenmaro, Kerry, Ire- E. Dodson (c). 94. 3. 27. 1.

land.

1. Kerrv. J. M. Gurney (p). 72. 1. 10. 1.

1. Scotland. (McLcaij.) Hargitt Collectiou. 8G. 9. 9. 1.
St. Keswick, Cumberland, W. R. Wilson (c). 96. 11. 6. 1.

England.
6. Cockermoutli, Cum- W. R. Wilson (c). 97. 1. 15. 1.

berland.

5. Pontrilas, Hereford- Mrs. St. John A. 85. G. 10. 1.

shire. Matthews (p).

6. Ludford Park, Here- H. J. Bailey (c & p). 0.2.23.1.

fordshire.

2. Wales (1828). Earl Cawdor (p). Gl. c. d.
2(5,19, Egersund, Stavanger, K. H. Schaanning 11.6.3.19-21.

skulls. Norway. (c).

2 skulls. Southern Germany. Dr. A. Giinther (c). 188i, 1299f.

6. Bohemia. Lord 0. Russell 50. 12. 23. 1.

(c & p).

Martes MARTES LATiNORUM Barrett-Hamilton.

1904. Mitstela martes latinorum Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., xiii, p. 889, May, 1904 (Nurri Mountains, Sardinia).
1910. Mustela martes latinorum Trouessart, Faune Mamm. d'Europe,

p. 72.

Tyjye locallfi/. — Nurri Mountains, Sardinia.

Geograi)liical distrihntkm. — Mediterranean region (Italy, Sar-
dinia, Balearic Lslands).

Diagnonig. — Throat j^atch buft-yellow ; general colour lighter
than in true martes.

Colour. — Upper parts a rather light brown, the longer hairs
between the raw-umber and mars-brown of Ridgway, the
general effect darker in certain lights, e.specially on posterior
half of body ; underfur a pale bluish grey, essentially as in true
martes, the tijas of the hairs ranging from pale wood-brown to
clear butf; face Isabella-colour streaked with whitish hairs, the
muzzle and lips between mars-brown and j^i'out-brown ; ear
edged with bufly drab ; tail like back at base, darker at tip ;
feet and legs washed with dark brown ; throat-patch varying
considerably, but as a rule approaching buff-yellow.

Measurements. — Adult male from Porlezza, Como, Italy :
liead and body, 470; tail, 235; hind foot, 94:'6; ear from
meatus, 42. Adult female from San Cristobal, Minorca, Balearic
Islands : head and body, 430 ; tail, 230 ; hind foot, 87 ; ear
from meatus, 45. For cranial and dental measurements see
Tables, pp. 376, 378.

Specimens examined. — Nineteen, from the following localities : —
Italy : Porlezza, Como, 11 (Ghidini) ; near Rome, 4 ; Milan, 1.
Sakdinia : Nurri Jlountaius, 1 (type).
Spain : Maucor, Majorca, Balearic Islands, 1 ; San Cristobal, Minorca, 1.

Itemarlcs. — The status of this form is very unsatisfactory. So
fai- as can be judged from flu- few specimens examined there is

374

an average difference, however, between the Mediterranean
animal and that occurring north of the Alps. Two of the
specimens from Porlezza are darker than the others, in this
respect agreeing with the northern animal.

1. Italy. (C. Coli.)

1. Milan.

1. iSTurri Mountains, Sar-
dinia.
9. JIancor, ilajorca, Bale-
aric Islands. (Riutoii.)
9. San Cristobal, Minorca.

G. Barrett-Hamilton

E. Caveudish Taylor

(P)-
E. N. Buxton (p).

{Type of subspecies.)
0. Thomas (p). 1. 3. G. 1

11. 1. 2. 29.

5. 6. 6. 8.

95. 4. 16. 1.

0. Thomas & R. I. 0. 7. 1. 43.
Pocock (c & p).

MARTES FOINA Erxlehen.

(Synonymy under subspecies.)

Geograpliical distrihuiion. — Central and southern Continental
Europe, from the Atlantic coast eastward, and from the
Mediterranean to the Baltic.

Dkt(jn<isls. — Third upper premolar with outline of crown
nearly biconvex, the outer side occasionally
flattened, the gieatest transverse diameter
at middle ; upper carnassial with inner
lobe slender, its diameter in line of tooth-
row equal to only about half greatest
width of trenchant poi'tion of tooth behind
middle of crown ; upper molar not so large
as in M. martes, its greatest diameter less
than outer length of carnassial, the meta-
cone less than half as large as paracone ;
external form as in M. marten, but fur
of less fine quality ; colour usually more
greyish or drab than in 31. martes and seldom with the rich
brown tints of the related species, the throat-jjatch never strongly
tinged with yellow. Mammte : i.

Skull. — The skull resembles that of Marion martes, but may
usually be distinguished by its greater breadth and less depth.
Brain-case noticeably wider than high, the general outline when
viewed from behind nearly as in Mimtela erminea and distinctly
less elevated than in Martes inartes. Interorbital region and
rostrum wider than in Martes martes, and concavity ()f dorsal
profile in nasal region much more pronounced. Anteorbital
foramen usually smaller than in the related species, though in
the same position. In other respects the skulls of the two
animals show no tangible diflerences.

Teeth. — Except as already pointed out the teeth agree with
those of Martes martes.

KiG. so.

Lfirgei' cheek-teeth of Marte-i
7nartes (re) and M.foina {h).
JS'at. size.

MAETES 375

BemnrJiS. — Though readily distinguishable from Martes marles
by tlio character of the skull and teeth iff. foina is sometimes
difficult to recognize by external peculiarities alone. Usually
the colour has a slaty or drab cast that is highly characteristic,
and the <|uality of the fur is inferior to that of the pine marten ;
but I have seen tanned skins of animals killed in winter which
were impossible to identify with any degree of certainty.

Two geographical races have. been described.

Martes foina foina Erxleben.

1777. {Mustela] foina Erxleben, Syst. Regni Anim., i, p. 458 (Germany).
1792. Martes domestica Pinel, Actes Soc. d'Hist. Nat., Paris, i, p. 55 (France).
1801. M[;ustela] f[oina] alba Bechsteiu, Gemeinu. Naturgesch. Deutsch-

lauds, I, 2nd ed., p. 759 (Thiiringen, Germany).
1857. Mustcla foina Blasius, Siiugethiere Deutschlands, p. 217.
18G9. Mustcla martes var. fagorum Fatio, Famie Vert. Suisse, i, p. 318

(Sweden) : name wrongly attributed to Linnaeus, Syst. Nat., i,

12th ed., p. ()7.
1910. Mustcla foina Trouessart, Faune Mamm. d'Europe, p. 72.

Type locality. — Germany.

Geographical distrihution. — Range of the species except
southern Spain.

Diagnosis. — Longer hairs of back tipped with sepia, the
general hue of upper parts drab.

Colour. — Underfur very light grey (about grey No. 10 of
Ridgway) with an evident buffy cast on distal third of hairs;
long hairs ranging from wood-brown to mars-brown, becoming
darker at tips. As the long hairs nowhere conceal the underfur
the general effect is a light drab resulting from the blending of
the two colours, very uniform thi'oughout dorsal surface, though
usually a little paler on neck and darker on posterior portion of
back. Tail essentially like body, but darkening toward tip, the
pencil often blackish. Legs and feet washed with dark brown.
Ear huSy drab externally, creamy white along rim, the inner
surface greyish white. Muzzle, lips and chin bi'occoli-brown.
Throat-patch buffy white, darkening to cream-buff' along outer
edges.

Measurements. — Adult male from near St. Gallen, Switzer-
land : head and bodj?^, 453 ; tail, 260 ; hind foot, 85 ; ear from
meatus, 3-1. For cranial measurements see Table, p. 377.

Specimens examined. — Twenty-three, from the following localities : —

France : ]\Iauonville, Mourthe-et-Moselle, 2.

Germany: Ingelheini, Rheinhessen, 2; Nuremberg, Bavaria, 1
(U.S.N.M.) ; southern Germany, 7 (skulls).

Switzerland : Geneva, 1 (Mottaz) ; Vallee-de-Joux, Vaud, 1 (Mottaz) ;
Thavngeu, SchaShauseu, 3 (B.M. and U.S.N.M.) ; Aarburg, Aarau, 1
(U.S.N.M.); Oberrich, St. Gallon, 1.

Spain : Silos, Burgos, 1 ; Donasantos, Burgos, 1 (U.S.N.!M.).

Italy : Near Rome, 1.

376

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380

CARNIVORA

Greece : Mt. Vuno, Cophalonia, 1 (colour normal ; size apparently
rather small).

?, juv. IManonviUe, Meiirthe-et- Lord Lilford (p). 95. 9. 5. 1-2.

Moselle, France.
6. Ingelbeim, Eheinhessen, C. Hilgert (c). 8. 11. 2. 17.

Germany.
6. Ingelheim, Eheinhessen. G. Barrett-Hamilton 11. 1. 2. 102.

7 skulls. South Germany. Dr. A. Giinther (c). 1299, a-g.

2 S juv. Schaffhausen, Switzerland. 0. Thomas (p). 2. 8. 4. 22-23.

(E. H. Zollikofcr.)

6. Oberrich, St. Gallen. 0. Thomas (p). 2. 8. 4. 21.

{E. H. Zollikofer.)

S. Silos, Burgos, Spain. Rev. Saturio Gonza- 8. 7. 7. 21.

lez (c).

1. Rome, Italy. B. Cavendish Taylor 5. 5. C. 7.

1. Mt. Vuno, Cephalonia, J. I. S. Whitakcr (p). 8.10.1.9.

Greece. (C. Mottaz.)

Martes foina mediterranea Barrett-Hamilton.

1898. Mnsfcla mediterranea Barrett-Hamilton, Ann. and Mag. Nat. Hist.,

7th ser., i, p. 442, June, 1898.
1910. Mustela foina mediterranea Trouessart, Faune Mamm. d'Europe,
p. 73.

Type locality. — Sierra de Jerez, Cadiz, Spain.

Geographical clistribution. — Southern Spain.

Characters. — Longer hair.s of back tipped with a light
yellowish brown, the general hue of upper parts lighter, yellower
and less drab than in true/o/)(«.

Measurements. — For cranial and dental measurements see
Tables, pp. 377, 379.

Specimens examined. — Two, the type, from the province of Cadiz, and
a second specimen (mummy) from the Sierra Nevada.

BemarTcs. — If the two individuals of Martes foina mediterranea
are not abnormal in colour the race is rather well characterized.
I have seen only one skin of true foina that ajaproaclies them, a
specimen from Rome.

1 juv. Sierra de Jerez, Cadiz, Spain. A Chapman (c & p). 98. 3. 18. 1.

(Type of subspecies.)

MARTES BUNITES Bate.

1899. Mustela foina leucolaclin:ia Barrett-Hamilton, Ann. and IMag. Nat.

Hist., 7th ser., iv, p. 383 (Not of Blauford).
190G. Mustela foina bunites Bate, Proc. Zool. Soc. London, 1905, ii, p. 318,

April 5, 190G.
1910. Mustela foina bunites Trouessart, Faune ^Nlamm. d'Europe, p. 73.

Tyjje locality. — Kontopalo, Kania, Crete.

(reograjMcal (listrihution. — Crete.

Diagnosis. — In general like Martes foina, but size not so

MUSTELA 381

large (condylobasal length of skull in adult male not reaching
80 inm.), rostral portion of skull not so broad, and pale throat-
patch so much encroached on by brown of surrounding jiarts,
that it is occasionally absent.

Colour. — The colour is lighter and more yellowish (less drab)
than in Maries foina folna, as the long hairs continue a clear
wood-brown to tips, producing a general effect more nearly a
light isabella-colour than drab. The difference is, ho\\ever, not
very great. Tail and feet essentially as in M. f. foina, though
not so dark. Throat-patch greatly reduced, absent in one of the
se^■en skins,* and in its most extreme development represented
by a mere horseshoe-shaped mark, one extremity of which lies in
front of each fore leg, while the anterior portion broadens out
to form an irregular patch covering entire throat immediately
behind interrainial region.

Slodl and teeth. — The skull and teeth resemble those of
Martrs foina except for their uniformly smaller size and the
relatively somewhat less breadth of the rostrum.

Meamrrmcnts. — External measurements of tyjae (young-adult
male) : head and body, 403 ; tail, 255 ; hind foot, 79 ; ear from
meatus, 39. For cranial and dental measurements see Tables,
l^p. 377, 379.

Specimens examined. — Seven, all from Crete.

Miss D. Bate (c). 5. 12. 2. 18-20.

Miss D. Bate (c). 5. 12. 2. 21.

Miss D. Bate (c). 5. 12. 2. 22.

(Type of species.)
Miss D. Bate (c). 5. 12. 2. 37.

H. 0. Jones, K.N. (c). 99. 6. 13. 2-3.

Genus MUSTELA Linnreus.

(Sj-nonymy under sub-genera.)

Type species. — Mustela erminea Linnaeus.

Geoijrapliiral distribution. — Northern hemisphere fi'om the
Arctic coast south in the Old World to northern Africa and the
Malay Archipelago, and in America to the Andes ; in Europe
east to Ireland.

Characters. — 8kull in general resemljling that of Martes, but
rostrum so shortened that distance from orbit to gnathion is less
than width of rostrum between anteorbital foramina ; auditory
bulla without meatal tube, its outline variable but never Hask-
shaped ; paroccipital process small and flattened, closely applied to
posterior martrin of bulla ; dental formula : i 1"-, c — , mm —

* .\ small tuft of white hair persists behind each angle of mouth, and
another in front of each fore leg, the largest of .these about 15 mm. in
diameter.

6,2.
1.

i.

Kanea, Crete

Katharo.

Kontopalo.

skull.
2.

Crete.
Crete.

382 CAENIVORA

m ^1 = 34 ; upper carnassial as in Maries, its posterior (;usp low
but well developed, the height of its main cusp about half outer
border of crown ; upper molar between pyriform and pandurate in
outline, the constriction evident though not deep, the main axis of
the crown nearly perpendicular to sagittal line ; lower carnassial
without metaconid, the posterior lieel crossed by a longitudinal
trenchant ridge ; other teeth essentially as in Maries ; external
form slender, the muzzle obtuse, the ears low and rounded, the
legs short, the tail variable in length and in quality of hair, but
never so bushy as in Martes.

Bemarks. — The genus Mustela, the most widely distributed of
the strictly terrestrial Mustelinse, is also the richest in species.
These fall into three main groups, extreme members of which
might be regarded as generically distinct. The existence of
species with intermediate characters makes it impossible to
define these groups by anything more than their average
differences.* They are therefore here treated as sub-genera. All
three are represented in Europe.

KEY TO THE EUROPEAN FORMS OP MUSTELA.

Brain-case broad, the mastoid width decidedly
greater than distance between basion and
palation ; auditory bullae triangular in outline

(Sub-genus Pntorius, Polecats) M. putorms, p. 419.

Underfur usually buffy grey (Central Europe) M. p. putoHus^ p. 423.

Underfur usually yellowish (Southern Spain) il.p- aureolus^ p. 425.

Brain-case narrow, the mastoid width less than
or about equal to distance between basion and
palation ; auditory buUfe not triangular in
outline.
Rostrum flattened above; inner margins of
auditory bulte strongly divergent pos-
teriorly ; tail bushy ; habits semi-aquatic

(Sub-genus Lutreola, 'Minkii) M. lutnola, p. 415.

Rostrum convex above; inner margins of auditory
bullffi nearly parallel ; tail not bushy ; habits
terrestrial (Sub-genus J\lustda, Stoats and
Weasels).
Tail with conspicuous black terminal area in-
cluding more than pencil ; skull with ros-
trum usually not so wide as interorbital
region; condylobasal length of skull never
less than 43 -4 mm. in adult male, or than
35 '4 mm. in adult female; hind foot
usually more than 43 mm. in males and
34 mm. in females (Stoats).
Brown of sides noticeably encroaching on
yellowish or whitish of underparts, the
line of demarcation irregular ; upper lip
always brown M. Jiibcniica, ]). 398.

* At least so far as Lidrcola and true Mustela are concerned. Putorms
appears to be sharply circumscribed, so that it might readily be regarded
as a distinct genus (see remarks on p. 419).

MDSTELA 383

Brown of sides not encroaching on yellowish
or whitish of undcrparts, the line of
demarcation straight ; upper lip wholly

or partly white.. m. crminca, p. 385.

Kegion extending from front of rostrum to
and including postorbital processes
relatively broad and short ; size below
the maximum for the European races ;
condylobasal length of skull in adult
males, 43-4 to 48-6 mm., mandible in
adult males, 23 to 26-4 mm. (Scandi-
navin Peninsula, except extreme south

of Sweden) .,/. ,. ,,„„■ gg^

Kegion extondmg from front of rostrum
to and including postorbital processes
relatively narrow and long ; size maxi-
mum for the European races ; condy-
lobasal length of skull in adult males,
47 to 52-5 mm., mandible in adult
males, 25 to 30 mm.
Zygomatic arches broadly spreading
(ratio of zygomatic breadth to con-
dylobasal length about 59) ; posterior
extension of palate unusually short
and broad (Islands of Islay and Jura

Scotland) ,V. ,. ,,-,,-„^ 3,j7_

Zygomatic arches moderately spreading
(ratio of Z3'gomatic breadth to condy-
lobasal length about 54); posterior
extension of palate not unusually
short and broad.
White winter coat normally assumed ;
condylobasal length of skull in
adult male seldom over 50 mm. ;
in adult female seldom over
45 mm.; upper carnassial not
enlarged and thickened (Conti-
nental Europe from southern

,,r,--?^'^'^®" *° *^^ -"^'PS ^^^ Pyrenees) M. c. «jun, p 3S'J
White winter coat normally not
assumed ; condylobasal length of
skull in adult male often over
50 mm. ; in adult female often over
45 mm. ; upper carnassial enlarged

m •, •., , ^"tl tJiickened (Great Britain) il. c .tabilU^n Z2Q

iail without conspicuous black terminal area,
though a tuft of black or blackish hairs
may bo present in pencil ; skull with ros-
trum usually wider than interorlntal
region ; condylobasal length of skull, except
m specimens from the Mediterranean
region, often less than 40 mm. in males, and
34 mm. in females; hind foot -(except in
M. africana) usually less than 41 mm. in
males and 25 mm. in females (Weasels).
Size about as in the smaller stoats, condy-
lobasal length of skull in adult male,
48 to 50 mm., mandible in adult male
about 28 mm., its depth at front of car-
nassial, 4 • 5 to 5 mm. ; tail nearly one-half
as long as head and body (afiicanus
group).

384 CARNIVORA

Brown of sides noticeably encroaching on
yellowish or whitish of underparts,
the line of demarcation irregular

(Malta, Azores) M. afiicana, 412.

Brown of sides not encroaching on yellowish
or whitish of underparts, the line of

demarcation straight (Crete) .1/. fjalintliias, p. 414.

Size less than in the stoats, condylobasal
length of skull in adult male, 37 to
47 mm. ; mandible in adult male not
more than 24 mm., its depth at front
of carnassial less than 4-5 mm. (true

Weasels) M. nivalis, p 401.

Winter pelage often white ; size small, hind
foot of males, 29 to 34 mm. (Northern

and central Europe) M. a. nirallf, p. 402.

Winter pelage very rarely white ; size

medium or large, hind foot of males

usually 32 to 41 mm. (Mediterranean

region).

Skull of adult male not infrequently

44 to 4G'6 mm. in condylobasal

length ; hind foot of adult male 34 to

41 mm. ; colour rather dark (Central

Mediterranean region) M. n. boccamda, p. 405.

Skull of adult male seldom if ever exceed-
ing 44 mm. in condylobasal length ;
hind foot of adult male 30 to 37
mm. ; colour rather light (Iberian
Peninsula) M. n. iberka^ p. 407.

8ub-genus MUSTELA Linnanis.

1758. Mustcla Linnteus, Syst. Nat., i, 10th ed., p. 45 (Type by tautonymy

M. erminea LinniBUs).
1829. Ardocjalc Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierw., i,

p. 30 [erminea).
1829. Idis Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierw., i, p. 35

(vulgaris = nivalis). Not of Schinz, 1824-1828.
1841. Gale Wagner, Schreber's Saugthiere, Suppl., ii, p. 234 {uiilgaris =

nivalis).
1857. Fcctorius Blasius, Saugethiere Deutschlands, p. 219 (part).
18G5. Ncogale Gray, Proc. Zool. Soc, London, p. 114 {brasiliensis, aureo-

vcntris and xanthogenys).
1871. Mustclina, Bogdanow, Tpvj. Oiiuu Eciei-TB. Kasaiici;. .ViiiiBppc. I, Mem. i,

p. 167.
1899. Enmustcla Acloque, Faune de France, MammifcTcs, p. G2 {vulgaris

and enninca).
1911. Mitstda Thomas, Proc. Zool. Soc. London, p. 138, April, 1911.

Tijpe species. — 3Instcla eniiiiica Linn;eus.

Gcograpli'tcal (listrihutum. — Northern liemisphere, south in the
Old World to northern Africa, and in the New World to the
Andes ; in Europe west to Ireland.

Charartcrs. — Form very slender ; tail not bushy ; fur not
modified for aquatic life ; skull without noticeably projecting
mastoid processes ; auditory bulhe not triangular, the inner and
outer borders nearly parallel ; posterior boi'der of jmr^ shearing

MUSTELA 385

against anterior border of jj)«3 ; other small premolars not
capable of trenchant action.

RrmarJcs. — The sub-genus Mustela is the most widely-
distributed group in the genus. It is also the richest in species
and local races. About seventy of these are now recognized,
eleven of which occur in Europe.

MUSTELA ERMINEA Linnseus.
(Synonymy under subspecies.)

GeograpTiical disfrihufion. — Europe from the Arctic coa.st
to the Pyrenees and Alps, and from Great Britain eastward
into Asia.

Diagnosis. — Skull with narrow brain-case and slightly pro-
jecting, scarcely angular mastoid region, the mastoid breadth
less than distance from basion to palation ; rostral width across
canines usually less than interorbital width ; auditory bulhe much
longer than broad, nearly parallel sided, rounded posteriorly,
truncate anteriorly ; form very slender, the tail slender and not
bushy ; colour brown above, whitish below, the line of demarca-
tion straight ; a white winter pelage assumed in colder parts of
range ; tail always with black tip including more than terminal
pencil ; upper lip always at least partly white ; ear usually with
a whitish rim.

External characters. — General form long and slender, the
legs short, the body cylindrical, the neck long and nearly as
thick as body, tail considerably longer than outstretched hind
leg. Ear short but appearing distinctly above fur of head, its
outline evenly rounded, both outer and inner surface densely
clothed with short hair ; muzzle rather broad and short, the
nostril pad sharply defined, entirely naked, separated from upper
lip by a narrow hairy area. Palm and sole completely furred in
winter, the tubercles bare in summer ; on both palm and sole
there is a trilobed, heart-shaped tubercular mass at base of
median digits and a small round pad at base of inner digit ; on
palm there is an additional posterior rounded pad near wrist.
Fur dense and soft, the longest hairs on back about 10 mm.
in winter, less in summer ; tail rather closely haired, the pencil
full, usually wider than basal portion of tail, its longest hairs
about half as long as vertebra3 ; fur turning white in winter
except in warmer parts of range. MammsB : i 4 — 4 = 8.

Colour. — Upper parts and outer surface of legs yellowish
brown, usually a little darker along middle of back and on head,
the underfur and bases of the longer hairs much paler ; under-
parts, inner surface of legs and upper surface of feet whitish or
yellowish in strong contrast, the line of demarcation between

2 c

386

CAKNIYORA

the two colours straight and definite, extending along middle of
sides of body and neck and passing a little below ear and eye to
lower edge of muzzle pad, normally leaving upper lip white ; ear
brown, the rim whitish ; no dark spot behind angle of mouth ;
tail with terminal tifth and entire pencil black, elsewhere
concolor with back except for an ill-defined yellowish or whitish
median area which occasionally extends along under surface from
base toward or to black tip. In white winter pelage the entire
animal is white or whitish (often tinged with yellow), with the

Fig. 81.
Muatda erminea. Nat. size.

excej)tion of the black area of the tail, which remains as in the
dark coat.

Skull. — In general appearance the skull resembles that of
MtisfcJa jmtorins (p. 421). This is especially true of the dorsal
profile, the form of the palate, rostrum and interorbital region, in
all of which the differences between the two animals are very slight.
Breadth of rostrum over canines rarely equal to least interorbital
width. Position of auteorbital foramen as in M. 2)i(tor{us, but
orifice relatively larger, the width of plate sej^arating it from

MUSTELA 387

orbit distinctly less than that of foramen. Postorbital process
short, not evidently directed backward. Zygomata more widely
spreading than in M. putorins, strongly and evenly bowed when
viewed from the side, the orbital process barely indicated and
posterior widening absent. Owing to shortness of postorbital
process and virtual absence of orbital process of zygoma, the
orbit is less margined with bone than in M.puturius. Brain-case
longer and narrower than in Mustela putorius, the outline when
viewed from above elongate ovate not distorted by the presence
of conspicuously projecting mastoid regions ; sagittal crest slight,
even iii old individuals ; lambdoid crest moderately developed,
scarcely overhanging, the condyles usually visible from above.
Floor of brain-case nearly flat, though with the usual median
ridge and lateral and posterior depressions present. Auditory
bulla? moderately inflated throughout, about three-quarters as
wide as long, nearly parallel sided, squarely truncate in front,
rounded behind, the meatus not tubular ; anterior extremity of
bulla not in contact with haraular, and separated from foramen
ovale by a broad nearly tiat area, the length of which is
about one-half distance between foramina ; inner extremity of
glenoid surface marked oft" by a wide, shallow notch. Postoi;bital
region abruptly constricted, short, without indication of the
neck-like elongation which forms so conspicuous a feature of
the skull of M. putorius. Mandible with lower border slightly
convex, its posterior fourth somewhat elevated above general
outline ; angular process small but better developed than in
M. putorius.

Teeth. — Except for their smaller size the teeth are essentially
like those of Mustela putorius. Enamel of lower canine not
rugose. Second upper premolar and third lower premolar more
compressed than in the larger animal, and with base of cusp
shorter, so that there is a slight but evident concavity or
flattening at both anterior and posterior base. Upper carnassial
with inner lobe projecting more forward than in M. putorius, so
that its anterior border extends nearly to level of that of outer
margin of tooth. Upper molar and posterior lower molar as in
Mustela putorius, but rii^ with, metacone more reduced, and m.,
with crown area distinctly smaller than that of heel of carnassial.

Mustela ermine a erminea Linnaeus.

1758. [Mustela] erminea Linnajus, Syst. Nat., i, 10th ed., p. 46 (Sweden).
1792. M^ustela] erminea hyhcrna Kerr, Anim. Kingd., p. 181 (name applied

to the northern true ermine).
1816. ]S[\jistcla] hcrminea Oken, Lehrb. d. Naturgesch., iii, pt. 2, p. 1026

(Renaming of erminea).
1827. [Mustela. erminea'] 0 macnlata Billberg, Synopsis Faunae Scandinavise,

p. 8 (Scandinavia).
1857. Fcetorius erminea Elasius, Siiugethiero Deutschlands, p. 228 (part).

2 c 2

388

CARNIVORA

1877. Putortus erniinea Goues, Fur-bearing Animals, p. 109 (part).
1910. Putorius (Ictis) ermincus Trouessart, Faune Mamm. d'Europe, p. 78
(part).

Ti/2)e locaUty. — Upsala, Sweden.

Geogra^jliical distribution. — Scandinavian Peninsula, except
extreme south of Sweden ; eastern limits of range unknown.

Characters. — Region extending from front of rostrum to
and including postorbital processes relatively broad and short;
size below maximum for the European races : condylobasal length
of skull in adult male, 43 "4 to 48 • 6 mm. ; mandible in adult
male, "23 to 26*4 mm. ; basal portion of tail (in summer pelage)
usually lighter below than above.

Measurements. — Hind foot in each of two males from the
vicinity of Christiania, Norway, 43 mm. ; in a female from the
same region, 35 mm. Male and female from Nordre Fron, Lower
Gudbrandsdal : hind foot, 43 and 34 mm. Two males from
Lappmark, Sweden : hind foot, 44 and 46 mm. Adult female
from Upsala, Sweden : hind foot, 36 mm. Adult male from
Stockholm, Sweden : hind foot, 44 "6. For cranial measurements
see Table, p. 392.*

Specimens examined. — Twenty-five, from the following localities : —
Sweden: Karesuando, Lappmark, 1 (U.S.N.M.) ; Wilhelmina, Lapp-
mark, 1 (U.S.N.M.) ; Jemtland, 1 (U.S.N.M.) ; Upsala, 1 (U.S.N.M.) ;
Stockholm, 2 (U.S.N.M. and Stockholm).

NoEWAY : Aker, near Christiania, 2 ; Brekkebygden, Trondhjem, 1 ;
Nordre Fron, lower Gudbrandsdal, 2; Marestuen, Fillefjeld, Bergen, 1,
Egersund, Stavanger, 13.

HemarTcs. — The Scandinavian form of Mustela erminea is
distinguishable from the other European races by its slightly
smaller size and by the broader rostro-frontal region of the skull,
characters which appear to be fairly constant. Of the other
races it perhaps most closely resembles M. erminea ricinae from
the islands of Islay and Jura, west Scotland. The white winter
coat is normally assumed throughout the range of this form.

c5, 6 juv. Aker, Christiania, Nor- Christiania Museum 93. 3. 1. 2, 4.
way. (e).

5. Brekkebygden, Trond- Christiania Museum 93. 3. 1. 3.

hjem. (e).

6, ?. Nordre Fron, Gudbrands- Christiania Museum 93. 3. 1. 5-6.
dal. (e).

?. Marestuen, Fillefjeld, E. K. Alston (p). 79. 9. 25. 21.

2,.50O ft.

13skulls. Egersund, Stavanger. K. H. Schaanning (c). 11.6.3.22-34

* In eight skuUs of males from. Egersund, Stavanger, Norway (all
sutures closed, teeth slightly or not worn), received after the Tables were in
type, the averages and extremes are as follows : condylobasal length, 44-2
(43 -4-45 -2); zygomatic breadth, 24-2 (;23-6-25-4) ; mandible, 23-8 (23-0-
24 '8). Two females from the same locality: condylobasal length, 35'4
and 38-0; zygomatic breadth, 18-6 and 20-0; mandible, 18-2 and 19-2.

389

MUSTELA ERMINEA iESTIVA Kerr.

1792. Mlustela] crminca .rstiva Kerr, Anim. Kingd., p. 181 (Germany;

based ou Schreber's pi. 137a).
1820. Mustela erminea major Nilsson, Skand. Fauna, i, p. 34 (Carlskrone,

Blekingc, Sweden).
1857. Fatorius erminea Blasius, Siiugethiere Deutscblands, p. 228 (part).
1910. PiitoriiiH (Ictis) ermineiis Trouessart, Faune ]\Iamm. d'Europe, p. 78

(part).

Type locality. — Germany.

Gcoijraphical distrihniion. — Continental Eui-ope from southern
Sweden to the Alps and Pyrenees.

Characters. — Region extending from front of rostrum to and
including postorbital processes relatively narrow and long ; size
larger than in M. erminea erminea ; condylobasal length of skull
in adult male, 47 to 51 mm. ; mandible in adult male, 25 to
29 mm.

Measurements. — Two young adult males from Pic du Midi,
Hautes- Pyrenees, France : head and body, 235 and 240 ; tail,
82 and 85 ; hind foot, 44 and 44 ; ear from meatus, 17 and 17.
Three males from Strass, near Burgheim, Bavaria : head and
body, 265, 270 and 280 ; tail, 90, 100 and 102 ; hind foot, 44,
47 and 48. Two females from the same locality : head and body,
240 and 240 ; tail, 80 and 90 ; hind foot, 38 and 40. Average
and extremes of eight adult males from the vicinity of
St. Gallen, Switzerland: head and body, 271 (251-292); tail,
104 (94-111); hind foot, 47 (45-50). Three adult females
from the same region : head and body, 226 (218-242) ; tail, 89
(85-98); hind foot, 38-8 (37-6-41).

Specimens examined. — Seventy-nine, from tbe following localities : —

Denmark : No exact locality, 2 (skulls, Copenhagen) ; Amager, 1
(skull, Copenhagen) ; Faaborg, Fyn, 4 (skulls, Copenhagen) ; Vestervig,
Jutland, 1 (skull, Copenhagen).

Holland: Texel Island, 1.

France : Pic du Midi, Hautes-Pyr6n6es, 2 ; Manonville, Meurthe-et-
Moselle, 1.

Germany: Neustadt, Wied, 1 ; Ingelheim, Bheinhessen, 8 ; near Magde-
burg, 2; Bleiche, Saxony, 5; Moritzburg, Saxony, 1 (U.S.N.M.) ; Rudol-
stadt, Thiiringen, 3 ; INIarxheim, Bavaria, 4 ; Strass, near Burgheim,
Bavaria, 5 ; no exact locality, south Germany, G (skulls).

Austria-Hungary : Csallokoz-Somorja, Pressburg, Hungary, 1.

Switzerland : Frete-de-Sailles, Vaud, 1 (U.S.N.M.) ; Frutigen, Bern, 1
(U.S.N.M.); Rorbas, Bern, 1 (U.S.N.M.); Ziirich, 1 (U.S.N.M.); St.
Gallen, 4 (U.S.N.JI.) ; Degersheim, St. Gallen, 5 (U.S.N.M.) ; Rorschach,
St. Gallen, 1 (U.S.N.M.) ; Ziiberwangen, St. Gallen, 5 ; Untervatz, Grisons, 3
(B.M. and U.S.N.M.) ; Priittigau, Grisons, 1 (U.S.N.M.) : Vals, Grisons, 1
(U.S.N.M.); Obersaxen, Grisons, 1 (U.S.N.M.); Pontresina, Grisons, 1
(U.S.N.M.) ; Scanfs, Grisons, 1 (U.S.N.M.) ; Poschiavo, Grisons, 1 (U.S.N.M.);
Ems, Grisons, 2; Osogua, Ticiuo, 1 (U.S.N.M.).

Hemarks. — The ermine of central Europe differs from that of
Scandinavia in sligiitly greater size and in the narrower rostro-
frontal region of skull. The white winter pelage is normally
assumed throughout the ranse of the form.

390 CARNIVOEA

As shown by both external and cranial measurements the two
specimens from the Pic du Midi are unusually small. It is
possible that they represent a peculiar Pyrenean race.

5. Texel Id., Holland. J. L. Bonhote (c & 8. 10. 26. 1.

P).
2 S. Pic du Midi, Hautes Pyre- O. Thomas (p). 8. 9. 1. 52-53.

n6es, France. (A. Bobert.)

6. Manonville, Meurthe - et - Lord Lilford (p). 11.1.1.139

Moselle. (Lomont.)
?. Untervatz,Grisous,Switzer- 0. Thomas (p). 4. 4. 5. 33.

land. {E.H.ZoUikofcr.)
6,9. Ems, Grisons. (£. Jf. ;?oZiJ- 0. Thomas (p). 2.8.4.25-26.

kofer.)
i. Neustadt, Wied, Germany. Lord Lilford (p). 11. 1. 1. 138.
(Schneider.)
5 <?, 3 9. Ingelheim, Eheinhessen. C. Hilgert (c). 8. 11. 2. 22-28.

2. Magdeburg, Prussia. Dr. W. Wolterstorff 0. 2. 8. 6-7.

(P)-

4 i. Bleicbe, Saxony. [H. Tor- Lord Lilford (p). 11. 1. 1. 140-

mtz^ 143.

6. Marxbeim, Bavaria. Lord Lilford (p). 11.1.1.145.

{Wolterstorff.)

$. Strass, Burgbeim. {Kor- Lord Lilford (p). 11. 1. 1. 144.

hitz.)

6. South Germany. Dr. A. Giintber (c). 59.9.6.64-69.

1. Csallokoz - Somorja, Press- Budapest Museum 94.3.1.1.

burg, Austria-Hungary. (e).

MusTELA ERMiNEA STABiLis Barrett-Hamiltou.

1904. Putorius ermineiis stabilis Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., sin, p. 394, May, 1904.

1910. Putorius {Ictis) ermineus stabilis Trouessart, Faune Mamm. d'Europe,
p. 79.

Ti/jje locality. — Blandford, Dorset, England.

Geographical distribution. — Mainland of Great Britain.

Diagnosis. — Size slightly greater than in 31. erminea Kstiva :
condylobasal length of skull in adult male, 49 to 52 '4 mm.;
mandible in adult male, 27 to 30 mm. ; teeth usually larger than
in the continental races, a peculiarity especially noticeable in the
upper carnassial ; colour averaging slightly darker above and
less strongly yellow below than in the continental forms (though
in extreme instances the underparta are between buff and straw-
yellow, quite as in the brightest continental specimens) ; tail
usually without distinct lighter area below.

Measurements. — ■ External measurements of type (adult
female) : head and body, 244 ; tail, 105 ; hind foot, 43 ; ear from
meatus, 22. Another adult female from the type locality : head
and body, 245 ; tail, 102; hind foot, 43; ear from meatus, 21.
Two adult males from Banstead, Surrey : head and body, 270
and 278 ; tail, 119 and 119 ; hind foot, 47 and 49. Two adult
males from Wales : head and body, 274 and 280; tail, 120 and

MUSTELA 391

120; hind foot, 48 and 48-6. Adult male from Farr, Daviot,
Inverness : head and body, 254; tail, 110; hind foot, 48 "5.

Specimens exanmicd. — Seventy-four, from the following localities : —

Scotland : Thurso, Caithness, 1 ; Suthorlaudshire, 1 ; Cromarty, 4
Annadale, Skye, 2 ; Farr, Daviot, Inverness, 1 (Wilson) ; Stockbriggs
Lanark, 9; Wyseley, Dumfries, 2; Ecclefechan, Dumfries, 1 (U.S.N.M.).

Wales: North Wales, no exact locality, 1; Nannerch, Flintshire, 2
Vaynol, Carnarvonshire, 2 ; Usk, Monmouthshire, 2 ; St. Brides, Pembroke
shire, 1 ; St. Fagan's, Cardiff.

England: Riding-Mill-ou-Tyne, Northumberland, 1; Westmoreland, 1
Doncaster, Yorkshire, 2 ; Leeds, Yorkshire, 1 (U.S.N.JI.) ; Gainsborough
Lincolnshire, 1 ; Derbyshire, 1 ; Sandringham, Norfolk, :3 ; Lowestoft
Suffolk, 1 ; Bury St. Edmunds, Suffolk, 1 ; Friswell, Cambridgeshire, 8
Rugby, Warwickshire, 1 ; Graftonbury, Hereford, 3 ; Triug, Hertford, 2
Pelden, Hertfordshire, 1; Banstead, Surrey, 2; Laseley Park, Guildford
Surrey, 1 ; Buckland, Somerset, 2 ; Blandford, Dorset, .3 ; Eversley
Hampshire, 1; Whitechurch, Hampshire, 1; Selborne, Hampshire, 1
Staplehurst, Kent, 1 ; Horsham, Sussex, 2 ; Mayfield, Sussex, 3.

Bemarks. — In the British stoat the change to the white
winter coat does not take place so regularly and completely as in
the continental forms. This is particularly true in central and
southern England. Apart from this character, of problematical
value, the animal is distinguishable by its heavy teeth. A few
small though apparently well developed skulls from the north
of Scotland indicate the possible existence there of a local form
somewhat resembling true erminea.

9. Thurso, Caithness, Scot- W. R. Sherrin (p). 9. 1. 9. 1.
land (Mrs. J. Edis).

3 9. Cromarty. W.R.Ogilvie-Graut 11.1.3.175-177.

(c & p).

<5, 9. Annadale, Skye. Dr. Hastings (c&p). 11.1.3.395-396

<^. Sutherlandshire. W. Paterson (c&p). 8.11.26.1.

3 6, St. Stockbriggs, Lanarkshire. B. R. Alston (c & p). 79. 9. 25. 16, IS,

20.

3 6, 9. Stockbriggs, Lanarkshire. E. R. Alston (c & p). 79. 9. 25. 14, 15,

17, 19.

9. Stockbriggs, Lanarkshire. E. R. Alston (c & p). 79. 9. 25. 84.

(5,9. Naunorch, Flintshire, A.Richardson 11.1.3.192-193.

Wales. (c&p).

2 9. Vaynol, Carnarvonshire. J. E. Harting (p). 11. 1. 3.194-195.

2 6. Usk, Monmouthshire. J. S. Phillips (c & p). 87. 8. 6. 1-2.

St. Brides, Pembroke- Hon. C. Edwardes 90. 12. 5. 1.

1. St. Pagans, Cardiff. J. Cording (c). 90. 6. 19. 1.

st. Derbyshire, England. Index iluseum. 94. 4. 8. 1.

St. Sandringham, Norfolk. H.M. King Edward 96. 4. 13. 1.

VII. (p).

2 St. Sandringham, Norfolk. H.M. King Edward 96. 11. 24. 1-2.

VII. (p).

9 St. Lowestoft, Suffolk. F. S. Worthington 98. 2. 15. 1.

(c & p).

St. Bury St. Edmunds, Suf- T. Harcourt-Powell 87. 2. 28. 1.
folk. (c & p).

3<J,4 9. Friswell, Cambridgeshire. W. Farren (c). 4. 5. 3. 1-8.

i. Rugby, Warwickshire. E. E. Austen (c&p). 11.1.3.181.

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MUSTELA 397

6,2 9. Graftonbury, Hereford- W. E. de Winton (p). 11.1.3.178-180.

shire.
2 9. Tring, Hertfordshire. Hon. N. C. Boths- 11.1.3.182-183.

child (p).

6. Felden, Hertfordshire. 11. 1. 3. 184.

1. Nr. Hounslow, Middle- W. Phillips (p). 60. 1. 23. 2.

sex.
9. Guildford, Surrey. G.Dalgleish (c &p). 4.4.3.1.

6 St. Horsham, Sussex. Sir E. G. Loder, 6. 7. 14. 1.

Bart, (c & p).
6,9. Mayfield, Sussex. C. H.B. Grant (c&p). 11.1.3.186-187.

6. Selborne, Hampshire. Dr. B. Bowdler 2. 11. 29. 1.

Sharpe (c & p).
9. Whitchurch, Hampshire. W. E. de Winton 11. 1. 3. 185.

(c & p).
6, 9. Buckland, Somerset. B. Hooper (c & p). 11. 1. 3. 190-191.

9. Blandford, Dorset. J. C. ManseU Pley- 98. 5. 13. 2.

dell (c & p). (Type of subspecies.)
2 9. Blandford, Dorset. W. T. Blanford 11.1.3.188-189.

(c & p).

MuSTELA EEMINEA EICIN^ Miller.

1904. Puforius erminea Barrett-Hamilton, Aim. Scottish Nat. Hist., v,

p. 203, October, 1904.
1907. Putorius erminea ricime Miller, Ann. and Mag. Nat. Hist., 7th ser.,

XX, p. 395, November, 1907.
1910. Putorius ermineus richix Trouessart, Faune Mamm. d'Europe, p. 80.

Type locality. — Islay House, Islay Island, Scotland.

Geographical distribution. — Islands of Islay and Jura, off coast
of south-west Scotland.

Characters. — Size a little less than in M. erminea stahilis and
about equal to that of M. e. aestiva : condylobasal length of skull
in adult male, 47 to 50 mm. ; mandible in adult male, 27 to
29 mm. ; zygomatic arches unusually wide-spreading ; posterior
extension of palate relatively wider than in any of the other
known forms.

Measurements. — External measurements of type (adult male) :
head and body, 254 ; tail, 105 ; hind foot, 43 ; ear from meatus,
22. Two other males from the type locality : head and bodv,
270 and 270 ; tail, 111 and 114 ; hind foot, 47 and 46. Two
adult females from the type locality : head and body, 23 1 and
234 ; tail, 95 and 105 ; hind foot, 38 and 39. Two adult males
from Jura : head and body, 220 and 225; tail, 120 and 125;
hind foot, 45 and 46 "5.

Specimens exatnined. — Thirteen, six from Jura and seven from Islay
(B.!M. and Cambridge).

Itemnr1c.<>. — While readily distinguishable from the large true
atubilis of England, the Islay and Jura stoat will probably prove
to be more nearly related to the smaller form occurring on the

398

CARNIVORA

mainland of Scotland, when the status of the Scotch animal is
more clearly understood.

1. Islay, Scotland. H. Morrison (c & p). 7. 10. 9. 1.

(Type of subspecies.)
6, 2 ?. Islay, Scotland. H. Morrison (c & p). 11. 1. 3. 196-198.

MUSTELA HIBERNICA Thomas and Barrett-Hamilton.

1895. Putorius Jdbcrnicus Thomas and Barrett-Hamilton, The Zoologist,

.3rd ser., xix, p. 12.5, April, 1895 (Ireland).
1895. Putorius hibernicus Thomas, The Zoologist, 3rd ser., xix, p. 226,

June, 1895 (Isle of Man).
1900. [Mustdrc] hihernica Schulze, Zeitschr. fiir Naturwissensch., lxxiii,

p. 221.
1910. Putorius (Ictis) hibernicus Trouessart, Fauna Mamm. d'Europe, p. 80.

Ty2)e locality. — Enniskillen, Co. Fermanagh, Ireland.

Oeogra])hiccd distribution. — Ireland and the Isle of ^lan.

Diagnosis. — ^Like Mustela erminea but slightly smaller (hind
foot in adult male rarely attaining 50 mm. ; condylobasal length
of skull in adult male, 44 to 50 mm.) ; ear and upper lip without
whitish edging ; brown of sides of body usually encroaching on
yellowish or whitish of underparts, frequently so much so as
to extend completely across throat and middle of belly, the
line of demarcation, except in rare instances, very irregular.

External characters. — The general external characters, includ-
ing the proportion of tail to head and body, and the long,
full pencil, are as in Mustela erminea ; manmipe usually more
numerous : a2 — 2, « 3 — 3 or 4 — 4 = 10 or 12.

Colour. — Except in the rare instances in which a partial or
complete white coat is assumed in winter, the actual colour is
very constant throughout the year, and such variation as there
is appears to be due chiefly to the greater or less abundance of
the long hairs on back. These hairs are a dark glossy brown
very nearly the bistre of Ridgway. The hairs of the dense
underfur are smoke-grey at base, changing to a yellowish broccoli-
brown distally. From the varjang coralnnations of these elements
diflerent shades of bi'own result which range from a yellowish
bistre toward isabella-colour, the bistre usually more pure along
median dorsal region and on head, the isabella-colour on sides,
legs and tail. Upper lij) and entire ear concolor with dark
area. No dark spot behind angle of mouth. Feet usually dark
but frequently varied with whitish, this colour appearing at tips
of toes and spreading upwards. Tail without evident lighter
area on under side. Underparts varying from a creamy white
to a pale straw-yellow. The line of demarcation between the
dark and light areas is very variable in both form and position.
In the extreme development of the light area the line of demarca-
tion extends along sides in the same position as in 3lusicla
erminea, except that it passes somewhat further below ear and
eye and ends at or just above angle of mouth. In the extreme

399

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400

CARNn'ORA

extension of the dark area the brown forms a band 20 mm. wide
across throat just in front of fore legs, and spreads over whole
median region of belly, so that the white is confined to three
patches, one extending from chin to throat band, another occupy-
ing chest between front legs and spreading on inner side of
forearm, and the third in region between hind legs and spreading
on inner surface of thighs. Between these extremes every
gradation may be found, including the most irregular arrange-
ments of isolated dark spots in the regions where the dark cross
bands occur.

Slull and teeth. — Except for their slightly smaller size the
skull and teeth are indistinguishable from those of Miistcia
erminea. The difference is, however, particularly well marked as
compared with the large British M. erminea stahilis, the nearest
geographical ally.

3Ieasure)aents. — External measurements of type (adult male) :
head and body, 228 ; tail, 88 ; hind foot, 42 ; ear from meatus,
21. Average and extremes of four males fi-om Cappagh, Water-
ford : head and body, 271 (256-283) ; tail. 111 (103-117) ; hind
foot, 48 '2 (46-51). Average and extremes of three females
from Powerscourt, Co. Wicklow : head and body, 209 (205-221) ;
tail, 78 (77-81) ; hind foot, 37 "6 (36-40). For cranial measure-
ments see Table, p. 399.

Specimens examined. — Forty-four, from the following localities : —
Ireland : Carrick, Donegal, 1 ; Colebrooke, Fermanagh, 8 ; Enniskillen,
Fermanagh, 2 ; Clandeboj'e, Down, 1 ; Glaslough, Monaghan, 1 ; Board-
mills, Down, 2 ; Castle Hamilton, Cavau, 1 ; ]Mountainstown, Meath, 3 ;
Clonlarock, Galway, 1 ; Woodfair, Galway, 1; Carna, Galway, 1 ; Temple-
more, Tipperary, 1 ; Geashill, Kings, 1 ; Powerscourt, Wicklow, 4 ;
Bagenalstown, "Carlow, 1 ; New Ross, Wexford, 4 ; Arthurstown, Wex-
ford, 2 ; Cappagh, Waterford, 4; Lismore, Waterford, 1.

Isle or Man : Lewaig, Ramsey, 1 ; Santon, 2 ; Tholt-y-Will, Snaefell, 1.

Remarks. — The Irish stoat is strikingly distinct from the
other European species of the sub-genus Mustela, though super-
ficially resembling certain North American members of the
group. It is at once recognizable by the combination of black-
tipped, heavily pencilled tail with entirely dark ear and upper
lip. It is also the only European stoat in which the line of
demarcation on sides of body is irregular and in which the dark
colour of sides tends to invade the light ventral area. In
exceptional instances* the colour pattern of body is like that of
Mustela erminea. While the colouring of the body and head
suggests that of Mustela nivalis, the animal is evidently a true
stoat, as shown by the heavily-pencilled, black-tipped tail and
the form of the skull.

<5 St. Carrick, Donegal, Ireland. Hon. N. C. Roths- 0.5.17.1.

child (P).
3i. Colebrooke, Fermanagh. Sir D. Brooke (c & p). 11. 1. 3. 199-201.

In four of the forty-one skins examined.

MUSTELA 401

S. Enniskillen, Fermanagh. J. E. Harting (c&p). 95.4.5.1.

(Type of species.)
£ St. Clandeboye, Down. Hon. N. C. Roths- 0. 5. 17. 2.

child (p).
<J. Carna, Galway. Col. J. W. Yerbury 93. 1. 6. 1.

(c & p).
St. Templemorc, Tipperary. E. Lynan (c & p). 99. 7. 3. 1.
2 6. Arthurstown, Wexford. G. Barrett-Hamilton 11. 1. 3. 202-203.

(c & P).
9. Ramsay, Isle of Man. P. M. G. Kermode 95. 5. 30. 1.

(c & p).
9. Suaefell, Isle of Man. P. M. C. Kermodo 11. 1. 3. 204.

(c & p).
6, 1 al. Sauton, Isle of Man. J. 0. Bacon (c & p). 1. 5 12. 1-2.

MUSTELA NIVALIS Linnaeus.

(Synonymy under subspecies.)

Geographical 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.

Diaijiiosis. — Size usually less than in Mustela 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 vertebi'ie ; 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 vertebrte ; palms and soles hairy, the
pads bare in summer in northern fonns, 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. Mamma? : i i — i = 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

CARNIVORA

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 che(;k, usually present behind angle of
mouth ; ear brown like surrounding parts, 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 j)elage 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.

ShiU and teeth. — The skull resembles that of Mustela ermhiea
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 upward, its upper border
flattened or sometimes with a distinct median
concavity and posterior widening as in
M. putorius. Teeth not distinguishable from
those of Mustela erminea except by their
smaller size.

Remarks. — Although the most variable of
the European Mustelidse this species is always
readily distinguishable externally from the
other memljers of the genus Mustela with
which it is associated by the short thin pencil.
The discrepancy in width between the rostrum and interorbital
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

Mustela nivalis (male),
Nat. size.

MusTKLA NIVALIS NIVALIS Linnwus.

1766. [Mustela'] nivalis Liunseus, Syst. Nat., i, 12th ed., p. 69.

1777. [Mustela'] vulgaris Erxleben, Syst. Regni Anim., i, p. 471

(Temperate Europe).
1811. Mustela gale Pallas, Zoogr. Rosso-Asiat., p. 91 (Renaming of

vulgaris).
1820. Mustela minor Nilsson, Skand. Fauna, i,p. 35 (Renaming of nivalis).
1853. PUdorius] mimitus Pomel, Catal. ]M6tli. et Descr. Vert. Foss. Loire,

p. 51 (Neighbourhood of Paris, France).

MDSTELA 403

1857. Fcelorius vjilgaris Blasius, Saugethiere Deulschlands, p. 231.

1869. Faforius piisillus Patio, Paune Vert. Suisse, I, p. 332 (Not of Audubon

and Bachman ; substitute for vulgaris).
1895. Putorius nivalis Tbomas, The Zoologist, 3rd ser., xix, p. 177,

May, 1895.
1900. Putorius nivalis typicus Barrett-Hamilton, Ann. and IMag. 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. Plntorius'] nivalis var. monticola Cavazza, Ricerche sui " Putorius

nivalis " e sui "Putorius ermineus" d' Italia, Bologna, p. 37 (High

valleys of the Alps).
1910. Putorius (Icfis) nivalis, P. (J.) nivalis vulgaris and P. (I.) vulgaris

minutus Trouessart, Faune Mamm. d'Europe, pp. 81-82.

Type locality. — Province of Vesterbotten, Sweden.

Geographical distribution. — 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) ;
Cortachy, Forfar, 1 (Wilson) ; Stockbriggs, Lanark, 3 ; Lamancha, Peebles,
1 ; Ecclefechan, Dumfries, 2, Wyseby, Dumfries, 1 ; Jardiue 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 Walden,
Essex, 4; Southall, Middlesex, 1; Eashiug, 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 Christiania, 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, Pyren^s-Orientales, 4;
Ax-les-Thermes, Ari^ge, 1 ; Barcelonnette, Basses- Alpes, 1.

Germany: Brunswick, 1 (U.S.N.M.); Konigsberg, 8 (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).

Austria-Hungary : Haida, Arva, Bohemia, 1 ; Hainspach, Bohemia, 4
(U.S.N.M.) ; Csall6kdz-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, St. Gallen, 1 (U.S.N.M.) ; Watt-
wil, St. Gallen, 1 (U.S.N.M.) ; Weiern, St. Gallen, 1 (U.S.N.M.) ; Ziiber-
waugen, 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 suthcient to show
whether or not the northern animals are all referable to a
single form.

S.

3<5.
£.

Thurso, Caithness, Scot-
land.
Cromarty.

Mrs. J. Edis (p).

11. 1. 3. 300.

2 i, ?.

i.

4(5.

6.

S.
6, 9.

2 St.

2.

Lamancha, Peebles.

Stockbriggs, Lanark-
shire.

Aberia, Merionethshire,
Wales.

Pembrokeshire.

G. St. Quentin

(c & p).
J. L. Bonliote (c & p).
E. R. Alston (c & p).

11. 1. 3. 20.5,

207, 432.
11. 1. 3. 433.
79. 9. 25. 23,
85-86.
G. H. Caton Haigh 11. 1. 3. 443.

(c & p).
G. H. Mills (c & p). 11. 1. 3. 444-
447.
Grimsby, Lincolnshire, G. H. Caton Haigh 11. 1. 3. 434.

England. (c & p).

Norfolk. Dr. C. Hose (c & p). 11. 1. 3. 208.

Sandringham, Norfolk. H.M. King Edward 96. 4. 13. 2. 3.

VII. (p).
Sandringham, Norfolk. H.M. King Edward 96.4.28.14-15.

VII. (p).
Suffolk. Col. E. S. Butler (p). 94. 1. 6. 6-7.

6. Histon, Cambridgeshire. Dr. S. F. Harmer 11. 1. 3. 206.

(c & p).
9 (albino) Soham Fen, Cambridge- F. Bond (p). 90. G. 21. 1.

shire.

2 (5, 2 9. Saffron Walden, Essex.
6. Southall, Middlesex.

S. Froyle, Hampshire.

A. Wright (c & p).

R. C. W^roughton

(c & p).
W. R. Ogilvie-Grant

(c & p).
W. L. S. Loat(c&p).
Christiania Museum

9. Stratton, Cornwall,

juv. Asker, Christiania, Nor

way. (e).

2 i. Guines, Pas-de-Calais, O. Thomas (c & p).
France.
S. L u c h o n. Haute- O. Thomas (p).

Garonne. (-4. Robert.)

5. L e g u e V i n, H a u t e- O. Thomas (p).

Garonne. (A. Hohert.)

6. Ax-les-Thertnes, Ariege. V. Builles (p).

2 (5, 1 9. Port6, Pyrenees - Orien- Cr. S. Miller (c).
tales.
9. Barcelonnette, Basses- 0. Thomas (p).
Alpes. (C. Mottaz.)

11. 1. 3. 4.3G-

4.39.
11. 1. 3. 440.

11. 1. 3. 441.

11. 1. 3. 442.
95. 5. 29. 1.

94. G. 6. 10-11.

6. 4. 1. 36.

G. 4. 1. 37.

8. 3. 27. 8.
8.8.4.16G-168.

8. 8. 10. 44.

MUSTELA 405

6 juv. Strass, Burgbeim, Ba- Lord Lilford (p). 11. 1. 1. 96.

varia. (KiJvhitz.)

3. Ingelheim, Rheinhes- C. Hilgert (c). 8. 11. 2. 29-31.

sen, Germany.

4. South Germany. Dr. A. Giinther (c). 59. 9. G. 71-73,

122.
3. Csallokoz - Somorja, Budapest Museum 94.3.1.2-4.

Pressburg, 400 ft. (e).
Hungary.
3d,?. Hatszeg, Hunyad, Trau- 0. G. Danford (c). 3.2.2.17-19.

sylvauia. 3. 11. 8. 19.

MusTELA NIVALIS BOCCAMELA Bechstein.

1801. Mustela boccamcla Bechstein, Gemeinn. Naturgesch, Deutsehlands,

I, 2nd ed., p. 819 (Sardinia).
1869. Putorius vulgaris var. mcridionalis Costa, Annuario del Mus. Zool.

della R. Univ. di Napoli, 1865, p. 40 (Southern Italy).
1900. Putorius nivalis 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 Mag. 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) dombroioskii Matschie, Sitz.-Ber. GeseUsch. Natur-

forsch. Freunde, Berlin, p. 231 (Siulnita, Roumania).

1905. [Fcetorius piisillus] major Fatio, Arch. Soi. Phys. et Nat. Geneve,
4th ser., xix, p. 512, May 15, 1906 (Poschiavo, Grisons, Switzer-
land ; see Jlottaz, Bull. Soc. Zool., Geneve, i, p. 169, November 15,
1908). Type in Mottaz collection.

1910. Putorius (Ictis) boccamela, P. (J.) nivalis meridionalis, P. (I.) nivalis
dombroioskii, and P. (J.) nivalis siculus, Trouessart, Faune Mamm.
d'Europe, pp. 81, 83.

Type locality. — Sardinia.

Geographical distrihiction. — Italy aucl coast of France as far
as the Department of Var ; islands of Sicily, Malta and Sardinia :
Corsica 1

Diagnosis. — Similar to Mustela nivalis nivalis, but decidedly
lai'ger, 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 ; hintl 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 : —

Feance. — Valescure, Var, 2; Agay, Var, 1.

Switzerland : Somvico, Ticino, 1 (U.S.N.M.) ; Bogno, Ticino, 1
(U.S.N.M.); Buggiolo, Ticino, 1 (U.S.N.M.); Sau Lucio, Ticino, 1;
Poschiavo, Grisous, 4 (U.S.N.M. and Mottaz, including type of major).
These Swiss specimens are intermediate between boccamcla 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.

RoDMANiA ; 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
hoccamda are strikingly different from the animal occurring
north of the Alps.

6.

Valescure, Var, France.

G. S. Miller (c).

8. 8. 4. 170.

6.

Agay, Var.

G. S. Miller (c).

8. 8. 4. 169.

9.

St. Lucio, Ticino, Switzer-
land. {E. H. Zollikofer.)

0. Thomas (p).

2. 8. 4. 27.

i.

Turin, Italy.

E. Cavendish
Taylor (p).

5. 5. 6. 11.

6.

Genoa, Liguria.

Lord Lilford (p).

95. 11. 4. 1.

2 6,19.

Verona (Conte degli

Oddi)

0. Thomas (p).

99. 11. 11. 1-3.

(99

11.11.1. Tiipeoiita

iciis Barr.-Ham.)

6.

Balestrate, Sicily.

J.I.S.Whitaker(p).

95. 3. 4. 4.

4 6.

Palermo.

J. I.S.Whitaker(p).

98. 10. 6. 3.
95. 3. 4. 1-3.

2 6, 19.

IMarsala.

J.I.S.Whitaker(p).
(95. 3. 4. 5. Type of

95. 3. 4. 5-7.
siculus B.-Ham.)

6, 2 al.

Cagliari, Sardinia.

E. N. Buxton (p).

95. 4. 16. 1-3.

3al.

Sardinia.

Genoa Museum (p).

86. 11. 3. 1-3.

Sardinia.

Purchased (Linnsea,
Berlin).

86. 7. 10. 2.

juv.

]\Ialta.

C. A. Wright (p).

94. 11. 26. 1.

9.

:Malta. (Micallcf.)

C. A. Wright (p).

95. 1. 2. 1.

3 <5, 1 9.

Ghallis.

C. A. Wright (p).

7. 7. C. 1-4.

9.

Bingemma Port.

C. A. Wright (r).

7. 7. 6. 5.

* studied in much detail by Cavazza, Ricerche sui " Putorius nivalis "
e sui " Putorius ermineus " d' Italia (Bologna, Zanichelli), 1908, Studien
fiber die in Italien vorkommenden Wleselarten der Untergattung Arctogale
(Zool. Anzeiger xxxiv, pp. 582-603), 1909, SuUo Donnole e suU' Ermellino
in Italia (Boll. Soc. Zool. Ital. xviii), 1909, and Contributo alia conoscenza
della vita e dellc abitudini della Donnola, Putorius nivalis, Linn. (ibid,
xix, pp. 65-82), 1910.

407

MusTELA NIVALIS IBERICA BaiTett-Hauiilton.

1900. Putoriiis nivalis ibericus Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., v, p. 45, January, 1900. Type in British Museum.
1905. Putorius hibericus Seabra, Bol. da DirecQ. Gen. da Agricultura,

Lisboa, viii, no. 2, p. 69.
1910. Putornis nivalis iberica Trouessart, Faune Mamm. d'Europe, p. 83.

Tyjje locality. — Seville, Spain.

Geographical distrihution. — Iberian Peninsula and Balearic
Islands.

Diagnosis. — Intermediate in size between Mustela nivalis
nivalis and 31. 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 buify clay-colour,
not so dark as in the other races.

Measurements. — Adult male and female from the neighbour-
hood of Silos, Burgos, Spain: head 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 fi'om 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): hind foot (dry), 31 •4.
For cranial measurements see Table, p. 411.

Specimens examined. — Twenty-seven, from the following localities : —
Spain : Paj4res, Leon, 1 ; Panticosa, Huesca, 1 ; Arrechavaleta,
Vitoria, 2 ; near Burgos, 1 ; Silos, Burgos, 1 ; Castrillo de la Eeina,
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.
Portugal : Alcochete, 1.

Bemarks. — The Iberian weasel appears to be more nearly
related to Mustela nivalis nivalis than to 31. n. hoccamela. It is
distinguishable from both of these by its paler colour.

9.

Pajares, Leon, Spain.

{N. Gmizalcz.)

0. Thomas (p).

8. 2. 9. 54.

6.

Panticosa, Huesca.

{N. Gonzalez.)

0. Thomas (p).

8. 2. 9. 58.

2 9.

Arrechavaleta, Vitoria.
(A^. Gonzalez.)

0. Thomas (p).

8. 2. 9. 59-60.

i, 9 juv.

Burgos.

G. S. Miller (c).

8. 8. 4. 50-51.

1 al.

La Granja, Segovia.

M. de la Escalera (c).

8. 7. 30. 8.

S, 9.

Villalba, Madrid.

{N. Gonzalez.)

0. Thomas (p).

8. 2. 9. 56-57.

i.

Barracas, Castellon.

{N. Gonzalez.)

0. Thomas (p).

8. 2. 9. 55.

3 6.

Dehesa de Valencia.

[N. Gonzalez.)

0. Thomas (p).

8. 2. 9. 51-53.

6,9.

Bormujo, Seville.

(Dr. A. Ruiz.)

Lord Lilford (p).

95. 9. 4. 12-13.

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CARNIVOEA

?. Seville. {Dr. A. Ruiz.) Lord Lilford (p). 95. 3. 3. 10.

{Ty2Je of sub-species.)

S. Jerez. A. Chapman (c & p). 9. 4. 24. 1.

1. Alcochete, Portugal. Lisbon Museum (e). 3.11.13.1.
<5. Muro, Majorca, Balearic 0. Thomas (p). 1. 6. 1. 5.

Islands. {M. Biutort.)

2. Inca, Majorca. O. 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. Mustela 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 nivalis africanus Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., v, p. 47, January, 1900.
1908. ? Plutorius] nivalis var. corsicanus Cavazza, Ricerche sui "Putorius

nivalis " e sui " Putorius ermineus " d' Italia, Bologna, p. 37

(Corsica).
1910. Putorius (Ictis) nivalis subpalmatus Trouessart, Faune Mamm.

d'Europe, p. 85.

Type locaJittj. — iSaid to be Africa (no exact locality stated),
but more probably the Azores, as the type came to Paris through
the museum of Lisbon, Portugal.

Geographical distribution. — Egypt, Malta, San Thome, Azores.

Diagnosis. — Like Mustela nivalis hoccaniela 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 brown of sides usually spreading on underparts, the
line of demarcation very irregular.

Colour. — The colour does not differ noticeably from that of
Mustela nivalis hoccatnela. 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-buft'or bufly
white, the light area on chest and belly much encroached on by
brown of sides, which occasi<mally pa.sses 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
boccamela 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 3T. erminea ; but the much bi'oader
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. boccamela except
that the carnassial is more robust and the small premolars in

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414

CARNIVORA

both jaws tend to be more crowded, so that their crowns are
usually set more obliquely to axis of tooth-row.

Measurement a. — 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 Thom6, 1 ; Terceira, Azores, 2 ; St. Michaels, Azores, 1.

BemarJcs. — This African species probably owes its presence in
Malta and on the Azores to introduction by man. So far as can
he judged from the description, the Corsican weasel to which
Cavazza has applied tlie name cursicanus, is the same animal.

1. Malta. C. A. Wright (p). 75. 4. 6. 1.

1. San Thomfi. Hon. W. Rothschild 4. 1. 1. 5.

Terceira, Azores. Hon. W. Rothschild 3. G. 5. 25.

{W. R. Ogilvic-Grant.) (p).
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.

190G. Putorins nivalis galintliias Bate, Proc. Zool. Soc. London, 1905, ii,

p. 319, April 5, 1906.
1910. Putorius (Ictis) nivalis galintliias Trouessart, Faune Mamm.

d'Europe, p. 14.

Type locality. — Crete.

Oeograplncal 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.

<?. 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 Wagner.

lS-41. Lutreola Wagner, Schreber's Siiugtlnere, Suppl., ii, p. 239 (lutreola).
1843. Vison Gray, List. Spec. Mamm. Brit. ]\Ius., p. 04 {lyison).
1857. Fa'torius Blasius, Siiugethiere Deutsohlands, p. 219 (part).
1871. Hydromnstela Bogdauow, Ipyj. OOin. EcrecrB. l\';i;!iiiici;. ,Viiiibi'|ii-. I,
'Mem. I, p. 167.

Tifpe species. — Mustela lutreola Linnteus.

Geographical distribution. — Northern portion of northern
hemisphere from eastern Germany to the Atlantic coast of North
America.

Characters. — Form moderately slender ; tail bushy ; fur modi-
fied for aquatic life ; skull without noticeably projecting mastoid
processes ; auditory bulL-e sub-triangular ; premolars more prehen-
sive than in Putorius ; inner lobe of upper carnassial functioning
against apex of ptii^ ; anterior border of pm^ shearing against pos-
terior border oipm^ ; point of pm- opposed to posterior border of jjmJj-

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 sihirica, M. cani-
ijula and others. So complete is this intergradation that it may
well be (juestioned vvhether the name Lutreola should be allowed
to remain in use. The only typical Old World species ranges
westward into south-western France.

MUSTELA LUTREOLA Linmeus.

1766. IMusteUC] lutreola Linueeus, Syst. Nat. i, 12th ed., p. 66 (Finland).
1777. [Lutra] minor Erxleben, Syst. Regni Anim., i, p. 451 (Renaming of

lutreola).
1792. M\iistela'\ Lutra ftilva Kerr, Anim. Kingd., p. 173 (Renaming of

lutreola).
1839. ? [Mustela lutreola^ var. alba De S61ys-Longcliamps, Etudes de

Micromamm., p. 46 (nomcn nudum).
1857. Foitorius lutreola Blasius, Siiugethiere Deutschlands, p. 234.
1863. ? Putorius alpinus Ogerien, Hist. Nat. du Jura, iii, p. 59 (highest

portions of the Jura).
1879. Lutreola exi,rop;va Homeyer, Zool. Garten, xxvi, p. 184, June, 1879

(Substitute for lutreola).
1910. Putorius (Tjutreola) lutreola Trouessart, Faune Mamm. d'Europe,

p. 75.

Type 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.

Dia;/Ho.s7'.s-.- -General 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

CARNIYOEA

400 mm., condylobasal length of skull ranging about from 57
to 65 mm.

Colour. — Back, sides and underparts a rich glossy dark brown,
nearly 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.

FIO. S3.
Mustela lutreola. Kat. size.

Skull. — The skull is about as large as in females of Mustela
pittorius. In form it is narrower and less deepened than in
M. ijiitorius, especially in interorbital region, the dorsal profile

417

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Number. Sex.

2975 Lataste 6
3270 „ i
3G78 ,, 6
3679 ,, 9

Locality.

France : Vinsac, Gironde

Laval, Mayenne

2 E

418

CARNIVORA

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 apprecialjly greater ; orbit larger and more widely
open posteriorly than in M. imtorius, 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 Imrely lialf its length, the meatus
slightly projecting but not tubular.

Teeth. — The teeth are less robust but more j^rehensive than
those of Mnstela putorius, the former peculiarity best shown by
the upper cai'nassial, 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 M. imtur'ms,
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 carnas.sial and posterior surface
of pm?, instead of being widely removed from carnassial and
fitting distinctly inside of ptm^.

Measure)iientsi. — 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: —
Fkance : Vinsac, Gironde, 2 (Lataste) ; Soulac, Girondc, 1 (Lataste) ;
Laval, Mayenne, 2 (Lataste).

Roumania : Malcoci, Dobrudscha, 1 (Schliiter).

Sub-Genus PUTORIUS Cuvier.

1317. Putorius Cuvier, Eegne Animal, i, p. 117 (Type by tautonymy

Mnstela imtorius Linnaeus).
1810. Fcctorius Keyserling and Blasins, Wirbelthiere Europas, p. G6

{jmtorins).
1857. Fcctorins Blasius, Silugethiere Deutschlands, p. 219 (part).
1877. Cijnomiuiiax Coues, Pur-bearing Animals, p. 99 (Putorius nigriiKS

Audubon and Bachman).

Type sjicclcs. — Mustela putorius Linna-us.

Geographical distribution. — Mediterranean region and central
Europe from Great Britain eastward into China; central United
States ; exact limits of range not known.

Characters. — Form moderately slender ; tail bushy ; fur not
modified for aquatic life ; skull with noticeably projecting

MUSTELA 419

angular mastoid processes ; auditory Inillfe distinctly triangular in
outline ; inner lobe of upper carnassial functioning against
paraconid of lower carnassial ; interrelationships of teeth essen-
tially as in Mustela, but small premolars more robust and less
trenchant.

Bemarks. — The subgenus Putorius, 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 Mnnfela is now
divided. No intermediate forms are known connecting it with
either Lutreola or true Mustela, and if none are found 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 Linnaius.

(Synonymy under subspecies.)

Geo(p-aphical (listrihutlon. — 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 bullaj
slightly longer than broad, triangular in outline ; size nearly
equal to that of the martens (head and body of adult male about
400 mm.) ; foi'm slender, fur coarse, tail rather bushy, tapering :
colour bully grey or yellowish overlaid with black, the under-
parts darker than back.

External cliaracfers. — Form much as in the members of the
sub-genus Mustela, but appearing less attenuate on account of
the longer fur and somewhat bushy tail. Underfur rather dense
and woolly, about 25 mm. long in winter, l.o 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,
liare 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? : i 4 — 4 = 8.

Colour. — Feet, tail, chest and intercrural region blackish, rest
of body %vith long black hairs producing a clouded effect over

2 E 2

420 CARNIVORA

buiiy grey or yellowish under fur, the two elements of the
pelage distinct and not intimately Vjlending, the black more pre-
dominant in winter than in summer, the exact shade of underfur
ranging from a light buff}^ grey with pale ecru-drab bases to
the hairs to a clear buti'-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 bufty grey, this light area, some-
what dulled by admixtuie 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.

Skidl. — The skull is robust and strongly built, rather Hat,
with short rostrum, short, broad brain-case, and long, nearly
parallel-sided postorbital constriction. Dorsal profile nearly fiat
from slightly 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 visiVjle 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 longitutlinal ridge
and shallow lateral depressions ; a transverse depression in front
of lip of foramen magnum. Auditory bull* moderately inflated,
e-ipecially along inner edge, about three-quarters as wide as long ;
the general outline triangular with anterior border' shortest, the
meatus not tubular ; anterior extremity of bulla not in contact
with hamular, and separated from foramen ovale by an oblique
ridge which is marked off from glenoid process by a deep
conspicuous notch, the distance betsveen 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 Liitrii Intra anumg European Mii!<ti-U<Jse. Interorbital region
slightly broader than rostrum, moderately convex longitudinally,
a little more convex laterally ; postorbital processes .short but
distinct, directed slightly backward. Nostrum short and heavy,
the width across canines ecpial to <_ii.stance from gnathion to
anteorbital foramen. Orifice of anteorbital foramen moderately
large, over anterior 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 by a broad, shcallow concavity and a rather abrupt
posterior convexity. Palate moderately broad, its width between
molars equal to a little less than twice greatest diameter of

Fig. Si.
Muatela putorius. JJat. size.

iiKjlar ; incisive foramina small, l)roadly ovate, oblique, in front
of canines and very close behind alveoli of incisors, the minute
median foramen between oi- behind posterior border ; posterior

422 CAENIVORA

extension of palate longer than broad, emarginate postei'iorly ;
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 faintly convex from broad,
ill-detined angular 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 martes,
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 Marten and enamel of lower tooth usually
less rugose. Anterior premolar both above and below (pm'^ and
jpjH,,) small, single-rooted, the crown area about equal to that of
outer upper incisor, the ill-developed cusp at front of crown, its
height scarcely etjual to \vidth of tooth. Middle upper premolar
(j^fu-^) two-rooted, compressed, the
outline of crown flattened-elliptical
when viewed from below, triangular
/'^'~" "^-^ when viewed from the side, the

{^B (^ 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^D-i)
essentially like pitfi but larger and
relatively higher, its crown less com-
pressed posteriorly. Middle lower
j-iQ g,- premolar (pw'a) intermediate in size

Mu.idnvutoriH^. Teeth. X 1^ <iiic^ fo™i between pm.^ and prn^.
Upper carnassial essentially like that
of Martes fu'tna, 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 2h times as
long as wide, the main ti'enchant portion of the tooth formed by
the paraconid and jarotoconid, which are essentially similar to
those of Maries, 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 pandurate 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

MUSTELA 423

smaller, the outer border of crown slightly notched in region
between cusps.

Bemarks. — In certain conditions of pelage Mustela puforius
bears a superficial resemblance to Marten foina, though the two
animals may always be distinguished by the different form of the
ears. Two geograpliical races ai-e at present recognized, though
their status is far from well understood.

The ferret, Martcs furo Linnaeus,* though usually assumed to
be a domesticated variety of Mustela j^yutoriiis, 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 buUte, 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 putorlus, but which render them practically
identical with those of the Eastern form.

Mustela putorius putorius Linnaeus.

1758. [Mustela] putorius Linnaeus, Syst. Nat., i, 10th ed., p. 46.
1785. Mustela iltis Boddaert, Elenchus Animalium, p. 87 (Renaming of
2)utorius).

1797. ]\[[usteld] furo-putorius Link, Beytrjige zui- Naturgesch., i, p. 83

(Based on a melanistic putorius popularly regarded as a hybrid
between furo and putorius).

1798. Viverra foetens Thunberg, Beskrifning pS Svenske Djur, p. 15

(Renaming oi putorius).

1801. M\icstela~\ pliitoriusl albiis Bechsteiu, Gemeinn. Naturgesch. Deutsch-
lands, I, 2nd ed., p. 782. Not Mustela foina 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 S61ys-Longchamps, Etudes de
Micromamm., p. 145 (jiomen nudum).

1839. ? [M^istela putorijis] var. vison De S^lys-Longchamps, Etudes de
Micromamm., p. 145 [nonien nudum).

1843. Putorius fcetidus Gray, List Spec. Mamm. Brit. Mus., p. 64 (Re-
naming of piutorius).

1857. Fcetorius putorius Blasius, Saugethiere Deutschlands, p. 222.

1863. Putorizis infectus Og6rien, Hist. Nat. du Jura, iii, p. 59 (Substitute
ioi putorius).

1904. Putorius putorius Barrett-Hamilton, Ann. and Mag. Nat. Hist.,
7th ser., siii, p. 389, May, 1904.

1904. Putorius putorius manium Barrett-Hamilton, Ann. and Mag. Nat.
Hist., 7th ser., xiii, p. 890, May, 1904 (Teufen, Appenzell, Switzer-
land). Type iu British Museum.

1910. Putorius putorius and P. putorius manium Trouessart, Fauna
Mamm. d'Europe, pp. 76, 77.

Type locality. — Sweden.

Geo(jrapMcal 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.

Measuronents. — 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,
Rheinhessen, C4ermany : head and body, 400 and 440; tail, 160
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, Ajjpenzell, Switzerland (type of mmiium 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, Sutherlandshire, 1 ; no exact locality, S.

England: No exact locality, 2; Oundle, Northamptonshire, 1; BuUnose
Coppice, Northamptonshire, 1 ; Wisbech, Cambridgeshire, 1 ; Swaffam Fen,
Cambridgeshire, 1 ; Cardiganshire, G ; Aberystwyth, Cardiganshire, 6.

Sweden : No exact locality, 1 (skull ; U.S.N.IM.) ; Slsane, 1 (Stockholm).

France : Scientrier, Haute-Savoie, '2 (IMottaz).

Ctermany: Ingelheim, Rheinhessen, 6; Ummerstadt, Thiiringen, 9;
ilagdeburg, Saxony, 2 ; Marxheim, Bavaria, 1 ; Niesky, Silesia, 1.

Switzerland : Teufen, Appenzell, 1 (type of maniuni Barrett-Hamilton) ;
St. Gallen, 2; Urmisch, 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.

Roiiarl-s. — The Swiss form, to which the name mnninin has
been given, appears to be identical with true Miistcla iiutorhm.*

6. CHencassley, Suther- Mrs. Flower (p). 92. 10. 21. 2.
landshire, Scotland.

i, ?, 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.

ganshire, Wales. child (p).

^, 2 9 St. .\bervstwvth. Hon. N. C. Roths- 1.5.22.5 7.

child (p).

* The names Piitorius ti/pus P. Cuvier and P. communis G. Cuvjer
mentioned by Trouessart as synonyms of M. pyforiux I have been unable
to verify.

MUSTELA 4-2")

(?, 9. Aberystwyth. {Riiskin- G. Barrett-Hamilton 11. 1. 2.105-lOG.

Buttcrjiehl.) (v).

S skeleton. Boneath, Cardigan- W. E. de Vinton 0. 10. 9. 1.

shire. (c & p).

2. England. Purchased. 39. 7. 15. 2-.S.

3 S, 9. Ingelheim, Rhein- C. Hilgert (c). S. 11. 12. 18-21.
hessen, Germany.

6,9. Ingelheim, Ehein- G. Barrett-Hamilton 11. 1.2. 107-108.

hessen. (Erlangcr.) (p).

2 6, juv. Ummerstadt, Thiirin- Lord Lilford (p). 11. 1. 1. 94-95.
gen. {Schuchardt.)

9. Magdeburg, Saxony. Lord Lilford (p). 11. 1. 1. 93.

{Woltcrstorff.) 0. 2. 8. 5.

V. Niesky, Silesia, Ger- Lord Lilford (p). 97. 12. 4. 19.
many. (Baer.)

5. Teufen, Appenzoll, O. Thomas (p). 2. 8. 4. 24.

Switzerland. {Zolli- {Type of Putorius p. inankim

kofer.) Barrett-Hamilton.)

6. St. Gallen. (Zolli- 0. Thomas (p). 4. 4. 5. 32.

kofcr.)

9. Florence, Italy. E. Cavendish Taylor 5. 5. G. 9.

(c & p).

3(?,2c5juv. Viterho, 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 Mag. Nat.
Hist., 7th ser., xiii, p. 389, May, 1904. Type in British Museum.

Type hicnlity. — Ferrol, Province of Coriina, Spain.

(!eo(irapliical 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.

Measnrevients. — Type: hind foot, 6 1 • 5 ; ear, 19. For cranial
measurements see Table, p. 426.

Specimens examined. — Four, from the following localities in Spain :
Ferrol, Corufia, 1 (type) ; Seville, 1 ; no exact locality (probably vicinity of
Seville), 2.

Remarks. — 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
Mnstela pidorins 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, Coruna, Spain. Dr. V. L. Seoane (r). 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. G. 11. 2 3.

426

CARNIVORA

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Genus VORMELA W. Blasius.

1857. Fcetorius Blasius, Siiugethiere Deutschlands, x">. 219 (part).
1884. Vonnela, W. Blasius, Bericht der Naturforscb. Gesellsch. in Bemberg,
XIII, x^. 9.

Type species. — Mnstela snntudira Pallas = M. peregnsna
(itieldenstaedt.

Geoi/raphiral distrihiition. — From soutli-eastern Europe to
central China.

Chdracters. — Skull narrow, somewhat flattened (depth of
hrain-ca.se a little more than half mastoid breadth), the dorsal
pr(jfile slightly arched posteriorly, rather strongly bent down
from orbital region forward, the zygomatic arches not specially
widened, and postorbital region not unusually narrowed (distance
fi'om jioint of greatest narrowing to zygoma less than breadth of
])Ostorbital constriction) ; rostrum rather short aiid 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 reaching level of foramen ovale ; dental formula :
i ^± c ti ynn ^~, m '-^ = 34 ; teeth relatively somewhat larger
than in Mustela, the canines, especially those of upper jaw,
relatively higher than in any of the other European MnsteJid-ce ;
upper carnassial I'obust, the width of its cutting portion about
half length, the anterior cusp so developed that its greatest
height is nearly 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 })o.sterior border extending about
to middle of crown ; upper molar elongate-pyriform, tlie outer
portion only a little narrower than inner, the median constriction
slight, the axis of crown strongly obli([ue 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 p)olecat-like ; fur rather soft and den.se, varied with black,
brown, and whitish or yellowish ; tail about half as long as
body, bushy.

Remarks. — This strikingly characterized genus is readily
distinguished from Mustela, apart from its peculiar external
appeai-ance, by the structure of the lower carnassial and by the
relationships of the hamular, auditory bulla and foramen ovale.
Two species ai'e known, imv of which ranges as fai- west as
eastern Hxnigary.

429

VORMELA PEREGUSNA Gueldenstaedt.

1770. rcrcgusna, nova Mustchv species, Gueldenstaedt, Nov. Comm. Acad.

iSci. Ivyp. Petrop., xiv, p. 441.

1771. Mustela sarmalica Pallas, Raise durch verschiedene Provinzen des

Russischen Reichs, i, p. 453 (along the Volga in southern Russia).
1857. Fcetorius sannaticus Blasius, Siiugethiere Deutschlands, p. 226.
1910. Putorius sannaticus Trouessart, Faune Mamm. d'Europe, p. 77.
1910. Vormcla peregusna Miller, Proc. U.S. Nat. Mus., xxxviii, p. 385,

August 19, 1910.

Tjpc JocnJltj/. — Banks of the River Dun, southern Russia.

Geographical distribution. — From central Asia west through
Asia Minor and southern Ru.ssia to Roumania, Bulgaria and
eastern Austria-Hungary (Bukowina).

Diagnosis. — Size about as in Mustela piitorlitx (head and body
in an adult male, 340 mm., condylobasal length of skull about
50 mm.) ; aviditory bulla sub-triangular in outline.

External form. — In general external characters (dry .skins
only examined) the animal appears to agree closely with Mustela
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 jjalms thickly furred except on the pads,
all of which are naked ; palm with two posterior tubercles, the
outer somewhat smaller than in M. pmtorius, the inner about
half as large as outer. INIammaj : a 4-4, i 2-2 =12.

Colour. — Ground colour of upper parts raw-umber, most of
the hairs with dai-ker tip and lighter subterminal band, a few
blackish throughout, the combination producing a slight effect 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 partly 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 transverse bands, their
total combined area slightly less than that of brown background ;
a stripe extending from Ijack 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-draV), in light areas similar to long hairs but paler, so that
the colour pattern becomes more sharply defined on areas whei'e
the fur has been nearly worn off. Head, feet, legs and under-
parts varjnng from dark seal-brown to lilack. Lips and inter-
ramia whitish or buffy. A broad 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

OARNIVOKA

suffusion of whitish behind it. Ear white or bufl'y except at
extreme base, which is blackish. Tail black at tip, elsewhere
white set oiF by underlying black ; the individual hairs cream-
colour through basal third, then darkening rapidly through raw-
umber to Ijlack, the terminal third white.

SJcuJI. — The skull is smaller than that of MiisteJa imtorius,
though resembling it in its general strength and raassiveness.

Via. 86.
Vormela pere(jiit<na. Xat. size.

particularly in lateral view. Doi'sal profile almost exactly as in
Mustela putorius, flat from level of anterior portion of orbit to
occipital region, the rostrum bent downward at an evident angle.
Depth of rostrum relativel}^ great, as in M. putorius, and notice-
aljly exceeding that of Mustela erminen. Postorbital constriction
abi'upt, as in Mustela eversmanni, giving the skull when viewed
from above a very peculiar aspect as compared with that of

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CARNIVORA

3T. j)ufortiifi. In fully adult individuals the pustorbital 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 ])utorius and
Mnsfela ermhiea the teeth are through-
out I'elatively heavier and the cusps
more elongated. Height of upper
canine slightly more than depch of
rostrum, instead of decidedly less than
rostral depth as in Mustela putorius.
Upjaer carnassial with brotxder antero-
internal projection than in 31. pntorhi!<,
and upper molar set more obliquely to
main axis of skull. Lower carnassial
strikingly different from that of all
species of Mustela in the presence of
a well developed third cusp on anterior

I oniiela pereijmiia. . , , /. ^.,1,11.

Teeth, x u. elevated portion or tooth, the torm ot

which is thus essentially as in the
martens. Interrelationships of small premolars much as in
3Iustela putorius.

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 Malcoci,
Dobrudscha, Eoumania (B.M. and U.S.N.M.) ; also others from various
localities in Asia.

BemarJiS. — Vormela 'peregusna is so well characterized both
cranially, dentally and externally as to require no detailed com-
parison with other European carnivores.

i. IMalcoci, Dobrudscha, Rou- Hon. N. C. Rothschild 10. 6. 4. 1.
mania. (Schliltcr.) {v).

Sub-Pamiia- GULUNINzE.

GeDijraplik'dl disty'ihutinn. — Xorthern forested portions of the
ISTorthern Hemisphere.

Characters. — Dentition essentially as in the MusteVnise but
much le.ss trenchant, the small premolars opposite and not capable
of shearing action, the points of all but pm-^ and pm^ widely
separated when jaws are closed ; upper carnassial very robust,
its posterior cusp broad, almost flat-topped, its height much more

GULo 433

than lialf that of main cusp ; skull robust and heavy ; external
form heavy, not in the least cat-like ; feet sub-plantigrade.

Remarks. — The sub-family Gidanhise, consisting of the genus
Gido alone, is well characterized by its peculiarities of skull,
teeth and external form. Though usually regarded as a near
relative of the MtiKiclln.',r, the genus more probably finds its true
affinities in the African McUirova.

Genus GULO Storr.

1780. Gulo Storr, Prodr. Jleth. Mamm., p. 34.

1857. Gulo Blasius, Saugcthiere Deutschlands, p. 208.

Type species. — Mustela gulo Linnteus.

GeorirapMcal distribution. — Northern forests of the Northern
Hemisphere ; in Europe confined to Scandinavia and noi-thern
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 downward
anteriorly, the zygomatic arches not specially M'idened, and
postorbital region not unusually narrowed (distance from point
of greatest narrowing to zygoma less than breadth of postorbital
constriction) ; rostrum short and very robust, its width about
equal to that of interorbital region, the distance from lim 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 narrowing tube, the transverse
diameter of bulla greatest ; paroccipital process I'obust, long,
standing out conspicuously behind bulla ; dental formula :

*' :i ii) '■ 1-1 > P'"- 4^1 "* a^'j = 38 ; teeth relatively lower, wider and
more i-obust than in Mastela (their actual size conspicuously
greater) ; small premolar's almost directly opposed, not fitting
between each other when jaws are sliut, 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-fifth 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.

ItemarJcs. — The genus Gulo is represented bj^ two closely
related species, G. hiscus of the northern portions of America,
and G. (julo of the Old World.

434 CAENIVOKA

GULO GULO Linnajiis

1758. [Mnstela] giilo Liunseus, Syst. Nat., i, 10th ed., ■p. 45.

181G. Gnlo vulgaris Oken, Lehrbuch d. Naturgesch., iii, pt. 2, p. 1001 (Re-
naming of (julo).

1820. Galo borealis Nilsson, Skaud. Faun., i, p. 95 (Renaming of gHlo).\

1820. Gulo ardicns Desmarest, Mammalogie, p. 174 (Renaming of gulo).

1829. Gulo ardos Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierw., i,
p. 68, described on p. G9 (Renaming of gulu).

1857. Gulo borealis Blasius, Siiiigethiere Deutschlands, p. 209.

1910. Gulo luscus Trouessart, Faime Mamm. d'Enrope, p. 71. Not Ursvs

luscus Linnteus, 1758.

1911. Gulo gulo Collett, Norges Pattedyr, p. 38G.

Type locality. — Lapland.

Geograp]i!cal distribution. — Northern forests of the Old
World ; in Europe, confined to Scandinavia and northern Russia.

Diagnosis. — Largest of the European MustcJ idne (head and
Ijody, 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.

E.xternal 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 gi'owth 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 verteljras 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 vertebrte. 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 increa.sing
abruptly to twice as long on flanks and rump, thus adding greatly
to the heavy appearance of the animal ; long hairs on wrist and
sides of fore foot tending to become almost bristle-like in texture.

Colour. — Upper parts and belly a rich dark brown becoming
Ijlackish 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, ami face to

(iL!LO 435

between eyes, suffused with ereaiii-buir, 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.

SJluH. — The skull is larger and more massive than in any of
the other European MitstcJldse, its general form somewhat inter-
mediate ))etween that of Melea and Mdvfi's. Dorsal profile with
slight anterior concavity over middle of short nasals, then ratliei-

Fig. 8S.
Gulo gulo. Reduced.

strongly convex to bregmal 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 moderatel)^ developed
in old individuals and joining rather low larabdoid 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 ntartcK but relatively wider.

2 F 2

436 CARNIVOEA

Auditory bullae inflated along inner border, elsewhere rather flat,
their general form elongate flask-shaped (transverse diameter
greatest), much like that in ilfe/cs melen, the meatal tube rather
long but not sharply difTerentiated from main portion of bulla.
Postorbital region less abruptly contracted than in Melcs meles
and Martes martcs, its width slightly more than half that of
brain-case. Interorbital region about as wide as rostrum,
smoothly convex both longitudinally and laterally. Postorbital
proce-sses short but relatively better developed than in Maries.
Rostrum short and heavy, the width across canines equal to
distance from gnathion to infraorbital foramen. Oritice of

I'lo. so.

Gulo (juln. llcdiiced.

anteorbital foramen relatively small, its actual size scarcely
greater than that in 3Iartcs martcs, its situation over posterior
root of iinfi 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 compressed
and deepened, especially over glenoid region. Palate rather
broad, its width lietween molars equal to nearly 2^ 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 ill 3Tartes. Mandible very robust and massive, but with Jio
special peculiarities of form. It is distinguishable from that of
Meles viclcH, 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 martes. Small premolars both
above and below with crowns relatively lower and broader than
in Martes, and less distinctly triangular when viewed from the

Fig. 00.
Gido (julo. Eecluced.

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 p/u^ and jmi^^ l)arely coming in contact, those of the
other teeth separated by a wide space ; jjjh^ with slightly raised
cingidum at posterior border, but without trace of the secondary
cusp present in 3Iartes. Upper carnassial like that of Maries in
genei'al outline, but posterior cusp relatively much larger, its
height distinctly more than half that of main cusp, its outer

438

CARNIVORA

surface at first nearly flat, then sloping almost perpendicularly
to very rudimentary cingulum ; commissure relatively shorter than
in Maries. Lower carnassial slightly more 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 cingulum ; a similar ridge at outer
side of protoconid and across posterior heel slighth' external to

Fin, !il.
(iiiloijuln. Tffth. ^'at. 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
tliat of j)JH2) ''^^^ ^ little larger than that of posterior heel of m.,.
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 e.ssentially as in Maries
martes, but paracone and metacone tending to become confluent.

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440 CARNIVOEA

3Ieasurements. — Adult (summer pelage) from Sweden (from
skin): head and body, 825; tail, I'io (pencil, 75); hind foot,
140±. For cranial measui-ements see Table, p. 439.

Specimens examined. — Sis, from the following localities : —

Norway : Vaage, G^udbrandsdal, 1 (Genoa) ; Egersund, Stavanger,
1 skeleton and 1 skull.

Sweden: Frastvik, 1; no exact locality, 1 skull (U.S.N.INI.).

Lapland: No exact locality, 1 skull (U.S.N.M.).
6 skeleton, Egersund, Norway. K. H. Schaanning (c). (10. 11. 26. 1.
? skull. 111. 10. 2:j. 1.

Family VIVEREID^.

1821. Vivcrrich: Gray, London Med. Repos., xv, p. 301, April 1, 1821.

GefKjrapliical distribution. — Warmer portions of the Old World
(e.xclusive of Australia) ; north to the Iberian Peninsula and
southern France.

Characters. — Dentition essentially as in the Mu»tdiflR', but
upper 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.

BemarliS. — The family Vicerridse is a tropical Old World
group, containing about 25 genei'a, the ranges of two of wliich
extend into the Mediterranean region of Europe.

KEY TO THE EUROPEAN GENERA OF VIVEnRIDJE.

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 (^Mou-

goose) Mangos, p. 140.

General form cat-like ; ear higher than wide ; body spotted ;

tail ringed ; bony palate extending behind molars in

median line mucli less than half distance to hamulars ;

orbit widely opening into temporal fossa (Genet) Gcnctta, p. 116.

Genus MUNGOS Geoffroy and Cuvier.

1795. Mnngos Geoffroy and Cuvier, ilagasin Encyclopedique, ii, p. 187

(tnungo, p. 184).
1811. Herpcstrs Illigor, Prodr. Syst. ilamm. ct Avium, p. 135 (misspelt

Hfipcrtes in text, corrected in list of errata, ]). 302) {ichneumon).
1824. Maiiijiista Horsfield, Zool. Researches in Java (Javanica).
1830. Maries Waglcr, Nat. Syst. dor Amphibien, p. 29 (Substitute for

lIer2Kstcs Illigor, preoccupied in botany). Not of Pinel, 1792.
1842. Mango Lesson, Nouv. Tabl. Regne Anim., Mamm., p. 63 {ichneumon).
1907. Mangos Thomas, Ann. and Mag. Nat. Hist., 7th ser., xix, p. 119,

January, 1907.

Tjipc sjiecie.'y. — Yin rra miuu/o Gnielin. "

Geographical iJi.s/ribuiioH. — Africa, southern Asia, and larger

MUNGOS 441

Malayan Islands ; in the INIediterranean 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
' 33' 1-1' ■' 4-4' 22 '

trenchant.

BemarJcs. — The widely dispersed genus Mungos, which is one
of the most characteristic features of the carnivorous 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.

18-12. HcrjKstes loiddnngtoiiii Gray, Ann. and Mag. Nat. Hist., 1st ser., ix,

p. 50, March, 1842. Type in British Museum.
1909. Herpestes ichneumon var. ferruginca Seabra, Bull. Soc. Portugaise

Sci. Nat., II, p. 286, May, 1909 (Altemejo, Portugal).
1909. Herpestes ichneumon var. dorsalis Seabra, Bull. Soc. Portugaise Sci.

Nat., II, p. 286, May, 1909 (Eibatcjo, Portugal).

1909. Herpestes ichneumon var. grisea Seabra, Bull. Soc. Portugaise Sci.

Nat., II, p. 286, May, 1909 (Ribatejo, Portugal).

1910. Herpestes ichneumon widdringtoni Trouessart, Faune Mamm.

d'Europe, p. 89.

Tijpe locality. — Sierra Morena, Spain.

GeograpMcal distribution. — Iberian Peninsula, principally in
the south ; exact limits of range not known.

Diagnosis. — Similar to the African Mungos icltnemnon, 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 buUai (lateral diameter of
swollen portion about 12 mm. instead of 10 mm.).

External characters. — Genei'al 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 rounded. 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 lai'ge

442

CAENIVOEA

and elongate, with long, non-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 naked, 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, thus
giving the tail its characteristic tapering form ; pencil long and
full. Mammte : a 2-2 = 4.

Colour. — The colour does not diflfer appreciably from that of
Moroccan and Egyptian specimens of Mnntjos ichtummon. Head,
Irnck, sides and tail (except the clear black tip) a coarse grizzle
of blackish and pale bufi", each hair on body and tail buff at base
and with three buff annulations 3 to 5 mm. in length interposed
between the Ijlackish 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 finer 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 entir-e fore leg except the uppermost
outer portion. On foi-efeet and outer portion of leg below elbow
it becomes blackish.

SJcuIl. — The skull is narrow and ratlier high, mastoid breadth
less than occipital depth including bullse, 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 zj^goma rising to meet it. Anteorbital foi'aman small, higher
than wide, over space between pm- and jjtn'-^. Median back-
ward prolongation of palate conspicuous, extending rather more
than half way to hamulars, its posterior border squarely truncate

MUNGOS 443

or irregularly angular-emarginate, never with median projection ;
niesopter^'goid space much less than twice as long as broad,
parallel-sided ; hamulars robust, the thickened extremities bent

I

Fig. 92.
3Iu)igos u'lddriivjlonii. Ruclucecl.

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 robust, 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 l)i-
lobed, the outer lobe largest, Ante-
rior premolar both above and below
simple, smaller than corresponding
outer incisor. iSecond 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 l)asal tubercle ; inner me-
dian projection well developed and
bearing a small cusp in ^jm-', slightly
indicated inpm'^ ; the two lower teeth
with longer base and less robust
main cusp, jjfflg 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 cusji
on posterior cutting edge large and well developed. Upper
carnassial with large antero-internal lobe somewhat exceeding
posterior heel in area, its cusp roljust ; antero-external corner
with large, occasionally almost cu.sp-like cingulum ; anterior
liorder of tooth slightly concave. Lower carnassial with the
three anterior cusps large, the outer exceeding the two inner,
which are of appmxiniately 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 Ijorder elevated
and trenchant. First upper molar with crown nearly twice as
wide as long, its anterior border convex, its outer and posterior
Ijorders concave ; protocone large, occupying entire inner Ijorder
of crown ; paraconc and metacone small, sub-equal, the latter

MiiiKjos widdritirjt.ou i,
Teeth. Nat. size.

posterior cutting edi

445

Observations.

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

„ slightly worn.

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362 B.H.

363 „

1

M. widdringtonii.

Spain : Ferrol, Coruiia
Sierra Morena
Seville .

Goto Donana,!
Huelva . /

M. ichneumon.

Morocco : near Tangier

446

(JARNIVOEA

occupying posteroexternal corner of crown, the former exceeded
by the hirge 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 Urst, its antero-external extremity
fitting into convexity behind metaconeof m^ ; in general structure
it is like iii^ 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 ;//j in height, its borders sometimes with four distinct
cusps, the two on inner side largest, but more often with postero-
external and postero-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 ; tad, 450; hind foot, 95 •5. Immature male
from Ferrol, Coruna ; hind foot, 86.

Specimens examined. — Eight, from the following localities in Spain :
Ferrol, Coruna, 1 ; Sierra Morena, 1 (type) ; Seville, 3 ; Goto Dofiana,
Huelva, 1 ; Audalucia, no exact locality, 2.

Remarks. — Among the smaller carnivores of Europe Muixjos
loiddringtonii is recognizable by its conspicuously grizzled coloura-
tion and long, coaivsely-haired, tapering tail. It is nearly related
to the African M. ichieumon, though sufficiently distinguished by
its larger teeth.

6. Ferrol, Coruna, Spain. Dr. V. L. Seoaue 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-G.

skull. Seville. Col. Irby (c & p). 85. 9. 1. 1.

S. Coto Dofiana, Huelva. A. Chapman (c &. p). 10. 7. 14. 1.

ad & juv. Andalucia. Lord Lilford (p). 7S. 7. 3. 2-3.

Genus GENETTA Oken.

1810. Genetta Oken, Lohrb. der Naturgesch., in, pt. 2, p. 1010 (genctta).
1841. Odmu'lnrus Ciloger, Genieinn. Hand- u Hilfsbucb der Naturgesch., i,
p. 72 (genetta).

Type species. — Vtverra genetta Linnaeus.

Geograj/Mcal 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

GKNKTTA 447

of palate short ; auditory bulla complete, its two divisions not
strongly contrasted in size and form ; dental formula : / -' ■', r ' ',
pm '—, m fl = 40 ; cheek-teeth trenchant.

Memarhs. — The genus Geneita, though practically confined to
Africa, extends its range into the Mediterranean region of Europe,
chiefly in Spain and the western portion of Frsmce. 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 OP GEN ETTA 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) G. (/. i-hndamca, 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 buffygrey; largest spots

about 20 mm. in diameter (Central and southern

Spain) G. g. genetta, p. 451.

Ground-colour of back a smoky grey ; largest spots

about 30 mm. in diameter (Balearic Islands)... G. g. balearica, p. 452.

GENETTA GENETTA Linnpeus.
(Sj-nonymy 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 taj^ering, the muzzle somewhat
pointed ; ear rising consi^icuously 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
bai-e area extending downward as a narrow line dividing middle
of upper lip ; whiskers well developed, reaching beyond base of
car when laid back. Feet cat-like, the digits short, armed with

448

CAENIVOKA

retractile clav/s, 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 ; ])alBi with large, cordate, obscurely
3-lobed tubercular mass at base of longer digits, a small roundish
pad at base of thumb, and two semi-contluent, 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 V)ack, 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 tijj. Mammas : a 2—2 = 4.

Colour. — Ground colour throughout a light grey, with or
without an evident bufty or brownish tinge, the slaty bases of
the hairs sometimes appearing at surface, particularly on under-
parts. Back and sides thickly .spotted with black, the spots
always confluent along median line, where they form a narrow
strijje 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-detined median dark streak and faint supraorbital shade ;
muzzle with conspicuous l)lackish area at base of whiskers and
extending back to eye, the extreme anterior region at side of
pad and beneath it shar])ly 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 10 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 Herpestes widdmigtonii
in general form, though much less robust, not unusually
deepened, the orbit large, widely open posteriorly, its vei'tical
diameter greater than depth of rosti'um at canine, the auditory
bulLe not much inflated ; mastoid breadth about equal to
occipital depth including bulhe ; 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. Reduced.

short but well developed median projection ; mesopterygoid
space a little more than twice as long as wide, parallel-sided ;
hamulars slender, with tapering, nearly horizontal extremities ;
auditory bulla with anterior annular portion rather large, slightly

2 G

450

CARNIVORA

but evidently inflated, the lari,'e meatus without trace of tube ;
posterior inflated ^jart 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 Miingos loiddrinf/tonii ; 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 more evident posteriorly than anteriorly ;
coronoid process with height above condyle about equal to width
at same level.

Teeth. — The teeth do not difler markedly from those of
Mungos wicldringtonii 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 wid-
dringtonii, its size greater than that of
corresponding outer incisor, jjni^ with
minute postei'0-ba.sal 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 pm^ often with a minute
lobe near base of posterior cutting edge ;
inner median projection well developed
and bearing a small cusp in jini^, slightly
indicated in pnr ; the two lower teeth with base scarcely longer,
both with small but evident anterior and posterior cingulum
cusp, and pjjjg with well develojjed median cusp on posterior
cutting edge ; fourth lower premolar like second and third
but larger, the crown less narrow behind, and the cusp on
posterior cutting edge better developed. Upper carnassial with
well developed, cusi>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 ; 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 jiortion 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

Geiietta genettn.
Teetli. 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 rather 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.

Beinarks. — 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 LinnseUS.

1758. IViverra] genetta Linnaeus, Syst. Nat., i, lOth ed., p. 45 (Spain).
1816. Viverra Genetta Jiispanica 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 France 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] communis 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?[ genetta Trouessart, Catal. Mamm. viv. fors., p. 325.
1905. ? Genetta peninsula; Cabrera, Bol. Real Soc. Espaii. Hist. Nat

p. 266, May, 1905 (El Pardo, near Madrid, Spain).
1910. Genetta genetta tnelas, G. genetta peninsulpe and G. afra Trou^sart,
Paune Mamm. d'Europe, pp. 88, 89.

Type locality. — Spain.*

Geographical 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 lai-gest about 20 mm. in diameter ;
ground colour of back a distinctly bufly grey.

Measure ruents. — 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.).

Remarks. — On the basis of the few specimen.s 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 d, 2 ?. 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 baleahica Thomas.

1902. Genetta genetta balearica Thomas, Ann. and Mag. Nat. Hist., 7th ser.,

X, p. 162, August, 1902. Type in British IMuseum.
1910. Genetta genetta balearica Trouessart, Faune Mamm. d'Europe, p. 88.

Tjijie hn-aJitij. — 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 underfur more whitish than in the continental Spanish
animal ; spots rather larger (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 ?. Majorca, Balearic Islands. 0. Thomas (p). 1. 6. 1. 1-4.

{Dr. Riutort.) (1. 6. 1. 3. Tijpe 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. ^

Geographical distribution. — South-western France from the
valleys of the Pthone and Loire to the Bay of Biscay ; perhaps
westward through the Asturias. I

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GENETTA 455

Diagnosis. — Ground colour as iii Genetta genetta geneita, 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 cvamined. — Four, from the following localities in France :
Sainte-Ci'cile, Vaucluse, 1 ; near Nimes, Gard, 1 (Mottaz) ; near Biarritz,
Basses- Pyrenees, 2.

Bemarhs. — An immature individual from Ferrol, Coruna,
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-Pyr6n6es. Zoological Society 8. 1. 26. 1-2.

1. Ferrol, Corufia, Spain. Dr. V. L. Seoane 94. 11. 3. 2.

(c. & p.)

Family FELID^.

1821. Felidx Gray, London Med. Repos., xv, p. -302, April 1, 1821.

Geographical 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 ci'owns compressed and high, without the crushing surfaces,
the last upper premolar and tirst 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 tiansverse 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.

Remarks. — The family Felidse is one of the most widely
distributed and sharply diflferentiated 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 commou 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

or not tufted (Cats) Felis, p. 450.

Small anterior upper premolar absent, teeth 28 ; ear con-
spicuously tufted (Lynxes) Lynx, p. 470.

456 CARNIVORA

Genus FELIS Linmeus.

1758. Felis Linnfeus, Syst. Nat., i, 10th ed., p. 41 (Type by tautonymy

Fells catus Linnseus).
1855. Catus Fitzinger, Wiss.-pop. Naturgesch. der Saugith., i, p. 2G5

(catus by tautonymy).

1857. Felis Blasius, Saugethiere Deutschlands, p. 159.

1858. Catolijnx Severtzow, Rev. et Mag. de Zool., 2nd ser., x, p. 385 {catus).

Tijpe species. — Felis catus Linnfeu.s.

Geograplncal distribution. — Tropical and northern temperate
portions of the Old World ; exact limits of range not known.

Characters. — Dental formula : i •^', c J}J, pm ||, m Jfl = 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.

Bemarks. — 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. silvcstris, p. 457.

Teeth enlarged (jmi^ and pm^ together 19 to 20 mm. ;
three lower cheek-teeth together 22-4 to 23-6

mm.; ?«, 9-2 to 10 mm.). Southern Spain F. s. tartessia, p. 465.

Teeth normal {pm-' and p7n* together 16-G to 19 mm. ;

three lower cheek-teeth together 18 • 8 to 22 mm. ;

?», 8-0 to 8-8 mm.).

General colour light, approaching the smoke-grey

of Ridgway ; stripes on sides and legs obsolete

(Central Europe) F. .s. ><ih-estri^, p. 462.

General colour dark, approaching the broccoli-
brown of Ridgway ; stripes on sides and legs
well defined (Scotland) F. s. grampia, p. 464.

457

FELIS SILVESTRIS Scliveber.

(Synonymy under subspecies.)

Geofjraphical 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
cafus 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 strijjes 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 jaarts 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 apjjearing heavier on account of the longer
fur, a peculiarity e.specially 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 incon.spicuous, 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. — Underfur 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

CARNIYORA

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
himl legs cream-bufF, becoming whitish or white on chin,
throat, foi'e 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 browji. Tail black at tip (about
70 mm.) and with two to four narrow (15 to 30 mm.) black or
brownish rings, 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. jSTape with four narrow longitudinal black stripes,
these extending to head where they tend to become confused
arid coniluent, and disajjpearing 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 continue.s 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 \isible and when fur
is smoothed, their general transverse and parallel arrangement.*
In continuation of this system of transvei'se 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
rostrum when skull is viewed from in front). Dorsal profile of
skull convex throughout, sometimes a little Battened in inter-
orbital region and along nasals, the latter occasionally terminating
in a slight anterior concavity ; nasals sloping very abruptly
forward and downward, the palattil dejjfh 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 Felis catus as
described and figured by Pocock, Proc. Zool. Soc, London, 1907, pp. 143-168,
pis. VIII and ix.

PELis 459

extremity of nasals the convexity ot" 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 oldei- specimens by the projecting lambdoid border, the
greatest width above zygomata slightly greater than depth at

Fig. 96.
Feb's ^ili'estris. x

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
bulhu large, highly inflated, longer than broad, formed mostly by

460

CARNIVORA

posterior chamber, the surface smooth, the meatus large, not
tubular ; width of basioocipital at middle less than that of bulla
in same region ; mastoid and 2:)aroccipital 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 very large, at iirst 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 off 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 (enci'oached
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
anterior 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 Europeaii
Carnivores of other families, the dentition appears rather weak,
owing to the complete suppiession of all crushing surfaces and
the consequent narrowness of the cheek-teeth. Incisors small
and weak, the crown area of P, the largest tooth, about equal to
that of /I and i'^ together and S(^arcely 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 i^ 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 Felia nlvestru. 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 al:)out 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.

Rcmarhs. — Externally Fells nilcestris 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 CAENIVORA

longer fur, this giving the animal the appearance of more robust
legs and less tapering tail. The skull and teeth differ only in
the greater average size of fully adult individuals, a character

FlO. 9S.
Felis siloestn'x (upper flgiiie) and F. cahis (lower figure). Nat. size.

most readily appreciable in the teeth. In twenty European
specimens of Felis sih'estris and a like number of F. catm the
extremes for certain dental measurements are as follows : —
Combined length of upper carnassial silredr!s. catus.

andpw^ lG-6to20-0 .. 15-0 to 17-8

Combined length of lower cheek-teeth 18-8 ,, 23-6 .. 18-0 ,, 20-4
Lowermolar 7-8 ,, 100 .. 6-6,, 8-6

Fklis silvestris silvestris Schreber.

1758. {Felis] catm Linnaeus, 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 catm Blasius, Saugethiere Deutschlands, p. 162 (Not of

Linnaeus).
1896. Catus ferox Martorelli, Atti Soc. Ital. Soi. Nat., Milano, xxxv, p. 253,

January, 1896 (Accidental renaming of Catm ferus Brehm, Illustr.

Thierleben, i, p. 275, 1863).
1907. Felis sylvestris Pocock, Proc. Zool. Soc, London, p. 150, June 12,

1907.
1910. Felis silcestris Trouessart, Faune Mamm. d'Europe, p. 98.

Tjjpe locality. — Germany.

Geogra^phical distribution,— Centva] 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 i-ange not known.

Characters. — Teeth not unusually large (combined length of
upper carnassial and pm^ 16 '6 to 19 mm.; three lower cheek-
teeth together, 18 "S to 22 ram. ; lower molar, 8"0 to 8 -8 mm.) ;
general colour light, the predominant hue approaching the smoke-
grey of Ridgway ; 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.).

Pbance : Manonville, Meurthe-et-Moselle, 1 ; Salavon, Haute-lMarne, 1 ;
Caterille, Haute-Garonue, 1.

Germany : North Germany (no exact locality), 1 ; Ingelheim, Rhein-
hessen, 1 ; south Germany (no exact locality), 3 (skulls).

AuSTRiA-HuNGAEY : Baranza, Hungary, 1 (skull).

Bulgaria : Varna, 1 (Andersen, skull).

Greece: Near Athens, 1.

Italy : Near Rome, 2.

Hematics. — 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).
i. Manonville, Meurthe-ct- Dr. E. Hamilton 95. 11. 9. 1.

Moselle, France. (Lomoiit.) (p).
6. Salavon, Haute - Marne. Dr. E. Hamilton 95. 11. 9. 2.

(Lomont.) (p).

6. Caterille, Haute-Garonne. O. Thomas (p). 8. 7. 15. 1.

(.4. Robert.)
6. N. Germany. Lord Lilford (p). 95. 5. 1. 1.

3 skulls. S. Germany. Dr. A.Giinther (c). 59. 9. 6. 55-57.

1. Ingelheim, Rheinhessen, G. Barrett-Hamil- 11. 1. 2. 104.

Germany. (Erlanger.) ton (p).

6. Baranza, Hungary. Dr. E. Hamilton 2. 6. 3. 1.

(p).
1. Athens, Greece. 0. W. L. Merlin 47. 7. 22. 2.

464 CARNIVORA

Felis siLVESTRis GRA5IPIA Miller.

1907. Felis 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.

Type locality. — Invermori.ston district, Inverness, Scotland.

Geographical distributioH. — 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. — TJnderfur of back and sides a light ochraceous-bufF,
the basal half of the hairs mouse-grey. Light annuiations of
longer hairs very nearly the cream-ljuff 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 l)lackish. 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.

Shull 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, Ross, 2 ; Aberdeenshire, 1 ;
Invermoriston district, Inverness, 5 ; Beauly, Inverness, 1 ; Fort William,
Inverness, 2 ; Kuoydart, 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.

Remarhs. — The British wild cat diff"ers from true Felis
silvestris in darker general colour and more pronounced 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.

skull. Glencassley, Sutherland- E. R. Alston (p). 79. 9. 25. 81.

shire, Scotland.
i. Glencassley, Sutherland- ilrs. Flower (p). 92. 10. 21. 1,

shire.

465

3 6, 9. luvermoriston, Inverness- A. H. Cocks (p). 4. 1. 25. 2-5.
shire. (4. 1. 25. B. 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.

(P)-

2 6. Knoydart, Inverness-shire. Sir Herbert Maxwell 98. 12. 2G. 1.

(p). 99. 2. 9. 1.

1. Arisaig, Inverness-shire. Dr. E. Hamilton (p). 2. 6. 3. 4.

?. 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). 8G. 7. 2. 13.

1. Scotland. 114b.

Felis silvestkis taetessia 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, Paune 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-buff. 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.

Skull. — 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
particulai'ly 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 INIorena, 1 ; Goto Douana, Huelva, 5 ; Andalucia (no exact
locality), 2 (skulls).

2 H

466

CARNIVORA

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468 CAENIVORA

Hemarlis.- — The Andalusiaii form of Felis silvestris is well
characterized by its large teeth aud rather dark colour.

1. Goto Dofiana, Huelva, A. Chapman (c & p). 7. 6. 4. 1.

Spain. (Type of subspecies.)

1. Goto Donana, Huelva. A. Chapman (p). 8. 3. 8. 1.

S, 9. Andalucia. (Chapman.) Dr. E. Hamilton (p). 2. 6. 3. 2-3.

FELIS SARDA Lataste.

1885. [Felis libijcn] var. sarda Lataste, Aetes See. 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
sxxv, p. 266, January, 1896 (Sardinia).

1910. Felis ocrcata sarda Trouessart, Faune Mamm. d'Europe, p. 101.

Type locaUtij. — Sarrabus, Sardinia.

Geographical 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 dor.sal line slightly elongated and
stiffened, forming a faint ridge or rudimentary mane ; dark
markings 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 noticeal)ly 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 group. Back of ear
yellowish clay-colour, the extreme tip black. Feet pale, dull
buff above, black below. Inner surface of hind legs bright
ochraceous-lDuff, 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 buUte 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 OF FELIS SILVESTRIS
AND F. SARDA.

Locality.

Number.

Sex.

lis:?

Three
lower
cheek-
teeth.

Lower
molar.

F. silvestris silvestris.

France: Manonville, Muerthe-|
et-Moselle . . /

95.11.5.1

S

18-2

—

—

Salavon, Haute-Marne

95. 11. 5. 2

6

18-2

21-2

8-8

Caterille, Haute - Ga- 1
roune . . . /

8. 7. 15. 1

£

17-6

20-2

8-0

Germany : southern Germany .

lliU

6

16-6

18-8

8-0

II 11

1143/

<5?

17-2

—

—

Austria-Hungary : Baranza

8. 7. 7. 10

6

18-8

21-8

8-4

Bulgaria: Varna

Andersen

S

17-8

21-0

8-2

Greece : near Athens

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

Fort William .

99. 2. 9. 1

6

17-4

21-0

8-0

Inverness-shire

Zool. Soc.

S

18-0

21-0

8-2

,,

4. 1. 25. 2

i

18-6

21-2

8-0

,,

4. 1. 25. 3*

i

16-8

21-0

8-0

1)

4. 1. 25. 4

6

18-0

21-0

80

,,

98. 12. 26. 1

S

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 Donana, Huelva .

7.6.4.1*

6

19-8

23-6

9-4

I) ji

2. 6. 3. 2

6

19-8

—

—

,, ,,

2. 6. 3. 3

9

19-8

23-4

10-0

„ .

8. 3. 8. 1

—

20-0

22-8

9-2

F. sarda.

Sardinia: Sarrabus

88. 12. 1. 1

6

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.M., Turin, Genoa,
and Lataste).

6. Ogliastra, Sardinia. {Dr. O. Thomas (p). 0. 5. 24. 1.

WoUcrstortf.)

i. Sarrabus. ' Marquis G. Doria(p). 88.12.1.1.

1. Sardinia. Purchased (Linneea, 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 distrihution. — Crete.

Diagnosis.- — Similar to Felis .^arcla, 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-bulf 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 ochraceous-buft' 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 buflfy
clay-coloui', 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.

Skidl and teeth not known.

Specimens 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. Kingd., Systematic catalogue inserted between

pages 32 and 83. (Type by tautonymy Lynx vulgaris Kerr =

Felis lynx Linna3us.)
1821. Lynceus Gray, London Med. Ftepos., xv, p. 302, April 1, 1821 {Felis

lynx LinnEeus).
1829. Pardina Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, i,

p. 53 {Felis pardina Temmiuck = Lynx par dellus Miller).

LYNX 471

1834. Lynchus Jardine, Naturalists' Library, Mamm., ii, p. 274 (Felis lynx
Linnaeus).

1857. Lynx Blasius, Saugethiere Deutschlands, p. 161 (Sub-genus of Fclis).

1867. Cervaria Gray, Proc. Zool. Soc, London, p. 276 (Lyncus pardinns =
Lynx pordcUus Miller; type fixed by Miller and Rehn, Proc.
Boston Soc. Nat. Hist., xxx, p. 199, December 27, 1901). Not of
Walker, 1866.

1903. Euccrvaria Palmer, Science, N.S., xvii, p. 873, May 29, 1903 (Substi-
tute for Cervaria Gray).

Type species. — Felis lynx Linnaeus.

Geographical distrihution. — Northern fore.sted portions of both
Old and New Worlds ; in Europe soutli to the INIediterranean
coast and west to the Atlantic (jierhaps within historic times to
Great Britain).

Characters. — Similar to Felis but without small upper pre-
molar, form heavier and less typically feline, ears tufted.

HemarJcs. — 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 part
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. pardellns, p. 475.

LYNX LYNX Linniieus.

1758. [Fclis] lynx Linnaeus, Syst. Nat., i, 10th ed., p. 48 (Sweden).

1792. Lynx vulgaris 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 mclina Kerr, Anim. Kingd., Systematic catalogue

inserted between pp. 32 and 83 ; described, p. 157 (Banks of the

Volga near Ca/.an, Russia).
1798. Flelis] borealis Thunberg, Beskrifning pa Svenske Djur, Mamm.,

p. 14 (Heavy forests of northern Sweden).
1798. Felis kattlo Schrank, Fauna Boica, i, p. 52 (Bohemia).
1820. Fclis lyncula Nilsson, Skand. Fauna, i, p. 14 (Wooded and moun-
tainous regions of Scandinavia).
1825. Felis lupulinus Thunberg, Denkschr. k. Akad. Wissensch., ^liinchen,

IX, p. 189 (Northern Scandinavia).
1825. Felis vitljnnus Thunberg, Denkschr. k. Akad. Wissensch., Miinchen,

IX, p. 192 (Near LTpsala, Sweden),

472

(JARNIVORA

1829. Felis virgata Nilsson, Ilium. Fig. Skand. Fauna, pis. 3 and 4
(Wooded and mountainous portions of Sweden).

1857. Felis lynx Blasius, Saugethiere Deutschlands, p. 173.

1870. Lynx ccrvaria Fitziuger, Sitzungsber. kais. Akad. Wissensch., Wien,
Math.-Naturwiss. Classe, "lx, Abth. i, p. 208 (Not Felis cervaria
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 portions 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 Vjelow 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
everywhere densely woolly.

Colour. — Upper jiarts 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 live
longitudinal rows, but spotting never so conspicuous as in Ljjnx
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 I'egion whitish ; contrast of whitish area sometimes
heightened by a dark shade along its lower border. Underparts
ranging from whitish to pale blufi". Feet intermediate between
back and belly ; front of forearm 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. Underfur usually brownish or buffy to
extreme base, but occasionally grey.

Skidl. — The skull of Lynx lyna: difi'ers in general aspect from

LYNX 473

that of Felts mlvestris in 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 Fells sihestris. Other general features so nearly as in the
wild cat as to require no detailed description. A few details,
however, are peculiar. Auditory bullse relatively smaller than in
Felis sihestris, the space between them fully as wide as bulla,
the surface less closely applied to mastoid and pai'occipital
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 j^m^ or
space between roots of j^wt^ 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 mj is distinctly oblique and
armed with a minute though evident basal cusp slightly above
the rudimentary cingulum. All the cheek-teeth are relatively
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 : —
NoBWAY : 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). ~^

Bemai-hs. — 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.

Museum.
1. Norway. Zoological Society's 69. 10. 19. 16.

Menagerie.

475

LYNX PARDELLUS Miller.

1827. Felis pardina Temminck, Monogr. de Mamm., i, ]}. 116 (Near

Lisbon, Portugal). Not Lynx pardina Oken, 1816.
1907. Lynx pardella Miller, Ann. and Mag. Nat. Hist., 7th ser., xx, p. 398,

November, 1907 (Goto Douana, Huelva, Spain). Type in British

Museum.
1910. Lynx {Eucervaria) parddla Troucssart, 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 line 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 sjsots 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-
parts 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 hea\y black band 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 parallel 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 huffy 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.

SJiuII. — The skull of Lynx pardeUus differs conspicuously from
that of L. lynx and Felis 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 representing the
external pterygoid j^rocess 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
interparietal.

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
Ml nearly as vertical as in Felis silvestris, and without secondary
cusp or evident cingulum. Crowns relatively a little narrower
and more trenchant than in Lijnx lynx, but less so than in the
wild cat, their length relatively as great as in the lynx.

Measurements. — Two males from Goto Donana, 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 Doiiana, 8 ; Andalucia, no exact locality, 1
no exact locality, 1.

jRemarhs. — 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. Widdring- 42.2.26.1.

ton (c & p).

2 6. Sierra Morena, Cordova. Lord Lilford (p). 89. 8. 27. 1.

94. 6. 11. L

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 Donana, Huelva. J. P. Gassiot (c & p). 72. 10. 26. 1-2.
head, st. Coto Donana, Huelva. A. Williams (c & p). 3. 3. 16. 1.

2 6,2 9. Coto Donana, Huelva. A. Chapman (c & p). 4. 12. 12. 1-2.

8. 3. 8. 2-8.
(4. 12. 12. 2. Tijpe of species.)
1. Coto Donana, Huelva. B. F. Buck (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 Mola (Boll. Soc. Zool. Ital., Roma, 2d ser., ix,
p. 48, 1908) as Lynx sardiniae. The more important characters

478

CARNIVORA

•

a

0

a

^ ^ ^

^

g ^ S

P-^ :::::;:;:; r

^ s ^

^

"

" " "ja

!M)

.Q

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r

OD

'H Tji

0

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10

-* in

tH

^^^^^^^^^^

Ol

000

0

0 0 0 CM cq Tt( (M 0 CC' 0

A'.iE(]!XBi\;

^

CM 0 10

COOOCJiOOCMCTioOiH

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'^i -H rH

'^tH

co-*'*coco-*'*coco^

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CO

9 9 1

00
C5

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CMC:3COC3CD^fMC.liO<M

0

t-

ooooaoaoco05C3coooc3

0

00 "O

lO

0 OCMOOOOOOO

03

lO -* 0

10

CO loOt-t-CJlCMCOt-O

IIIO^SBM

CD

O 10 CD

«3

in mininioomincD

CO

^00

0

0 OOOOCMO'*!

CO

<N CM (N

?5

(M CM (M CM CM (M C>^ (M

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479

DENTAL MEASUREMENTS OP LYNX LYNX AND
L. PABDELLUS.

Locality.

N limber.

Se.\.

Upper

car-

nassial.

rm3

and

2im*

together.

Lower
cheek-
teeth.

Lower
J car-
nassial.

Lynx lynx.

1

Sweden

J 1034 \
1 U.S.N.M.J

(5?

18-6

80-8

37-6

16-0

...

B.M.

juv.

—

29-0

—

15-6

Switzerland

1 668.32 \
\ Geneva /

ii

19-4

31-0

37-2

15-0

„

f 223.37 \
\ Geneva J

9?

18-6

29-4

—

—

. .

f 603.64 -1
\ Geneva j

juv.

17-0

-

-

14-0

Lynx pardellus.

Spain : Old Castile

94. 6. 11. 1

9

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 Dofiana .

72. 10. 26. 1

6?

14-6

24-4

29-6

12-8

.,

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

9

14-2

24-2

28-8

12-4

»

152618

6?\

14-6

24-6

28-8

12-S

" " • 1

152619

6 1

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 grey ; 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. Olires Linnaeus, Syst. Not. i, 10th ed., p. 5G (part).

1827. Bodentia Griffith, Cuvier's Animal Kingdom, Mamm., iii, p. 61.

Geographical distrihution.— 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 unguiciilate ; dentition consisting
of -J 3j 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.

Bemarks. — 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
manmials. They are all relatively small animals, the largest
member of the group, the South American Hydrochoerus hi/dro-
choeris, attaining a weight of only about 50 kg., while the largest
Eui-opean species, the beaver, is less than half this size. In
structure they show variety exceeded by the Insectivora 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 Euroj^e.

KEY TO THE EUROPEAN FAMILIES OP 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) Leporidv, p. 484.

Upper incisors 1-1, the enamel confined to the front
surface ; incisive foramina moderate or small,
distinct ; bony palate longer than mesopterygoid
space ; fibula not articulating with calcaneum.

* INIembers 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

RODENTIA

Anterior portion of zygomatic arch formed chiefly

by the jugal bone ; infraorbital foramen small.

Skull without postorbital processes ; molars

rootless ; tail broad and flat (Beavers) Casioridn', p. 947.

Skull with postorbial processes ; molars rooted ;
tail terete.
Fore and hind leg not joined by a fold of

skin (Squirrels) Sciurid.T, p. 897.

Fore and hind leg joined by a fold of skin

acting as a parachute (Flying Squirrels). Pctauristid.r, 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) Hystricidn-, 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 foramen 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 ; skuU wedge-shaped,
the sux^raoccipital sloping forward to
middle of brain-case, its area much
greater than that of parietals (Mole-
rats) Spalacidn', p. 887.

External form not specially modified for
underground life ; skull not wedge-
shaped, the supraoccipital vertical or
nearly so, confined to back of brain-case,
its area much less than that of parietals.
Cheek-teeth (in European genera) |- ;

caecum absent (Dormice) Muscardinidx , p. 549.

Cheek-teeth (in European genera) '^ ;

csecuni present Miifid.r, p. 591.

Molars j)i'ismatic, hypsodont, usually
rootless, their crovras flat (Voles

and Lemmings) Microtinx, p. 610.

]\Iolars tuberculate, brachyodont,
rooted.
Tubercles on crowns cf upper molars
arranged in two longitudinal

series Cricetlme, p. 592.

Tubercles on crowns of upper molars
arranged in three longitudinal
series Murinx, p. 791.

IJUPLICIDENTATA 483

ARTIFICIAL KEY TO EUROPEAN MEMBERS OF
RODENT FAMILIES.

(Based exclusively on the teeth.)

Incisors ~^^; cheek-teeth ''[^ Lrporichi-, p. AS-i.

Incisors \'_\ ', cheek-teeth not more than '^'•
Cheek-teeth 'j^^.

Crowns of mandibular cheek-teeth with con-
spicuous median concavity Sciurid;>\ p. 897.

Crowns of mandibular cheek-teeth without

conspicuous median concavity Petauristid.T, p. 940.

Cheek-teeth less than ^"j.
Cheek-teeth j^.

Cheek-teeth brachyodont, rooted Muscardinichr , p. 549.

Cheek-teeth hypsodont, rootless.

Enamel foldings transverse only Casfor'uh-, p. 947.

Enamel foldings transverse, oblique and

longitudinal Hystricid.-r, p. 542.

Cheek-teeth less than ^.

Cheek-teeth .p, Zapod'uhv,-^. 5.35.

Cheek-teeth Z~^^.

Enamel foldings sigmoid Spalacidx, p. 887.

Enamel foldings not sigmoid Mtiridx, p. 591.

Cheek-teeth hypsodont, prismatic Microtinx, p. 610.

Cheek-teeth brachyodont, tubercular.
Tubercles of upper molars arranged

in two main rows Crlcetinx, p. 592.

Tubercles of upper molars arranged

in three main rows Marinve, p. 701.

Sub-Ordkr duplicidentata.

1811. Duplicidentata lUiger, Prodr. Syst. Mamm. et Avium, p. 91

Geographical distribution. — Essentially that of the oi'der
Modentia except that it does not include Madagascar or Australia.
One genus now almost universally distributed in the warmei" 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 tlie 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.

ItemarJcs. — The well defined sub-order* Dtiplicidentata contains

* Not improbably an order distinct from the Rodentia, as there is
reason to l)elievo 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 rabbits, is represented in Europe.

Family LEPORID^.

1821. Leporidsi Gray, London Med. Repos., xv, p. 304, April 1, 1821.
1857. Leporina Blasius, Siiugethiere Deutschlands, p. 409.
1897. Lagid.v Schulze, Helios, xiv, p. 82.

Geograplncal distrihution. — As in the sub-order DiipUcidentata.

Characters. — Palatal bridge formed chiefly by the maxillary
bone ; supraoccipital processes present ; auditory bulla' rather
small, not inflated with spongy tissue ; clavicle rudimentary ;
hind legs elongated.

B,emarhs. — The members of the family Leporidse are now-
arranged in ten genera,* two of which occur in Europe.

KEY TO THE EUROPEAN GENERA OF LEPOEIDyE.

jMesopterygoid 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) Onjdolagiis, 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) Lcpus, p. 495.

Genus ORYCTOLAGUS Lilljeborg.

1758. Lepns Linnaeus, Syst. Nat., i, 10th ed., p. 57 (part).

1857. Lepus Blasius, Siiugethiere Deutschlands, p. 410 (part).

1867. Cunicnlus Gray, Ann. and Mag. Nat. Hist., 3rd ser., xx, p. 225

(Not of Brisson, 1762, or Wagler, 1830). Sub-genus of Lepus.
1874. Oryctolagus Lilljeborg, Sveriges og Norges Ryggradsdjur, i, p. 417

(Sub-genus of Lepns).
1899. Oryctolagus Major, Trans. Linn. Soc, London, 2nd ser., ZooL, vii,

p. 514, November, 1899. Genus (part).
1904. Oryctolagus Lyon, Smithsonian Miscell. Coll., xlv, p. 402, June 15,

1904 (Genus).

Tjijje sjjecies. — Lepus cuniculus Linnaeus.

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 limV) uf which (except in certain domestic forms)
is always free from cranium, persistence of sutures surrounding
interparietal, and the strongly narrowed choana?, the width of
which, immediately behind palate, is noticeably less than least
length of palate. Teeth essentially as in Lcjms, 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 filled 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 maxillai'y 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.

BemarJcs. — The genus Ori/dolagus contains the domestic
rabbits and the wild rabbit of central and southern Europe. In
addition to the more technical characters of the group its
members are readily distinguishable from the other Leposidae,
with which they are naturally associated, by the blind, helpless
condition of the young at birth. |

ORYCTOLAGUS CUNICULUS Linna^US.
(Synonymy 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 mediterraneus excepted), the hind
foot in dry specimens seldom exceeding 90 mm., the occipito-
nasal length of skull in lai'gest 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 brown ;
a distinct, dull buffy nape patch ; ear essentially concolor with
back though more finely grizzled, without distmct 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 Sylvilagus and perhaps also
in some of the less known genera.

486 RODENTIA

and inner surface of legs whitish except for dark inguinal
patches and collar.

External characfcrs. — 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 stiffened 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 nmch 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. Mammse : p 1 — 1,
a 2-2 = G.

Colour. — Upper parts a coarse 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 llidgway No. 6) basal area (13 mm.), the extreme
base somewhat paler ; (2) a brownish band (6 mm.) vaj-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 Lcpus. Nape patch a clear brown
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 bufty 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 gi-eyish 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-buft'. On hind legs the whitish
area extends over dorsum of foot to extreme tip of toes, though
sometimes suffused with the bufiy brown of sides of feet. On

ORYCTOLAGUS 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 cur\ es 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 squai-ish, 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 pi'ocesses descending below level of lip of large
foramen magnum, and closely applied to posterior surface of
auditory bullK. Floor of brain-case marked transversely by
persistent suture between basioccipital and basisphenoid ; 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 occiput, 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
l)ull;e ; height including tube nearly twice transverse diameter.
Interorbital region flat, somewhat depressed, its width about
equal to that of rostrum a little in front of middle ; postorbital
j)rocess 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
flrst cheek-tooth noticeablj' 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.
Oriictohi'jiis ciiniciihis. N;it. siz

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 breadth
about one-thii-d length. Least longitudinal diameter of palate
about two-thirds or three-quarters distance l^etween alveoli of
anterior premolars. External pterygoid plate well developed.

Teeth. — First upper incisor with root extending somewhat
more than half way from aveolus to suture between premaxillary
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 l)order ; posterior
face with wide, shallow longitudinal concavity, the beveled edge
abruptly angled near middle. Second 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 with
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 antei'ior 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 equal 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 KODENTIA

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.

Memarhs. — The well-known European rabbit needs no special
comparisons with other members of the fauna. In diflerent
parts of its range it is represented by two local races.

KEY TO THE SUBSPECIES OF OBYCTOLAGUS CUNICULUS.

Occipitonasal length of largest skulls, 78 to 82 mm. ;

hind foot, 83 to 93 mm. (Central Europe) 0. c cuniculus, p. 490.

Occipitonasal length of largest skulls, 71 to 77 mm. ;

hind foot, 72 to 82 mm. (IMediterranean region,

Azores, &c.) 0. c hiixleiji, p. 491.

Oryctolagus cuniculus cuniculus Linnteus.

1758. [Lc2ms] cuniculus Linnseus, Syst. Nat., i, 10th ed., p. 58.

1837. ? Lcpus vcrniciilaris Thompson, Tho Athenseum, p. 468. Nomen

nudum (Ireland).
1843. ? Lcpus vcrmicula Gray, List Spec. Mamm. Brit. Mus., p. 128.

Nomen nudum.
1857. Lcpus cuniculus Blasius, Siiugethiere Deutschlands, p. 426.
1867. Cuniculus fodicns Gray, Ann. and Mag. Nat. Hist., 3rd ser., xx,

p. 225, September, 1847 (Substitute for cuniculus). Type in

British Museum.
1904. Olryciolagus] cuniculus Lyon, Smithsonian Miscell. Coll., xlv,

p. 406, June 19; 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. — Size 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.

Pleasure iiients. — 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 > ear, 70 and 70.
Adult female from Lezayre, Isle of jNIan : head and body, 437 ;
tail, 66 ; hind foot, 91 ; ear, 70. Two adult females fi-om
Ingelheim, Rheinhessen, Germany : head and body, 407 and 418 ;
tail, 68 and 62 ; hind foot, 90 and N7 ; ear, 7(3 and 73. For
craneal measurements se3 Table, p. 492.

Specimens examined. — Forty-six, from the following localities : —
Ireland : Belfast, 3 ; 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 tamo) of Germany. That the wild rabbit of Germany was
considered by Linnrous as tho 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 ;
Pangbourno, Berkshire, 2 ; Croydon, Surrey, 2 ; Orleston, Kent, 1 ; Kent, 1
(skull) ; Ditchling, Sussex, 4 ; Seatowu, 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,Rheinhessen,2; Strassburg, 2 ; south Germany, 3
(skulls).

Remarlcs. — The wild rabbit of Ii'eland, 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.

6, 9. Belfast, Down. Marquis of Dufferiu 5. 2. 22. 1-2.

and Ava (p).

?. Dumdrum, Tipperarv. Major P. J. Waldron 5. 1. 15. 1.

(c & p).

6, 9. Kilmanock, Waterford. G. Barrett-Hamilton 9. 12. 15. 5.

(p). 11. 1. 2. 109.

skull. Ireland. C. Darwin (p). 68. 2. 19. 92.

skull. Shetland Islands, Scot- C. Darwin (p). 68. 2. 19. 96.

land.

?. 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 Ciinicidus fodiens Gray.)

6, 9. Pangbourne, Berkshire. Sir James Clark, Bt. 97. 2. 14. 1-2.

(c & p).

6, 9. Croydon, Surrey. C. H. B. Grant (c & p). 11. 1. 3. 398-399.

6, 3 9. Ditchling, Sussex. Guy Dollman (c & p). 8. 9. 25. 1-4.

6. Orleston, Kent. F. N. Garrard (c & p). 90. 1. 5. 1.

skull. Kent. C. Darwin (p). 68. 2. 19. 98.

2 9. Ingelheim, Rheinhossen, C. Hilgert (c). 8.6.15.10-11.

Germany.

3. S. Ciermany. Dr. A. Giinthcr (c). 59. 9. 6. 44-46.

Oryctolagus cuniculus huxleyi Haeckel.

I874. Lcpus 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 Troucssart, Faune Mamm. d'Europo,

p. 215.

Type locality. — Island of Porto Santo, Madeira.

GeograpMi-al 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

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494

RODENTIA

animal) ; colour more finely grizzled, the back showing less
tendency to be distinctly clouded with black.

Measuremenis. — 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
from Goto 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 tvi>va 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, oft' Candia, Crete (type of niosulus) : 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 : Goto Dofiana, 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, INIinorca, 2.

France: Poulx, Gard, 5; near Nimes, Gard, 14 (Mottaz) ; Digne,
Basses-Alpes, 8 (Mottaz).

Italy : Island of Capraja, Genoa, 5 (B.il. 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).

Remarks. — In colour the Mediterranean rabbit nearly agrees
with the central European form, but when series of skins are
compared it is at once seen that the smaller animal is the more
grey and finely grizzled oi the two, its back seldom distinctly
clouded with black, a condition that is usual in true cunicnlux.
The actual elements of the colour are essentially the same in the
two, but in hnxleiji the sub-terminal annulations of the longer
hairs are slightly paler than in the central form. Except for
the dift'erences 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 rabbits 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 j^rove to be exactly similar to the common
Mediterranean form.

5. Porto Santo, Madeira C. Darwin (p). 68.2.19.9.3,104,

Islands. 107, 111, 117.

1. San Miguel, Azores. F. du Cane Godman 73. 3. 21. 1.

(c & P).

c?, 9. San Miguel. Major F. A. Chaves G. 3. 25. 1-2.

(c & p).

LEPUS -195

^. San Miguel. Major F. A. Chaves 0. 3. 25. 3.

(c & p).

2. Goto Dofiana, Huelva, J. P. Gassiot (c & p). 72. 10. 2G. 3-4.
Spain.

6, 9. Goto Donana, Huelva. A. Chapman (c & p). 2. 3. 3. 1.

2 6,29. Seville. (Dr. A. Ruiz.) 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. Gastrillo do la Reina, G. S. Miller (c). 8. 8. 4. 122.

Burgos.

<J. Selva, Majorca, Balearic 0. Thomas (p). 1. 3. G. 3.

Islands. {M. Eiutort.)

2 9. Inca, Majorca. 0. Thomas (p). 1. 3. 0. 5 G.

(M. Eiutort.)

S. Alcudia, Majorca. 0. Thomas (p). 1. 3. G. 4.

(M. Riutort.)

i, 9. San Cristobal, Minorca. O. Thomas and R. I. 0. 7. 1. GG-67.

Pocock (c & p^

3 <5, 1 9. Poulx, Gard, France. O. Thomas (p). 8. S. 10. 135-1.38.

(C. Mottaz.)
1. Gapraja 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. cnossiits Bate.)

Genua LEPUS Linnteus.

1758. Lepus Linnseus, Syst. Nat., i, 10th ed., p. 57 (type by tautonymy,
L. timidus Linnasus).

1828. ? Lagos Brookes, Catal. Anat. and Zool. Mus. of Joshua Brookes,

p. 54 (Nomen nudum).

1829. Chionobates Kaup, Entw.-Geseh. u. Natiirl. Syst. d. Europ. Thier-

welt, I, p. 170 (variabilis and horealis).
1857. Lepiis Blasius, Siiugethiere Deutschlands, p. 410 (part).
1867. Eulagos Gray, Ann. and ]\Iag. Nat. Hist., 3rd ser., xx, p. 222,

September, 18G7 (mecliterraneus and judx'm).
1899. Lepus Major, Trans. Linn. Soc, London, 2nd ser., Zool., vii, p. 514,

November, 1899.
1899. Eiikpiis Acloque, Paune de Prance, INIammifferes, p. 52 (europi.Tus

and variabilis).
1904. Lepus Lyon, Smithsonian Miscell. Coll., xlv, p. 389, June 15, 1904.

Type species. — Lepus timidus Linnajus.

Geo<jrapliical 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 choame 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.

BemarJcs. — As now restricted the genus Lt^'pua contains about
eighty named species. Eight of these occur in Europe, most of
them represented by several geographic forms.

KEY TO THE EUROPEAN FORMS OF LEPUS.

Tail, including pencil, much shorter than hind
foot, its upper surface white or clouded with
hrowu 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 Ridgwav) ; winter pelage never

entirely white (Ireland) -.. L- hiberniciis, 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. timidus, p. 522.

Occipitonasal length of adult skulls ranging
from 95 mm. to 103 mm. (Scandinavian

Peninsula) i- t-timidus, 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. f. 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.vieclitcrraneus,-^. 513.

LEPus 497

Hiud 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 cinnamou-

rufous of Ridgway ; dorsal surface of foot

and wrist marked with pure white L. granatensis, p. 615.

Ear from crown about Da mm. (Basses -

Pyrenees) L. gratmtcneis iturissius,

p. 518.
Ear from crown, 105 to 115 mm.

Buffy tints pale, approaching the cream-
buff of Bidgway (Central and south-
ern Spain) L. grunatemis granatensis,

p. 516.
Buffy tints rich, approaching the ochra-
ccous-buff of Ridgway (North-western

Spain) L. granatensis gallaecixis,

p. 517.
Outer side of thigh not brighter than back,
its colour ochraceous, buffy, 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 10 mm. below tip ; skull with
unusually slender rostrum (Parnassus

region, Greece) L. parnassius, 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 126, 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) A. europeeus ti-anssylcanicvs,

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 parts a light cream-
buff ; size very large, hind foot about
155 mm. (Eastern Clermany 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.

■1 K

498 KODENTIA

Hind foot 124 to 147 mm. ; occipito-
nasal length of skull 93 to 102 mm.
(Central).
BuSy tints rather pale, approaching
the cream-buff of Ridgway (Ger-
many, Denmark, France, except

southern portion) L. e. eiuopaeuf, p. 502.

Buffy tints rich, approaching the
ochraceous - buff of Ridgway

(England) L. c occidentalis, 504.

Hind foot 113 to 125 mm. ; occipito-
nasal length of skull 89 to 94 mm.
(Southern).
Nape patch well differentiated ; base
of ear consj)icuously buffy be-
hind ; underfur not yellowish

(Pyrenees) L. c. pyrenaicus, 50G.

Nape patch scarcely indicated ; base
of ear not conspicuously buffy
behind ; underfur yellowish
(Corsica, Sicily, southern Italy) L. e. cvrsicaaiis, p. 507.

LEPUS EUROP.ffi:us Pallas.

(Synonymy under subspecies.)

Geoijraplilad iJintrihution. — Central Europe from Gre^t Britain
to Russia and from the Baltic south to the Pyrenees, Italy and
Greece.

Diagnosis. — Tail moderately long, its length including terminal
hairs about etjual 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 bufi'y 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 Ijlack 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 Ori/ctolagus.
Digits similarly H— 1, their proportional lengths as in the smaller
animal, but claws somewhat more flattened. Mammae : p 1 — 1,
rt 2 - 2 = 6.

Colour. — Upper parts buliy, nearly clear (m 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.
Rurap 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 grej'ish white suffusion.
Head essentially like body but with the darker and lighter
colours more finel}^ 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 clear buffy sometimes tinged with grey, the buffy clearer
and brighter just below terminal 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, M'hich is very pale Ijuff 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
Hne grizzle of the same elements as those of back, the edge
conspicuously fringed with clear buffy except at black-rimmed
tip ; posterior outer surface buffy 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
patch 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 j^'iler than outer surface (cf. L. meditcrraneus).
A clear buffy or ochraceous patch in inguinal region, the two
patches sometimes meeting in median line. Feet buffy, darker
on the toes, paler proximally, but never marked with pure white
(cf. L. yranatensis) ; soles an indefinite buffy grey.

The grey winter coat, when io occiu's, 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 Lrpus europsens and some form of varying hare.f

Skull. — The skull of Lejms europseus resembles that of L.
iimiduH 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. eurvpaeus the occipitonasal
length is usually more than 93 mm. (92 to 105 mm.), while in
L. tiviidux (except in the large L. t. iimidus) it is seldom more

* The absence of the white edging at tip of tail is nearly always an
indication 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. (86 to 93 mm.). Brain-case relatively narrower
and deeper than in Lepiis timiclus, and parietal region less
flattened, the dorsal ijrohle 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 interorbital region. Posterior
extremity of malar and relative length of suture between malar
and zygomatic process of scpiamosal as in L. timidus. Anterior
poi'tion of zygoma usually though not always diftering from that

Fig. uriA.
Lepuf: euro/}Kiix. Nat. size.

of L. timuhis in the greater width and relati^"ely 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 Ltytis tlmidns, its ventral profile a little less
convex and anterior border less rounded off, a pecularity not
easily described but which when once .seen is readily appreciated.
Terth. — In general the teeth resemble those of Oryctolagiis
cunindus, difTering in a few slight details only. Upper incisor
with groove on anterior face decidedly nearer internal border

LEPUS 501

than external border ; posterior face with longitudinal concavity
barely indicated ; root as short as in Oryctolagus, but 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.

etiropxus. Xat. size.

teeth not extending so far into orbit as in Ori/dolagus and in
Lepus timidiis, 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 Ori/ctolagns, but broader than in Lepus timidus ;
posterior section of this tooth as well as that of the succeeding
molariform teeth (including ik^) relatively larger than in
Oryctolagus.

502 EODENTIA

Semarks. — 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 surface 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 eueop.eus eup.op.fx's Pallas.

1777. [Le2nts] timidns Ersleben, S3'st. Regni Anim., i, p. 325 (Not of

Linnaeus, 1758).

1778. Lcjms europxus Pallas, Nov. Spec. Quaclr. Glir. Ord., p. 30 (Bur-

gundy, France).
1783. Lepus timidus Schreber, Siiugthiere, pis. ccxxxniA and ccxxxniB

(Germany). Not of Linnaeus, 1758.
1789. Lepus curop:vus Schreber, Saugthiere, iv, p. 865.
1801. L[cpus\ t\jnudus] albus Bechstein, Gemeinn. Naturgesch. Deutsch-

lands, I, 2nd ed., p. 1096 (Thiiringen, Germany).
1801. /^[epeis] t[imidus] flavus Bechstein, Gemeinn. Naturgesch. Deutsch-

lands, I, 2nd ed., p. 1096 (Thiiringen, Germany).
1801. L[€pus'\ t[imidus] nigcr Bechstein, Gemeinn. Naturgesch. Deutsch-

lands, I, 2nd ed., p. 1097 (Thiiringen, Germany).
1820. Lepus medius Nilsson, Skand. Fauna, i. p. 224 (Zealand, Denmark ;

based on specimens in grey winter pelage).
1857. Lepus timidus, b. ]Mitteleuropaischc Form, Blasius, Siiugethiere

Deutschlands, p. 417.
1859. '? Lepus campicola Gervais, Zool. et Paleont. Francaises, 2nd ed..

p. 47 (Nomen nudum based on the common hare of France).
1867. ? [Lepus timidus'] coronatus, rufus, cinereus, nigricans, mactilaius,

albus and nigcr iPitzinger, Sitzungsber. kais. Akad. Wissensch.

Wien, Math.-Naturwiss. Classe, lvi, p. 161 (Austria). Nomina

nuda.
1875. [Lepiis timidus'] var. argcnteogrisea Konig-Warthausen, Yerzeichniss

der Wirbelthiere Oberschwabens, Siiugethii;re, p. 86 (Uhn, Wiirtem-

berg, Germany). Probably based on an individual in grey winter

coat.
1910. Lcjms europ);i'US and L. medius Trouessart, Faune Mamm. d'Europe,

p. 219.

Type loccd'djj. — Burgundy, France. f

Geographical distribution. — Central Europe from Germany to

* The Lepus curop;rus carpatlioriim of Hilzheimer (Zool. Anzeiger, xxx,
p. 512, August 14, 1906) from the Carpathians may represent an eighth
race. The difficulty of clearly understanding the various local forms of
European hares is greatly increased by the frequency with which the
animals are transported from one region to another for the purpose of
restocking exhausted hunting grounds. Ij. e. transylvanicus has thus beeu
taken to Denmark, and L. e. occidentalis to Switzerland. Other instances
of the same kind will undoubtedly be found.

t Pallas refers to Daubenton's account of this animal for measurements
(Nov. Sp. Quadr. Glir. Ord., p. 2), and mentions no definite locality.
]?olonia and Pannonia are, he says (p. 5), inhabited by another 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.

Diaguoni.s. — Buft'y tints pale, approaching the cream-buff of
Ridgway ; rump usually not grey, always with evident darker
area continuous with black tail-stripe.

Colour. — Underfur (20 mm.) silvery white at ba.se (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 aloiig sides cream-
buff, darkening to a light, yellowish ochraceous-buff 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.

Skull and teeth, — 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.

Measure iitents. — Two adult males and an adult female from
Haslev, Zealand, Denmark : head and body, 660, 680 and 640 ;
tail, 80, 82 and 7i; 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, St. Gallen, Switzerland : head and bod)', 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 Gallen,
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 Plandei-s, 1.

France : Near Paris, 1 ; Ktupes, Doubs, 1 (Mottaz).

Germany: Brunswick, 3; Ingelheim, Rheinhessen, 1; Burg, near
Magdeburg, 1 ; south Germany, 6 (skulls).

AUSTEIA-HUXC4ARY: 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.).

Bemarlcs. — The typical form of Lepus europaeus is characterized
by large though not maximum size, and rather light, strongly
yellowish colour. It is probabl)' 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). 46. G. 2. 71.

504

RODENTIA

1. Slype, Flanders, Belgium. Andrew van Iseghcm 2.11.3.1.

(c & p).

?. Paris. A. Forsyth Major (p). 97. 2. 19. 1.

3. Brunswick, Germany. G. Barrett - Hamilton 8. 9. 29. 1-3.

i. Ingelheim, Rheinhessen. G. Barrett - Hamilton 11.1.2.110.

(Hilgert.) (p).

6. S. Germany. Dr. A. Giinther (c). 59. 9. G. 38-43.

Lepus europ^us occidentalis de Winton.

1898. Lepus europxus occidentalis de Winton, Ann. and Mag. Nat. Hist.,
7th ser., i, p. 152, February, 1898. Type in British ^Museum.

1906. L[e2ms] e\i(,ropxus] occidentalis Hilzheimer, Zool. Anzeiger, xxx,
p. 512, August 14, 1906.

1910. Lepus europiums occidentalis Trouessart, Faune !Mamm. d'Europe
p. 220.

Ti/jjc locah'fi/. — Herefordshire, England.

Gcoijrapliical distrihntiun. — 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* ami Switzerland.

Diagnositi. — Similar to Lejnis enropseiis rin-dpiens, but bufly
tints rich and dark, approaching the ochraceous-buff of Kidgway.

Colour. — Underfur as in L. eurojtseus europseun, 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-bluft", 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 buffy 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.

SJiull 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) from Pangbourne,

See Barrett-Hamilton, The Irish Naturalist, ilarch, 1898, pp. G9-7G.i

LEPUS 505

near Reading, Bei'kshire : tail, 75; hind foot, 135; ear from
crown, 99. For cranial measurements see Table, p. 510.

Specimens cxatnined. — Thirty-six, from the following localities : —

Scotland : Tulloch, Inverness, 1 (grey pelage) ; Eaith, 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 ; Pangbourne, near Reading, Berkshire, 1 (grey pelage) ;
Marley Common, Haslemere, Surrey, 1 ; Weston Sands, Somerset, 2 ;
Stokenhara 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.).

Hemarhs. — Though closely related to Lcjjus europseus europtmis,
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). 3.11.14.1.

Scotland.
6. Eaith, Fifeshire. W. R. Ogilvie-Grant 98. 12. 23. 2.

(c & p).
6. Kilwinning, Ayrshire. Hon. G. Montgomerie 92. 2. 15. 1.

(c & p).
6. Isle of ]\Ian. P. M. C. Kermode 9.5. 1. 19. 1.

(c & p).

2. Bangor. Carnarvonshire, G. W. D. Assheton 97. 3. 8. 1.

Wales. Smith (p). 3. 1. 28. 1.

1. Healey, Northumberland, Rev. H. H. Slater 0.2.24.1.

England. (c & f).

6- Swithland, Leicestershire. Earl of Lanesborough 92.1.29.1.

(c & p).
c?, 2 9. Thetford, Norfolkshire. Lord Walsingham (p). 98. 2. 11. 1-3.
2 st, Sandringham, Norfolk- H.M. King Edward 96. 5. 9. 3-4.
shire. VII. (p).

Moorhtimpton, 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. Sawbridge 0.11.16.1.

(c & P).

Haslemere, Surrey. S. J. Wigley (c & p).

Weston Sands, Somerset- Tomes Collection.

shire.
England.

Purchased (Lead-
beater).
Hon. L. Powys (c&p).

Giasford Castle, Armagh

Ireland.
South-east coast, Ireland. Miss E. Hope.
6, 2 9. St. Gallen, Switzerland. O. Thomas (p),

(E. H. ZolUkofcr.)

6. 9. 18. 1.

7. 1. 1. 175-176.

88. b.

76. 1. 25. 1-2.

6. 4. 12. 1.
2. 8. 4. 53-54.
4. 4. 5. 46-47.

506

Lepus europjEUS pyrenaicus Hilzheimer.

1906. Llcjnis] e[u7-opieiis] pyrenaicus Hilzheimer, Zool. Anzeiger, xxx,

p. 512, August 14, 1906.
1910. Lepus curopseus p)ijrenaicus Trouessarfc, Fauna ^lamm. d'Europe,

p. 220.

Type loralit;/. — Bagneres [de Luchon ?], Pyrenees, France.

GfO<jrapltieal flisfrlhnfion. — Pyrenean region, south-eastern
France.

Diagnosis. — Essentially like Lepus curojjcens europsevs, but
.smaller (hind footj 113 to 125).

Specimens examined. — Five, from the Department of Ariege, France,
and the neighbouring portion of Andorra.

B.i'inarlcs. — The tive specimens, though inijjerfect, do not
appear to be referable to any of the other forms of Lepus
cnropneus. They agree sufficiently well with pyrenaicus as
described by Hilzheimer.

2?, 1. Ax -les-Thermes, Ariege, V. Builles (c & p). 10.9.20.1-3.
France.

Lepus europ.eus meridiei Hilzheimer.

1859. ? Lepus meiidionalis Gervais, Zool. et Paleont. Fraucaises, 2nd ed.,
p. 47 (Nomen nudum, based on the hare of Languedoc and
Provence).

1906. Llcpus] eluropxus] meridiei Hilzheimer, Zool. Anzeiger, xxx, p. 512,
August 14, 1904 (Aveyron, France).

1910. Lepus europxus meridiei Trouessart, Faune Mamm. d'Europo, p. 220.

Type locality. — Department of Aveyron, France.

Geograpiliical (Vistrihutiuu. — South-central and south-eastern
France, northern Italy, Corfu. •

Diagnosis. — Size as in Lepus eiiropseus eiiropseus ; colour
))rowner and less buffy than in the typical form, and runi})
distinctly bluish grey, though less conspicuously contrasted with
back than in the large L. e. transsylvanicHS.

Measurements. — Four males from the neighbourhood of
Nimes, Gard, France, average and extremes: 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.

LEPOS '^07

Italy : Ceresole d'Alba, Turin, 1 (Turin) ; Porlezza, Como, 1 (Ghidioi) ;
Siena, 4.

Greece : Corfu, 1.

Bemarlx. — 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 S. Barcelonnette,Basses-Alpes, O. Thomas (p). 8. 8. 10. 133-134.

Prance. (C. Mottaz.)

2 i. Siena, Italy. (S. Brogi.) Dr. E. Hamilton. 98. 10. 2. 18-19.

6. Corfu, Greece. (C. ilfo/tei.) J.I. S. Whitaker (p). 8.10.1.51.

£. Corfu. C. Mottaz (c). 8. 11. 3. 23.

Lepus EUROPiEUS coRSiCANUs de Winton.

1898. Lepus corsicamis de Winton, Ann. and Mag. Nat. Hist., 7th ser., i,
p. 155, February, 1898. Type in British Museum.

1906. Llexms] m[editerraneus\ corsicamis Hilzheimer, Zool. Anzeiger, xxx,
p. 513, August 11, 1906.

1910. Lepus corsicamis Trouessart, Faune Mamm. d'Europe, p. 224.

Type locality. — Bastia, Corsica.

Geographical distribution. — Corsica, Sicily and Italy (Rome).

Diagnosis. — Size small, as in Lepus europeeus pyrenaicus (hind
foot, 114 to 125 mm., occipitonasal length of skull about 90 to
94 mm.) ; general colour more yellow than in true europeeus, the
underfur conspicuously buff.

Colour. — The colour does not differ appreciably from that of
Lepus europseus europseus, except that it is throughout somewhat
lirighter 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 sti'ong contrast by the black. Inguinal patches light
tawny. Rump dull ochraceous-buff, the bases of the hairs
l)luish grey (about grey No. 7 of Ridgway).

ShuU 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. T\vo 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 : —
Cobsica: Bastia, 1 (type).
Sicily : IMarsala, 2.
Italy : Rome, 1.

508 EODENTIA

BemarJcs. — The conspicuously yellowish underfur of this hare
is perhajw 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 (p). 78. 7. 3. 4.

{Type of subspecies.)
2 6. Marsala, Sicily. J. I. S. Whitaker (p). 98. 2. 9. 1-2.

i. Rome, Italy. (C. Coli.) G. Barrett-Hamilton (p). 8. 9. 30. 1.

Lepus europ.eus hybridus Desmarest.

1822. Lcpus hybridus Desmarest, .Mammalogie, p. 349 (Central Russia).
1842. Lcpus aquilonius Blasius, Amtl. Bericht xix Vers. Naturf. u. Aerzte,

Braunschweig, p. 89 (Central Russia).
18.50. Lepus nicdius Middendorff, Bull, de la Classe Phys.-Math. de I'Acad.

Imp. des Sci. Nat. de Saint-Petersbourg, i.x, p. 218 (part).
1857. [Lepus timidus] c. Nordostliche Form, Blasius, Siiugethiere Deutsch-

lands, p. 417 (part).
1898. Lcpus europneus (typical) de Winton, Ann. and Mag. Nat. Hist.,

7th ser., i, p. 150, February, 1898.
1906. L[epius] ni[edii(s] aquilonius Hilzheimer, Zool. Anzeiger, xxx, p. 511,

August 14, 1906.
1910. Lcpus mcdius aquilo7iius Trouessart, Fauna Mamm. d'Europe, p. 217.

l^i/pe JdCiditij. — Central Ru.ssia. 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 Lcpus cnropsens europseus
(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 Lcpus europseus europseus, the general effect
is noticeably lighter, owing to the greater length of the sub-
terminal cream-butf 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 (excejjt for a dull ochraceous-buft" area under
and behind eye and another at base of whiskers), in strong
contrast with the grizzled bufl'y 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. curopetus 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
(Forstzoologie, 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 from Litlmania : hind foot, 155; ear
from crown, 125.

Specimens examined. — One, from Lithuania.

llrinarJcs. — 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. Barey.) Branieki JIuseum (e). 94. S. 7. 30.

Lepus europ^us transsylvanicus Matschie.

190L Lepus transsylvanicus Matschie, Sitz.-Ber. Gesellsch. Naturforsch.

Freuncle, Berlin, p. 236 (Taslau, Roumania).
1906, L[epus'] e{i.iropxus\ transsylvaticiis Hilzheimer, Zool. Anzeigcr xxx,

p. 512, August 14, 1906.
1910. Lepus europxKS frajissylvaticiis Trouessart, Fauue JMamm. d'Europe,

p. 221.

Ti/pe locality. — Taslau, Roumania.

Geographical distrihiition. — From Roumania southward through
the Balkan Peninsula to the Peloponesus. Exact limits of range
unknown.

Diagnosis. — Size nearly as great as in Lepus europseus
hi/hridus (hind foot about 150 mm., occipitonasal length of skull
in largest individuals 105 mm.) ; general colour dark and
brownish, much as in L. europaeus 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.

Bemarlcs. — 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
europseus 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.

6. Visoko, Bosnia. Dr. O. Reiser (c & p). 7. 10. 15. 3.

i, 9. Surmancipolje, Herze- Dr. 0. Reiser (c & p). 7. 10. 15. 1-2.

govina.
2 c?. Opposite Corfu, Albania. J. I. S. Whitaker (p). 8.10.1.49-50.

(C. Mottaz.)

510

KODENTIA

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LEPUS CRETICUS Barrett-Hamilton.

1903. Lepus creticus Barrett-Hamilton, Ann. and Mag. Nat. Hist., 7th ser.,
XI, p. 126, January, 1903. Type in British ]Museum.

1905. Lepus europnus creticus Bate, Proc. Zool. Soc, London, 1905, ii,
p. 322, April 5, 1906.

1910. Lepus creticus Trouessart, Faune Mamm. d'Europe, p. 222.

Type locality. — Crete and Cephalonia.

Geographical distribution. — Crete.

Diagnosis. — General colouration light brownish essentially as
in Lepus europxus transsylvanicus and L. parnassins, but paler,
the rump conspicuously grey ; size rather small, about equal to
that of L. ruropseus 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-l)uff, 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-browir to dull ochraceous-buff. usually grizzled
with Ijuffy white, the former predominating. Head a fine grizzle
of same elements as body colour, the pale markings Ijefore and
behind eye silvery whitish though not very extensive. Cheeks
essentially like crown, distinctly grizzled and with 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 po.sterior 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.

Skull 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. curojjseus. Anterior
portion of zygoma essentially as in L. europieus, 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.

BemarJi's. — While resembling the brown races of Lepui^
europseus the Cretan hare is readily distinguisliable 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. O. Jones, R.N. (p). 2. 11. 9. 1.

{Type of species.)

6. Kanea, Crete. Miss D. Bate (c). 5. 12. 2. .34.

6, 4 9. ]\It. Vuno, Cephalonia, J. I. S. Wbitaker (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 mediterraneus Wagner, Miinch. Gelehrt. Anzeiger, p. 439,
March 17, 1841.

1857. [Lepus timidus'] a. Siideuropiiische Form, Blasius, Siiugethiere
Deutschlands, p. 417 (part).

1898. Lepus mediterraneus de Winton, Ann. and Mag. Nat. Hist., 7th
ser., I, p. 154, February, 1898.

1906. L[epus"\ m\editcrraneus'] typicus Hilzheimer, Zool. Anzeiger, xxx,
p. 512, August, 1906.

1910. Lepus mediterraneus Trouessart, Fanne Mamm. d'Europe, p. 224.

Type locality. — Sardinia.

Oeographical distrihntion. — 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 graiiatcnsis, 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 Lepms graaatensis 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-bufi
of Ridgway), strongly incroaching on white of underparts, and
continuous with inguinal patches and inner surface 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 thei-e is no evident line of demarcation between the
two surfaces of the legs. Rump slightly more bufFy 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 suftused with ochraceous-buff, the pale eye-ring and the
sjjot 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 parts. Ear dark and dull,
the colour pattern ill-defined, and the narrow whitish rim of
outer border thrown into unusually strong contrast ; anterior
outer surface and dark area on inner surface prout-brown slightly
grizzled with dull buify ; grey of posterior outer surface extending
nearly to base of ear ; black area normal ; no grey on inner
surface, this region, excejat on dark area, everywhere ochraceous-
bufF like fringe of inner border. Collar dull ochraceous-buff
irregularly washed and varied with cream-bufF. Interramial
i-egion 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
whitish ochraceous-buff like inner surface of fore legs.

SlcuU 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 europseu>:, though the auditory bullje
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.

Bemarhs. — 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,'i. Cagliari, Sardinia. (Mazza.) Genoa Museum (e). 98.5.12.1-2.

515

LEPUS GRANATENSIS Rosenliauer.

(Synonymy under subspecies.)

Geographical distribution. — Iberian Peninsula and Balearic
Islands. Exact northern limits of range not known.

Diagnosis. — Size rather small, about as in Lepus corsicanus,
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. viediterraneus,
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 ; dorsal surface of feet and wrists marked with
pure white ; lengthened whitish hairs on sides very conspicuous.
Mamma; as in L. europaeus.

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 buffy 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 europxus. 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-i'ing
about as in L. europieus, but pale area before and behind eye
barely indicated, and no distinct rusty suffusion below eye ;
muzzle clear dull butiy ; a darker area at base of whiskers. Ear
as in Lepus europseus, but colour pattern more strongly defined
and black terminal 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
foi'e 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.

Shull 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 EODENTIA

appreciable on inspection, and evidently of much importance in
a group showing such remarkable uniformity in cranial characters.
The auditory bullae 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 fiat 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.

Bemarks. — The Iberian hai"e 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.

Lep.us granatensis granatensis Rosenhauer.

1856. Lepus granatensis Eosenhauer, Die Thiere Andalusiens, p. 3
(Granada).

1867. \_Lcpus'] Uispanicus Fitzinger, Sitzungsber. kais. Akad. Wissensch.
Wien. Math.-Naturwiss. Classe, lvi, p. 161 (Substitute for grana-
tensis. " Natt." cited as authority).

1897. Leptis meridionalis Graells, jMem. Real 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[cpns] c[uropxus'] granatensis Hilzheimer, Zool. Anzeiger, xxx,

p. 512, August 14, 1906.

1907. Lepus granatensis granatensis MiUer, 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 llidgway 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 hairs (30 mm.) greyish at base, then
black to tip, each with a conspicuous, sharply defined sub-terminal
annulation (4 mm.) of pale cream-buft". Clear area along sides
(ill-defined) between cinnamon-rufous aud 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 con.spicuous
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 maximvim
size for the species, occipitouasal 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.

G. S. Miller (c). 8. 8. 4. 120-121.

A. Chapman (c & p). 8. 3. 8. 8-9.

Lord Lilford (p). 95. 3. 3. 11-14.
(95. 3. 3. 12. Ti/i)e of L. lilfonli de W.)

H. F. Witherby 0. 2. 17. 1.

(c & p).

Col.J.H. Irby(c&p). 96.7.29.1.

A. Chapman (c&p). 8. 3. 8. 16.

Lord Lilford (p). 75. 2. 11. 3.

0. Thomas (p). . 1. 3. 6. 2.

Lepus granatensis gall.ecius Miller.

1907. Lepus granatensis galhecius 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'Europe,
p. 223.

Type locality. — La Coruna, Province of Coruiia, Spain.

Geo(jrapMcal distribution. — At present known from extreme
north-west Spain only.

Diagnosis. — Similar to Lepus granatensis (jranatensis, but colour
throughout dai'k and rich, the ground colour of back nearly the
ochraceous-buflf 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

2?.
2.

Castrillo, Burgos, Spain.
Coto Doiiana, Huelva.

2 (?, 2 9.

Seville. (Dr. A. Ruiz.)

1.
1.

Las Marismas, near

Seville.
Granada.

juv.

Jerez, Cadiz.

1.
1.

Spain.

Selva, Majorca, Balearic
Islands. (M. Biutort.)

518 RODENTIA

slightly less extensive than in L. (jranaiensis granatensis. Collar
a dark 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
cranial measurements see Table, p. 521.

Specimens examined. — Two, one from La Corufia, the other from Vigo.

1. Vigo, Pontevedra, Spain. Lord Lilford (p). 82. 12. 8. 1.

i. La Corufia, Corufia, 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.

Type JocaJitij. — Basse.s- Pyrenees, near Biarritz, France. The
type was probably bought in the Biarritz market, but a note
published l)y the collector in the Field, Lxx.xix, January 30,
1897, p. 135, refers to the animal as occurring on the (Spanish
side of the border.

Oeographical disfrihiition. — 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 dor.sal 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-Pyr6n6es, J. E. Harting (p). 97. 6. 15. 1.

France. (Greig.) {Type of subspecies.)

519

LEPUS PARNASSIUS Miller.

1903. Lepiis iMrnassius Miller, Proc. Biol. Soc. Washington, xvi, p. 145,
November 12, 1903. Type in U.S. National Museum.

1908. Lepus eluropxus] parnassius Hilzheimer, Jahreshefte des Vereins
fiir vaterl. Naturk. in Wiirttemberg, p. 395.

1910. Lepus europxus parnassius Trouessart, Paune ]\Iamm. d'Europe,
p. 221.

Type localify. — Agoriani (Agokgianne), north side of Lyakuia
(Parnassus) Mountains, Greece.

Geographical distribution. — Known only from the type
locality.

Diagnosis. — General appearance much as in Lejnis 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 jji'^ in the two specimens
known ; ?»3 with the two enamel cylinders broadly communi-
cating.

Colour. — General colour of upper parts a coarse grizzle of
black and pinkish bufl', the latter slightly in excess, much less so
than the cream-buff of the corresponding region in Lepus europseus.
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 gi-izzled,
and with a faint blackish wash below ear. An indistinct greyish-
buft' eye-rmg and loi-al 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-
buft'area 10 ram. 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, fiank patches, and throat, dull ochraceous-buff. Under-

520

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522 KODENTIA

parts and inner surface of letjs white, much suffused with
ochraceous-buff on front legs. Collar very wide, essentially Hke
sides of body, but without black and with a decided tinge of
wood-brown.

ShiU. — The skull differs from that of Lepus europaens 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 othei- respects the skull shows no tangible characters to
distinguish it from that of L. europeeiis*

Teeth. —The teeth differ from those of all other European
hares in the absence, in the type, and a very young specimen
(in L. eiiropaeus 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 Liuna?us.

(Synonymy under subsxiecies.)

Geographical cUstrihution. — 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 buUse 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
Ridgvvay) in summer, white or pale grey in winter, the extreme
tip 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 nuich as in Lcpus europseiis
except for the shorter tail and ear, and more woolly fur.
Mammre : jj 1-1, a 3-3 = 8.

Colour. — Summer j^elage : general colour of upper parts
between hair-brown and sepia, inconspicuously grizzled with
cream-bufF, the sides not so dark as back, the rumj) sooty blackish ;
cheeks, muzzle and outer surface of legs tinged with buflf'y ; 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 pure 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 bufFy 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 frequently 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 ujjper parts sometimes more yellowish,
owing chiefly to the wood-brov/n of underfur brightening toward
ochraceous-buft". 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 bufl'y 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 Ridgway) 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 Mdiitish, 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.

Shdl. — In general form the skull resembles that of OrijctoJagus
cunicnlus. The size is, however, decidedly greater than in either
of the European 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 Ori/cfolaf/us, 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 Orydolagus
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 Orydolagus, 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 OrydoJacjm 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 ecjual to distance from latter
point to front of incisive foramen. Auditory bulla essentially as
in Orydolagus, 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. curopxus, its inner re-entrant angle appearing to

525

¥10. 103.
Le^ms timidus scoticus. Nat. size.

526

]ie on inner border of crown rather than on inner side of
anterior border. Anterior section of first lower premolar louder
and narrower than in the related species. Other teeth not
certainly distinguishable from those of L. europieus 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 alveolar length
of tooth-row.

Bema7-ks. — As a group the vary-
ing hares are readily distinguishable
from the other European 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 pecu 1 iar it ies . That these more
fundamental characters indicate a
higher degree of specialization than
that attained by Lejms europseus has been shown by AVinge
(Danmarks Fauna, Pattedyr, p. 58, 1908). Lcpus 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 diiierentiated from the
continental and British forms.

Fig. 104.
Lepus timidtis. Teeth.

Lepus timidus timidus Linna'us.

1758. [Lcjms'] timidus Linnseus, Syst. Nat., i, 10th ed., p. 57.

1777. [Lepus iimidus'] alpinus Erxleben, Syst. Regni Anim., i, p. 328 (the

varying hares in general, from Switzerland to Greenland). Not
of Pallas, 1773, or of Erxleben, p. 337.

1778. Le])us variabilis Pallas, Nov. Sp. Quadr. Glir. Ord., p. 2 (Renaming

of timidus).
1778. [Lcpus] 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[epus'\ septentrionaUs Link, Beytriige zur Naturgesch., Bd. i,

Sttick 2, p. 73 (Substitute for variabilis).
182C. Lep7is borealis Nilsson, Skand. Fauna, i, p. 211 (Substitute for

variabilis).
1827. [Lipiis timidus] /8 alba Billberg, Synopsis Faunae Scandinaviae, p. 7

(Northern Scandinavia). Not L. timidus alius 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 canescens Nilsson, Ofversigt af Kongl. Vetensk.-Akad. For-

handl., Stockholm, i, p. 133 (Renaming of sylvaticus).

LEPUS 527

1857. Lepus variabilis b. Form der Mittelregion und der Alpen, Blasius,

Saugethiere Deutschlands, p. 424 (part).
1910. Lepus timidus and L. timidus collinus Trouessart, Faune Mamm.

d'Europe, p. 216.

Type locality. — Upsala, Sweden.

Geographical distribution. — 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.) ; Kronon, Vesterbotten, 7

(U.S.N.JI.) ; 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, Skftne, 1 (U.S.N.M.) ; no

exact locality, 1.

Norway : Hammerfest, 1 ; Enebek, 1 ; Bidsvold, Christiania, 2 ;

near Christiania, 1 ; Jsederen, Stavanger, 1 ; Holme, Mandal, 2.

Remarlcs. — 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 i-egarded 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 Jtederen. 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 sylvaticus
and canescens of Nilsson were based on the southern animal,
while the horealis and collinus of the same author were ajiplied to
the northern form.*

Stockholm Museum (e). 46. G. 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.

R. J. Cuninghame (c & p). 8. 8. 9. 38-89.

1.

Sweden.

6.

2 6.

Enebek, Norway.
Eidsvold, Christiania.

i.

Christiania.

i.
i juv, 9.

Jsederen, Stavanger.
Holme, Mandal.

♦ A good account of the two races is given by Collett, Norges Pattedyr,
pp. 58-69, 1911.

528 RODENTIA

Lepus timidus varronis Miller.

1857. [Lepus variabilis] b. Form der Mittelregion und der Alpen, Blasius,

Saugethiere Deutschlands, p. 424 (part).
1901. Le^ms varronis Miller, Proc. Biol. Soc, Washington, xiv, p. 97,

June 27, 1901. Type in U.S. National Museum.
1906. Llepus] vi[edi>ts'] varronis Hilzlieimer, Zool. Anzeiger, xxx, p. 511,

August 14, 1906.
1906. Llcpus] m[edius'] breviauritus 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 mcclius varronis Trouessart, Faune Mamm. d'Europe, p. 218.

Type locality. — Hinzenberg, Grison.s, Switzerland.
Geoijraphical distribution. — Alps and adjacent mountain
chains, mostly at elevations above 1,300 m.

Diaqnosis.- — -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 greyisli wash ; anterior portion of outer border of ear
nearly always buffy grey ; 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.

Specimens examined ; — Twenty-four, from the following localities : —

Fbance : Col-de-Vars, Hautes-Alpes, 2 ; above Barcelonuette, Basses-
Alpes, 2 (Mottaz) ; Petit Bornand, Haute-Savoie, 2 (Mottaz).

Switzerland : Canton St. CVallen, 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.) ; Ceutovalli, Ticino, 1 (U.S.N.M.) ; San Lucio,
Val Colla, Ticino, 1 (U.S.N.M.).

Italy: Padola, Cadore, 2 (Turin); Dronero, Cuneo, 1.

Remarks. — The Alpine varying hare is a strongly characterized
race, which should perhaps be treated as a species distinct from
Lepus tivtidus. The winter coat appears to be always white or
incompletely white ; a special blue grey winter pelage like that
of '' sylvaticas" is unknown.

2 6. Col-de-Vars, Hautes-Alpes, 0. Thomas (p). 8. 8. 10. 1.31-132. [:

France. (C. Mottaz.)
9. St. Gallen, Switzerland. O. Thomas (p). 2. 8. 4. 52.

(E. H. ZoUikofer.)
rf, 2 9 CMmns.(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

Lepuh timidus scoticus Hilzheimer.

181G. ? Leptis albiis Leach, Syst. Catal. Spec. Indig. ]\Iamin. and Birds,
Brit. Mus., p. 7 (nomen nudum: ''White Hare"). Not Lepii^
timidtcs albus Bechstein, 1801.

183.3. Lepus albun Jenyns, Man. Brit. Vert. Anim., p. 3.5. Not Lcpus
timidus albus Bechstein, 1801.

18.57. \Lcpus variabilis] b. Form der Mittelregion und der Alpeu, Blasius,
Siiugethierc Deutschlauds, j>. 424 (part).

1900. Ijcpus timidus tiipicus Barrett-Hamilton, Proc. Zool. Soc, Loudon,
p. 88 (part).

190G. L'cpus'] vi[edius] scuticus Hilzheimer, Zool. Auzeiger, xxx, p. 511,

August 14, 1906.
1910. Lepus timidus scoticus Trouessart, Faune SJamm, d'Europe, p. 218.

Tj/pe locaUti/. — Nortlicrn Scotland.

Gedtjrajihii'al (listrihuiion. — Highl<and.s of Scotland ; range now
extended by artiticial introduction irregularly into Wales and
northern England, also in Ireland.

Diaguosii^. — Size even les.s than in Lepu^t tituidu-s rarronis, the
occipitonasal length of fully adult skulls ranging from 83 to
89 mm. ; ear .shorter than in varronis, its length from crown
80 to 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 Leinis 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 with collar. Winter pelage appaientlv
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 suffusing
the entire dorsal surface. It is i)roduced by the presence among
the white hairs of a few that ai-e entirely black, and others that
are black with a light cream-buff sub-terminal annulation as in
summer pelage. Underfur very pale buff, this colour ajjpeai'ing
at surface when hairs are disarranged. Ear with anterior outer
surface conspicuously grizzled (much more than in varronis),
though usually paler than in summer, the basal half often mostly
white. Sides of muzzle usually tinged with buffy.

Shdl and teeth. — The skull is immediately distinguishable
from those of the otlier Europe;in forms of Lcpus 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 Alt\'re, Morayshire : head and body, 501 ;
tail, 57 ; hind foot, 131 ; ear from crown, 90. Adult from New
Galloway, Kirkcudbright : head and body, 535 ; tail, 58 ; hind

2 M

530 RODENTIA

foot, 133 ; ear from crown, 77. For cranial measurements see
Table, p. 533.

Specimens examined. —Forty, from the following localities in Scotland
Borgie, Sutherland, 1 ; Nairn, 5 ; Forres, Elgin, 1 ; Altyre, ^Morayshire, 5
Dallas, Morayshire, 2; Morayshire, 3 ; GlengrouUie, 3; C'romlix, Dunblane
Perthshire, 8; Caparoch, Selkirkshire, 1; Thornhill, Dumfriesshire, 2
New Gallowav, Kirkcudbright, 2 ; Clonas, 4 ; no exact locality, 3 (B.M.
and U.S.N.M.").

Bemarhs.— While readily distinguishable from true Lepus
ihnidus 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. timiihis rarronis no sjjecial grey winter coat
is known.

Individuals supposed to be hybrids between this animal and
Lepus curopcBUS occidentalis are sometimes unusually heavy
examples of scoticiis, 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 scoticm, aud
its colour pattern nearly as in occidentnlis, though less distinct ;
the tail is short as in scoticm, 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.|

1.

Borgie, Sutherlandshire,

Rev. H. H. Slater

0. 2. 24. 3.

Scotland.

(c & V).

c5,2 ?.

Nairn.

Earl Cawdor (p).

98. 11. 21. 1-3.

99. 5. 1. 1-2.

1.

Knockie, Inverness-shire.

E. Hargitt (p).

8G. 9. 9. 3.

1.

Forres, Elginshire.

Sir W. Gordon Gum-
ming (p).

3. 1. 27. 1.

4 ^, 1 V.

Altyre, ^Morayshire.

Sir W. Gordon Gum-

93. 12. 4. 1-3.

ming (p).

94. 2. 15. 1-2.

3 St.

Morayshire.

Sir W. Crordon Gum-
ming (p).

97. 12. 27. 1-3.

2 St.

Dallas, Morayshire.

Sir W. Gordon Gum-
ming (p).

1. 1. 15. 1-2.

3.

GlengrouUie.

Earl Cawdor (p). ■

99. 3. 20. 1-3.

6 6, 2 ?.

Dunblane, Perthshire.

Capt. 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, Craigmyle, Aberdeenshire, Scotland.
No. G. 12. 20. 1. IMale, Capenooh, Thornhill, Dumfriesshire, H. S.
Gladstone.

t Swedish specimens combining the characters of timidus and euro-
p/eus have been described and figured by Lonnberg, Proc. Zool. Soc,
London, 1905, ii., pp. 278-287.

LUPUS 531

1. Thornhill, Dumfriesshire. H. S.Gladstone 7. S. 7. 1.

(C&1-).

2. New Galloway, Kircud- Col. Gordon Jlaitland 95. 10. G. 1-2.

brightshirc. (c & p).

6, 3 ?. Clonas. W. R. Ogilvie-Grant 99. 2. 17. 1-4.

LEPUS HIBERNICUS ]5ell.

1833. Irisli Hare Yarrell, Proc. Zool. Soc, London, p. 88. No technical

name.
1837. Lepiis hibernicus Bell, History of Brit. Quadrupeds, p. 341.
1857. [Lcpus variabilis] a. Form der warmeren Klimate, Blasius, Siiuge-

thiere Deutschlauds, p. 424 (part).
1900. Lcpus timidiis hibernicus Barrett-Hamilton, Proc. Zool. Soc,

London, p. 89.
1900. Lcpus timidus Intcsccns Barrett-Hamilton, Proc. Zool. Soc, London,

p. 89 (Douobate, Co. Dublin, Ireland). Type in British Museum.
190G. Llepus] t[i'midus'] hibernicus Hilzheimer, Zool. Anzeiger, xxx, p. 510,

August 14, 190G.
1910. Lcpus timidus hibernicus and L. timidus lutesccns Trouessart, Faune

Mamm. d'Europe, p. 216.

Tijpc local ifij. — Ireland.

(TCoyrajiJiieal i1isfriJ)utlou. — Ireland, chiefly in the more hilly
districts. Now introduced and established in iiortions of AVales
and Scotland (Carnarvonshire and the island of Mull).

Dia(/nosif<. — Differing from the other Euroijean members of
the Lcpus ti mill us group in its strongly russet colour and in the
jjartial or complete absence of the white winter coat. 8ize
noticeably greater than in its nearest geographical ally,
L. iiiuidun scoticus.

Co/o«r.— General colour of upper parts a finely and incon-
s]iicuoush' grizzled russet brown, approaching the russet and
wood-brown of Ridgway, brighter and more reddish on shoulders
and back, paler, though strongly russet, on crown and face,
duller and fading to ochraceous-buff" on cheeks, sides, and outer
surface of legs ; nape and back of neck to between shoulders
scarcely grizzled, the nape usually with a sufiusion of drab. The
colours on the back are arranged as follows : underfur (15 mm.)
light pearl-grey (about grey No. 10 of Ridgway) at base, the
terminal half of the hairs abruptly light russet ; longer hairs
(25 mm.) light pearl-grey at extreme base, becoming black at
level of russet portion of shorter hairs, each with a dull butt sub-
terminal annulation about 4 mm. long and a rather shorter black
tip ; a few still longer hairs (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 jiroducing a markedly clouded ett'ect. Tail
white, usually with a bluish tinge, the upper surface sprinkled
to a varying degree with blackish or brownish hairs, these
occasionally producing a decided grizzled effect, and never

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entirely absent. Ear grizzled ocliraceous-buff, rimmed with
black at tip, the inner surface becoming clear ochraceous-buff
sub-terminally, the posterior outer surface washed with grey,
and with a buffy or whitish border from base to black terminal
area ; anterior border occasionally whitish near middle. A
faintl}^ 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 with 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 ajipearing irregularly at surface.
Feet ochraceous-buft', duller and somewhat grizzled on meta-
podials, clear and brighter on toes. Soles an indefinite buiiy drab.

»S7i'm// and teeth. — The skull differs from that of Lipits
timidiis scoticus in the larger sizes attained by fully adult
individuals, the occipitonasal length in such specimens ranging
from 91 to 94 mm., while in the Scotch hare it seldom exceeds
88 mm. Similarly the mandible in Jiihernicus is usually more
than 70 mm. in length, while in Hcoticus 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.

Measurementx. — 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 (8.5-95). For cranial measurements see
Table, p. 533.

Specimens examined. — Twenty-nine, from the following localities: —

Ireland: Gosford Castle, Armagh, 2; Lisduff, Co. Cavan, 12; Eath-

rohbin, King's County, 2 ; Arthurstown, ^Vaterford. 1 ; Kilmanock,

Waterford, 8 ; no exact locality, 2. Donobale, Dublin, 1 (type of Intescens).

Walks : Yaynol Park, near Bangor, Carnarvonshire, 1 (introduced).

Heiiinrkn. — The Irish hare is strikingly different from the
other European members of the tiiuidvs group in its peculiar
russet colour. In size it noticeably exceeds the Scotch hare, but is
itself surpassed by the Scandinavian true timidus. The yellowish
specimens from Co. Dublin, to which the name hdcsccns 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, Hun. L. Powys (c & p). 70. 1. 25. 1-2.

Ireland.
4. Lisduff, Cavan.

4. Lisduff, Cavan.

Hon. B. E. B. Fitz-

7G. a. 26. 1-2

patrick (c & p).

70. 5. 8. 1-2.

Hon. B. E. B. Pitz-

70. 7. 12. 1-2

patrick (c & p).

70. 'J. 7. 1-2.

SIMPLICIDENTATA

535

4. Lisduff. Cavan. Hon. B. E. B. Fit/. 7G. 11. 18. 1-2.

Patrick (c & p). 77. 1. 18. 1-2.

2 6, St. Rathrobbiii, King's County. Col. M. W. Biddulph 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, Watorford. G. Barrett-Hamilton 99. 2. IG. 1-3.

(c & P).

2 .?, 9. Kilinanook, Waterford. G.Barrett-Hamilton 9.12.1-5.1-3.

(c & p).

t<, 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).

8ub-0rder SIMPLICIDENTATA.

1891. Simplicidentata Flower and Lydekker, Mammals, Living and
Extinct, p. 448.

Geof/rajihical tlistrihiition. — Same as that of the order
liodentia.

Characters. — Upper incisor.s 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 jif ; 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.

Kemarlcs. — This suborder contains the vast majority of exist-
ing rodents, among which about twenty families ai-e now recog-
nized. Eight of these occur in Europe (for kej's see pp. 481-483).

Family ZAPODID^.

1875. Za2)odidn' Coues, Bull. U.S. Cxeol. and Geogr. Surv. Terr., 2d ser., i,

p. 253.
1901. Zapodidiv Lyon, Proc. U.S. Nat. Mus. xxxiii, p. 666, May 2, 1901.

(ieographical distrihufion. — Boreal portions of both hemi-
spheres, east in Europe to Norway.

Chnrarters. — Skull and teeth with a general resemblance to
those of the Dipodidie though less highly modified ; but infra-
orbital foramen similarly large and with distinct supplemental
canal along inner boi'der, and jugal in contact with lachrymal ;
hind legs much less specialized than in the related family, the
foot with live distinct metatarsals.

Hemarks. — The family Za])odidse, though related to the

J536 ■ RODENTIA

Dipodidse* is readily distinguished l)y the more primitive murine
character of the skull and hind foot. It is divisible into two
main groups, the Zapodinse with two genera in North America
and one in China, and the Sicistinse with a single genus peculiar
to the Old World.

Sub-Family SICISTIN.^.

1857. Murina Blasius, Siiugethiere Deutscblands, -g. 299 (part).

1887. Smintlii (section of DiiMdini) Winge, Jordfundne og nulevende Gna-

vere {Rodent ia) fra Lagoa Santa, ilinas Geraes, Brasilien, p. 122.
1901. Sminthiiw Lyon, Proc. U.S. Nat. Mus. xxxiii, p. G66, 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 n(jt elongated,
cheek-teeth tuberculate.

Semarhs. — The sub-family Sicistinse, a strictly Old World
group less .sj)ecialized 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 {Mus

subtilis Pallas).
1839. Sminthus Natbusiiis in Nordmaun, Demidoff's Voyage dans la

Russie Meridionals, iii, p. 49 (S. longer Nordmann).
1857. Sminthus Blasius, Siiugethiere Deutscblands, p. 301.
1901. Sicista Allen, Proc. Biol. Soc. Washington, xiv, p. 185, December 12,

1901.

Type species. — Mus subtilis Pallas.

Geographical distribntion. — 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 Ishmds see Major, Zool. Garten, xlvi, pp. 129-134,
May 1905.

Characters. — As in the sub-family Sicistinpe. Dental formula :
i y , pm J ||, wt ;| :| =18; crowns of cheek-teeth with complicated
enamel pattern which does not form transverse ridges ; larger
teeth with four well developed, tubercles.

Beiitarls. — 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 tuljercles 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.

* Waterbouse, Ann. and ]Ma^. Nat. Hist., x., p. 203, November, 1842.

537

KEY TO THE EUROPEAN SPECIES OP SICISTA.

Least depth of rostrum behind incisors about equal to
breadth of rostrum in same region ; crown area of »i'
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 «t' about three times that of premolar ; sides
essentially concolor with back (Hungary, Denmark,
south-eastern Norway) S. iri^onrz, p. 539.

SICISTA LORIGER Nathusius.

18-40. Sminihus lorigcr 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).

Type Jocalifi/. — Odessa, Russia.

Geographical distrihution. — South-western Russia, eastern
Roumania, Bulgaria ; limits o£ range not known.

Diagnosis. — Least depth of rostrum behind incisors about
equal to width of rostrum in same region ; crown area 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 musculus, but size less (about as in 3Tus spicilegus), 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 lai'ge 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 sliort hairs ;
basal lobe conspicuously tufted ; anterior border of no.stril
thickened, a small wart beneath it ; space between nostrils
narrow, crossed by a well developed vertical ridge ; upi:)er lip not
grooved in front ; feet more slender than in Mus musculus 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 nail ; third digit longest, fourth slightly
shorter, second extending to middle of third, fifth just beyond
middle of fourth, j^ads as in Mus musculus ; hind foot with
relative lengths of digits as in Mus musrulus, 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. Mammae : p 2-2, i 2-2 = 8.

Colour. — Back a peculiar light brown perhaps best described
as a bufty, 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 defined black stripe
about 2 mm. wide thrown into relief posteriorly by a distinct
lightening of ground colour along its edges. Sides 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 l>urty 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 bufty brown rather lighter than belly, the
toes a little more pallid. Tail obscurely bicolor, sepia above,
dull cream-buft' below. Longer hairs oi back black to extreme
tip ; basal colour everywhere slaty.

SlaiU and tcefh.— The skull diflers from that of ,S'. trizoua

FIG. 105.
SIclKta longer. . Xat. size.

Ki.j. Kin.
Sicista longer. Teeth, x 0.

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 discrepancy in size between premolar
and first molar more pronounced, the crown area or the smaller
tooth about one-fourth that of the larger.

Measuremeiifs. — 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.

Spcci7nctis examined. — Sis, five from Dobrudscha (B.]\I. and U.S.N.]\I.),
and one from Bulgaria, no exact locality (Lataste).

Bcmarliif. — 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-
i^niishable by the contrast between the colour of sides and back.

1. Dobrudscha, Roumania. Purchased (Pruliere). 8G. 4. 2. 4.

SICISTA TRIZONA Petuuyi.

1S57. Smi)itliu.f: vagus Blasius, Siiugetbiere Deutschlands, p. 302 (probably
not Mils vagus Pallas).

1882. Mas trkoniis Petenyi, Termeszetrajzi Fiizetek, v, p. 103 (Hungary).

1882. Mus intersoniis Petenyi, Termeszetrajzi Fiizetek, v, p. 103 (Alterna-
tive for trizonus).

1882. Mus intcrstriatus Petenyi, Termeszetrajzi Fiizetek, v, p. 103
(Alternative for trizonus).

1882. Mus tripartitus Petenyi, Termeszetrajzi Fiizetek, v, p. 103 (Alterna-
tive for trizonus).

1882. Mus virgulosns 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 subtilis Trouessart, Faune ]Mamm. d'Europe, p. 205 (probably
not Mus subtilis Pallas).

Tyjyp local! tji. — Hun gaiy .

Geographical didrihntion. — Hungary ; Denmark ; * south-
eastern Norway.f Details of distribution ver\' imperfectly
known.

Diai/iwsis. — 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 buify tinge, this more noticeable on cheeks and sides of
neck than elsewhere ; back veiy inconspicviously " lined " with
black. The individual hairs of the underfur are slate-colour at
base (5 mm.), the tips (1 -5 mm.) bufly raw-umber ; longer hairs
(12 mm.) black, the tips (2 mm.) colourless and with a silvery
gloss. Median dorsal line with clear black stripe aboiri 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 oc.hraceous-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-buft' below.

Skull. — In general appearance the skull suggests that of

* .SVe Winge, Danmarks Fauna, Pattedyr, p. 65, 1908.
t See CoUett, Norges Pattedyr, pp. 68-77, 1911.

540 RODENTIA

Mils 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 bulhe rather large, well
inflated, their greatest transverse diameter along squarely truncate
anterior border. Interorbital region much brtmder than rostrum,
nearly flat, its edges faintly angular. Zygoma abruptly spreading
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 lachr^'mal, the nasal branches 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 lai'ge, flat throughout, extending at full width
conspicuously behind last mtjlars, rather more than half of its
surface formed by the palatine liones ; a noticeable transverse
ridge at level of middle of )ii'\ 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 antero-internal angle of bulhe ; ectopterygoid well
developed, forming a conspicuous, nearly horizontal jjlate, the
rather large pterygoid fossa, therefore, shallow. Mandible slender,
with no special peculiatnties, 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 ci'own terete, with
high, narrow antei'o-external cusp, and low ridge-like internal
cusji, the three cusps separated by two evident furrows. Its ai-ea
is about one-third that of succeetling tooth. First upper molar
3-rooted, the general outline of ci'own 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
(jutward Ijetween 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
crowns longer and less squarish in outline, and pattern of folding
reversed, the intermediate folds extending inward instead of out-
ward. Third lower molar larger than the cori-esponding upper
tooth.

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542

RODENTIA

Measurements. — Adult male from Vestervig, Denmark : head
and body, 65; tail, 87 ; hind foot, 16*4 ;^eai- 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.,
nortliern Hungary, 2.

Denmark : Vestervig, Thy, 1.

Remarl's. — 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. Zuborec, Hungary. Budapest Museum (e). 94. 3. 1. 73-74.

1 al. Tatra ]Mount dns. Zoological Society. 87. 1. 4. 1.

1 al. Tatra Mountains. Dr. R. Collett (p). 91. 1. 21. 3.

1 al. Vestervig, Thy, Den- Copenliageu IMuseum (e). 88. 1. 9. 1.
mark.

Family HYSTRICID^l

1821. Hystrid;f Gray. London Med. Repos., xv, p. 304, April 1, 1821.
1831. Hystricidn' Burnett, Quart. Journ. Sci. Lit. and Art, xxviii, 1829,
'p. 350, 1830.

Geographical disirilutiioii.. — Warmer poi'tions of the Old
World, Madagascar, Australia and the outer portions of the
Malay Archipelago excepted. In Euroj^e eontined 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 lai'ge, 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 (juills.

Memarhs. — The family i///.s//-/c/'?»' contains four or five genera,
one of which reaches the Mediterranean region of Europe.

Genus HYSTRIX Linnajus.

1758. Hijstrix Linnsus, Syst. Nat., i, 10th ed., p. 56 (criatatu by tautonymy).
1798. Hixtrix Cuvier, Tabl. Element, de I'Hist. Nat. der Anim., p. 170

(Modification of Hystrix).
186G. OJduceplialiis Gray, Proc. Zool. Soc. London, p. 308 {Acanfhiun

ciivieri Gray).

Type species. — Hysiri-r cristata Linna'us.

Geograpliicnl disfrihiition. — Afiica and locally in JNIediter-
ranean region of Europe.

Characters. — Inflation of facial region of skull maxinium fin-
the family, the contrast between depth at front of tooth-row and
that behind alveolus of incisor \ ery 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 ' \ pm — , m ;' ;'
= :20 ; tail short ; neck with conspicuous crest of long coarse
bristles.

RciiiarJis. — The genus Hi/sirix as here restricted contains the
African and Italian species only. Several forms have been
described, the status of which is imperfectly understood. One
occurs in Europe.

HYSTRIX CRISTATA Linu:eus.

1758. [Hi/stiix] cristata Linnaeus, Syst. Nat., i, 10th ed., p. 5G (near Eomc,

Italy).
1792. H\_ystrix\ cristata cnro^wa Kerr, Anim. Kingd., p. 213 (Renaming of

cristata).
18-39. ? [Hi/strix cristata] var. albi De Selys-Longchamps, Etudes de Micro-

mamm., p. 152 (nomen nudum).
1910. Hystrix cristata Trouessart, Faune Maram. d'Europe, p. 212.

Type localltij. — Near Rome, Italy.*

Geogrcqjhical distribution. — Northern Africa ; central and
southern Italy : Sicily. (Wrongly attributed to the Iberian
Peninsula.) f

Diagnosis. — General characters as in the genus Hi/sfr!x ;
size medium, the condylobasal length of skull about LSO mm.,
depth of skull at front of toothrow about 50 mm.

E.vternal characters. — Form heaA'y, the body appearing umch
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 tine 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 tifth 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 (Syu. Meth. Anim. Quadr., p. 209), whose
account is based on a specimen from the mountainous districts near
Rome (see Thomas, Proc. Zool. Soc. London, 1911, p. 14-1).

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
inedian 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 l)order 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
e.xtremities for the most part stiff and awl-like, but occasionally
th-awn out into a hair-like filament ; base of tail with stiff terete
quills like those of body ; tip Ijearing 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 Ijody 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. iii length, the general
effect of the quilled region black conspicuously spotted and
streaked with white.

SkuJl. — 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 Ijones (posterior margin
(jf 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

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-

FlG. 107.
Hystrix cristata. x I.

tiated as an evident region, its width greater anteriorly than
posteriorly, the entire surface of each parietal occupied by a
cleop 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 visil:)le 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 j^oints 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 : auditoi'y bulla
rather small, globular, the inflated portion longer than wide,
noticeably broader anteriorly than posteriori}-, the meatus with
a short though evident tube ; anterior border in contact with
hamular ; posterior border free from paroccipital process.
Interorbital region, the broadest portion of skull, highly arched
laterally, without evident ridges or depressions. Zygoma short
and heavy, not widely spreading, its anterior portion deep, the
posterior portion shallow, the lower border nowhere above
alveolar level thoiighsloping evidently ui^ward 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 7ir' ;
hamular short, obliquely expanded, the posterior, longer limb of

547

the expansion broadly 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 jji'ocess broad and low ; coronoid process small,
not rising conspicuously above level of posterior alveolar border,
widely sejjarated from articular process ; angular portion broad,
lying consjiicuously 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-posteiior diameter
jierceptibly greater than lateral diameter ; enamel surface yellow.
Upper incisor with root extending to immediately in front of
alveolus of premolar, where its position is indicated Ijy a slight
elevation on the surface of the bone :
anterior face of tooth slightly oblique,
nearly flat though a little rounded oti'
at edges ; posterior sui-face 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
producing any very noticeable swell-
ing on outer side of mandible, and
scarcely rising above level of molar
crowns ; anterior face of tooth verj-
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 impei'iectly
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
simple ; third curving backward and outward, its extremity
often becoming isolated as one or two minute secondary islands ;
lingual side of tooth with a single re-entrant fold usually directed
obliquely forward and often communicating with second or more
rarely with tirst outer fold. Pattern of premolar like that of

flG. lUS.

Hi/strix cristata. Teeth. X 1 \

548

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HYSTRIX 549

molars but sliowing more tendency to irre;ij;ulai'ities. Lower
teeth with the same pattern reversed, the outer side with one
simple, oblicjue, 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.

MeaHurenwnts. — Young adult (probably female) from Vizzir.i.
Sicily (apj)roximate) : head and body, 600 ; tail, 40 ; hind footj
75. Immature individual from Province of Rome, Italy (approxi-
mate) : head and body, GOO ; tail, 50 ; hind foot, 85. For cranial
measurements see Table opposite.

Specimens e.ramincd. — Seven, from the following localities : —
Italy: Near Rome, 4 (B.M. and U.S.N.M.) ; Vizzini, Sicily, 2; no
exact locality, 1 (Stockholm).

3. Rome, Italv. Genoa IMuseum (e). 97. 2. 22. 1-3.

2, juv. Vizzini, Sicily. J. I. S. Wliitaker (p). 3.4.21.1-2.

Family MUSCARDINID^.

1821. Mijoxidce (misprinted Myosidie) Gray, London Med. Repos., xv,

p. 303. April 1, 1821.
1857. Mijoxina Blasius, Saugetbiere Deutschlands, p. 285.

1898. Gliridx Lydekker, Zool. Rec. xxxiv. (1897), Mamm. p. 27, 1898.
Not of Ogilby, 1837.

1899. Mmcardlnidx Palmer, Science, N.S., x, p. 413, September 22, 1899.

Geographicnl distribution. — AVarmer 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 Maridye (p. 591) but
Ciecum absent, jugal bone larger and forming a more important
mechanical part of zygoma (in some genera approaching the
condition characteriatic of the Sciuridw), 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 .".ometimes obsolete, but referable to a
primitive 3-ridged pattern ; one premolar typically present in
each jaw.

Roiinrlcs. — Six genera of Mmcardinidse 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 Muridae

550

EODENTIA

by the presence <>f 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 MVSCARDINIDJi.

Outer margin of first and second upper molar with
two high cusps (occasionally one or two others
low and obsolete).
Crov/ns 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 Elionujs, p. 650.
Crowns of cheek-teeth slightly concave ; j)i'eiuolar
both above and below not noticeably cuspidate ;
tail uniformly haired, moderately distichous... Dijromys, p. 560.
Outer margin of crown of first and second upper
molar with five or sis low cusps.
Crowns of cheek-teeth very slightly concave ; first
and second upi^er 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 Miiscardimis, p. 588.

Genus ELIOMYS Wagner.

1843. Eliomys Wagner, Abhandl. k. Bayer, Akad. Wissensch. Miinchen,
Math.-Phys. CI., ill, p. 17G (Sub-genus of Myoxus).

1857. Eliomys Blasius, Silugethiere Deutschlands, p. 289 (Sub-genus of
Myoxus) part.

1890. Eliomys Reuvens, Die Myoxidae oder Schlaefer, p. 22 (Sub-genus of
Myu.vus) part.

Type ffjiccii'S. — Elioiiqis iinlduuriis ^\'agiier.

GcoyrapliicaJ distrihutinn. — Europe from the Atlantic coast
eastward into Asia Minor, and from northern Germany (Mecklen-
burg) south into northern Africa.

Vhararterti. — Skull deep, smooth and rounded, without evident
ridges or depressions in interorbital region ; brain-case deep, the
auditory bulhe large ; ectopterygoid well developed ; jugal short
anteriorly, not approaching lachrymal ; mandible with angular
portion fenestrate ; dental formula : i ^{|, pm ' ', m 't;; = 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 m^ and ni" with hve
transverse ridges, of which the third is incomplete ; tail not
uniformly haired throughout, the basal half terete, the terminal
portion slightly flattened.

Heiuarliit. — The genus Elioiiiys as now understood contains

ELIOMYS 551

about a dozen forms peculiar to the Mediterraneau and neigh-
bouring regions. Five of them occur in Europe.

KEY TO THE EUEOPEAN FORMS OE ELIOMYS.

Size large, hind foot often more than 30 mm,,
condylobasal length of skull in adults 36 to

39 mm. (Southern Spain and Portugal) E. lusitanicus, p. 560.

Size medium, hind foot never more than 30 mm.,

condylobasal length of skull in adults less than

36 mm.

Underside of tail normally * white from base

to tip.

Ear from meatus in adults 23 mm. or more

(Central and south-western Europe) E. quercinus, p. 551.

Ear from meatus in adults 20 to 22 mm.

(Balearic Islands) E. gyinjiesicus, p. 558.

Underside of tail normally f with white inter-
rupted by a black sub-terminal ring.
Condylobasal length of adult skulls 33 to
85 '6 mm.; brain-case slightly flattened,
the ratio of depth through bulla to
condylobasal length about 37 '5 (Sicily and

southern Italy) E. pallidiis, 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. sardus, p. 560.

ELIOMYS QUERCINUS Linnteus.

1766. [Mils] qiieycinus Linnaeus, Syst. Nat., i, 12th ed., p. 84. Germany.
1778. Mas nitcdida Pallas, Nov. Sp. Quadr. Glir. Ord., p. 88 (part ;

synonymy, not description).
1782. Myoxus nitela Schreber, Siiugthiere, pi. ccxxvi (text, iv, p. 833,

1787). Germany.
1857. Myoxus quercinus Blasius, Saugethiere Deutschlands, p. 289.
1890. Eliomys quercinu,s Reuvens, Die Myoxidae oder Schlaefer, p. 26.
1904. Eliomijs hortuaUs Cabrera, Bol. Real Soc. Espaii. Hist. Nat., iv,

p. 183, April, 1904 (Valencia, Spain). Type in Madrid JIuseum.

1907. Eliomys lunniUonii Cabrera, Bol. Real Soc. Espan. Hist. Nat. vii,

p. 225 (June-July), issued October, 1907. (El Pardo, near Madrid,
Spain.) Type in Cabrera collection.

1908. Eliomys quercinus Cabrera, Ann. and Mag. Nat. Hist. 8th ser., i,

J). 192, F-ebruary, 1908 {Jiortualis and hamiltonii united with
(j_ucrciniis).
1910. Eliomys quoxinus Trouessart, Faune Mamm. d'Europe, p. 136.

Type localitij. — Germany.

Geoijrapldcal distribution. — Central Europe from northern
Germany (Mecklenburg) to central Spain (Valencia) and northern
Italy ; west to the ^lantic coast.

Diagnosis. — Size medium (condylobasal length of largest
skulls rarely attaining 33 mm. ; hind foot usually 25 to 27 mcb.) ;

* 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 cliaraders. — 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 aljruptly 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-inuer
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.
Mammre : p\ — \ ; a\ — \; i 2 — 2 — S.

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 j^osterior portion becoming decidedly
broader ; a small but noticeable white spot immediately in front

* See Thomas, Proc. Zool. Soc. London, 1905, ii, p. 491.

ELIOMYS 553

of mecatus, this succeeded by a larger black spot just above base
of upper margin of conch. Ear dusky, sometimes inclining to
blackish along anterior 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 upi^er 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

KIG. 109.

Klvimys qncrcinus. 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 convex to interparietal across which it is again
flat ; ventral profile parallel with dorsal to front of tooth-row
then bent down to lowffit 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 deepei- anteriorly than posteriorly, its surface smooth
and without ridges other than a slight angularity along anterior

554

RODENTIA

purtion 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
upjier 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 antero-posterior diameter from one-
third to one-half transverse diameter ; auditory bulhe forming a
very conspicuous feature of brain-case when viewed from behind,
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 Iiullse laige, smoothly inflated, the area of each
IniUa 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 jarocess 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-
curyed lip in region nearest zygoma ; interorbital region slightly
hour-glass shaped, the upper surface flat or a little convex
laterall}-, the edges behind middle becoming somewhat angular in
old individuals ; zygomata rather weak, compressed, a little
expanded at middle, gradually spreading anteriorly, the greatest
breadth nearly at glenoid level, the lower border essentially
straight though sometimes a little concave ; anteorbital foramen
rather large, consideiably 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 foramen to fi'ont of alveolus of corre-
sponding premolar ; palate wider anteriorly than posteriorly,
the small jDortion 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 Ions; enouoh in fig. 100.

ELIOMYS 555

niecliaii spine, the slightly curved hamular in contact with audi-
tory bulla iit tip. External pterygoid plate evident though small,
forming 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 Math abrupt angle, above which the bone
is perforated by a conspicuous round vacuity about I'T) mm. in
diameter.

Teeth. — The teeth are moderately large relatively to size of
skull, the length of tooth-i"Ow contained about '2\ times in palatal
length ; tooth-rows wider apart anteriorly than posteriorly, the
alveolar length equal to distance between anterior roots of
premolars. Incisors with no special peculiarities, the anterior

KIG. 110.

Ktiomtjs querciims. f'heek-tettli. X ICl.

surface 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.j^ ; 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 largest, the crowns squarish, slightly wider than
long, the inner border a little rounded, the posterior border of
im? 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 succeedStl by a minute postero- internal cusp
early disappearing with wear ; outer border with two moderately
large subterete cusps, the second not so large as first (the
discrepancy greater in ni^ than in the other teeth) and two or

556 RODENTIA

three minute secondaiy cusps, one in front of first main cusp, the
others behind second ; surface of crown crossed by four complete
transverse ridges, the tirst, 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 postero-
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 cus])S and ridges, but crown
slightly smaller than that of wi\ 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 rii.j, to
become Isasin-shaped, their cusps and ridges less sharply defined,
those of III., often very ii-regular ; outer side of crown with three
low but well defined cusps, inner side irregularly crenulate in wt^
and ?»3, with two rather low but well defined cusps in jBj, the
230sterior 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
jHj ; 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.

Measurevicnts — Two adult females from Maredsous, Namur,
Belgium : head and body, 1.36 and 137 ; tail, 116 and 121 ; hind
foot, 28 and 29 ; ear from meatus, 24 and 24. Adult male from
I'Hospitalet, Aricge, France: head and body, 116; tail, 97-G;
hind foot, 25' 4; ear from meatus, 2-o. 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 (2.5-27);
ear from meatus, 23-6 (22-25). Five adult females from Silo.s,

Burgos, .Spain: head and body, 120 (115-130); tail, 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
meiisurements see Table, p. 562.

Specimens examined. — One hundrod and twenty, from the following
localities : —

Belgium : Maredsous, Namur, 2.

France : Pas-de-Calais, 2 ; I'Hospitalet, Ariege, 7 ; Porte, Pyr6n6es-
Orieutales, 2 ; Nimes, Gard, 14 (B.M. and Mottaz) ; Valescure, Var, S;
Chamonix, Haute-Savoie 3 (U.S.N.M.) ; Mounetier, Haute-Savoie, 1
(IMottaz) ; no exact locality, 2.

Spain : Cabanas, Corufia, 2 ; Jaca, Huesca, 4 ; Panticosa, Huesca, 1 ;
L^rida, 1; PajAres, Leon, 7; Silos, Burgos, 11; Castrillo de la Reina,
Burgos, 1 ; La Granja, Segovia, 4 ; Villalba, Madrid, 2 ; Eascafria, Madrid, 1 ;
Barracas, Castellou, 5; Dehesa de Valencia, 3.

Germany ; Schmilka, Saxony, 1 ; Rudolstadt, Thiiringen, 2 ; Marxheim,
Bavaria, 1 ; South Germany, 2.

Austria-Hungary : Spalato, Dalmatia, 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. Gallon, 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?.

Maredsous, Namur,

Rev. G. Fournier

3. 3. 30. 1-2.

Belgium.

(c & P).

2 9.

Boulogne, Pas-de-Galais,

C. G.Danford(c&p).

89. 10. 6. 2.

Franco.

10. 7. 17. 1.

4 9.

L'Hospitalet, Ariege.

G. S. Miller (c).

8. 8. 4. 172-175

2 6, 9.

L'Hospitalet, Ariege.
(A. Robert.)

0. Thomas (p).

8. 9. 1. 72-74.

9.

Port6, Pyrendes-Orien-
tales. {A. Robert.)

0. Thomas (p).

8. 9. 1. 75.

9.

Port6, Pyr6n6es-Orien-

tales.

G. S. Miller (c).

8. 8. 4. 171.

2 c$, 4 9.

Nimes, Gard. (C. Mot-
taz.)
Valescure, Var.

0. Thomas (p).

8. 8. 10. 57-62.

5 (5, 2 9.

G. S. Miller (c).

8. 8. 4. 176-182

9.

Valescure, Var.

Hon. N. C. Roths-
child (c & P).

10. G. 13. 1.

1.

Prance.

Dr. J. E. Gray (p).

43. 12. 29. 17.

6.

France.

Tomes Collection.

7. 1. 1. 76.

9al.

Cabanas, Coruna, Spain.

Prof. V. L. Seoane
(c & P).

94. 1. 1. 9.

1.

Cabanas, Coruna.

Prof. V. L. Seoane

(c & P).

94. 3. 12. 2.

2 6.

Jaca, Huesca. {N. Gon-
zalez.)

0. Thomas (p).

8. 2. 9. 75-76.

i.

Panticosa, Huesca.

(A''. Gonzalez.)

0. Thomas (p).

8. 2. 9. 77.

i.

L6rida. (N. Gonzalez.)

0. Thomas (p).

8. 2. 9. 80.

2 9.

Pajares, Leon. (JV. (fo7i-
zalez.)

0. Thomas (p).

8. 2. 9. 78-79.

2<},3 9.

Pajares, Leon. (N.Gon-
zalez.)

0. Thomas (p).

8. 2. 9. 81-85.

558

8 9.

Silos, Burgos.

?.

Castrillo, Burgos.

2 9.

Villalba, Madrid.

(N. Gonmh'z.)

6, 3 al.

La Granja, Segovia.

3 6.

Barracas, Castellon.

{N. Gonzalez.)

3S.

Dehesa de Valencia.

{N. Gonzalez.)

6, 9.

Eudolstadt, Tbtiringen,

Germany. {W. Schiilto

6.

IMarxheim, Bavaria.

{Wolterstoif.)

2.

S. Germany.

1.

Spalato, Dalmatia.

2 9.

Slittelberg, Austria.

{Dr. F. Major.)

9.

St. Cergues, Vaud, Swit-

zerland.

1.

Miirren, Bern.

1.

Switzerland.

G. S. Miller (c). 8. 8. 4. 57-64.

G. S. Miller (c). 8. 8. 4. 65.

O. Thomas (p). 8. 2. 9. 70-71.

M. de la Escalera (c). 6. 11. 4. 5-S.

O. Thomas (p). 8. 2. 9. 72-74.

0. Thomas (p). 8. 2. 9. 67-09.

Lord Lilford (p). 0. 5. 5. 1-2.

Lord Lilford (p). 0. 5. 5. 3.

Dr. A. Giinther (c). 59. 9. 6. 25-26.

Lord Lilford (p). 95. 4. 6. 1.

O. Thomas (p). 8. 11. 30. 5-6.

C. Mottaz (p). 0. 2. 6. 5.

W. Giirtner (p). 92. 10. 5. 0.

E. R. Alston (p). 79. 9. 12. 3.

ELIOMYS GYMNESICUS Thomas.

1901. Eiiomys quercinus Thomas, Proc. Zool. Soc, London, p. 41.

1903. Eiiomys gyninesicus Thomas, Ann. and Mag. Nat. Hist., 7th ser., xi,

p. 494, May, 1903. Type in British Museum.
1910. Eiiomys querdnus gymncsicus Trouessart, Faune Manim. d'Europe

p. 137.

Type locality. — San Ci'istobal, Minorca, Balearic Islands.

Geographical clisirihution. — Balearic Islands.

Diagnosis. — Similar to Eiiomys quercinus but with smaller
ear (height from meatus 20 to 22 mm. instead of 23 to 25 mm.).

Colour. — The colour is in all re.spects like that of Eiiomys
quercinus, though tlie 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.

Specimens examined. — Six, all from the Island of Minorca.

Bemarhs. — The Balearic Eiiomys appears to be well differen-
tiated from the mainland animal by its smaller ear.

5 S, S San Cristobal, Minorca- O. Thomas and E. I. 0. 7. 1. 44-49.
skull. Pocock (c & p).

(0. 7. 1. 47. Ty2ie of species.)

559

ELIOMYS PALLIDUS Barrett-Hamilton.

1899. Eliomys imllidus Barrett-Hamilton, Ann. and Mag. Nat. Hist.,
7th ser., iii, p. 226, March, 1899. Type in British Museum.

1901. Eliomys cincficauda Miller, Proc. Biol. Soc, Washington, xiv, p. 39,
April 25, 1901. (Sorrento Italy). Type in U.S. National Museum.

1010. Eliomys cincficauda and E. pallidus Trouessart, Faune Mamm.
d'Europe, pp. 137, 188.

Tiipf locality. — Palermo, Sicily.

Gi'ograplncal dhtrlhution. — Sicily and southern Italy.

Diagnosis. — Size greater than in Eliomys querclnns (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.

Measureinemis. — 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). Four
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). For 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, 3; Ficuzza, 7; San Giuglielmo, Castelbuono, 3;

Mondello, 2 ; Madonna del Alto, 4.

Remarks. — In general the South Italian Eliomys so closely
agrees with E. quertinus 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 comjaared with E. qucrcinus, 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 jsoint. 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 (J, G ?. Aspromonte, Calabria, O. Thomas (p). 6. 8. 4. 2-5.

Italy. (A. Robert.) 8. 9. 1. 33-35.

6, 9. Palermo, Sicily. J. I. S. Whitaker (p). 98. 10. 6. 6-7.

> (98. 10. 6. 6. Tijpe of species.)

? 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. Rohert.) 8. 9. 1. 32.

5G0 EODENTIA

2(5. Mondello, 1000m. Sicilv. 0. Thomas (p). 6. S. 4. 33-34.

(A. Robert.)
'2 6, '2 9. Madonna del Alto, Sicily. 0. Thomas (p). 8. 9. 1. 28-31.

(A. Bobcrt.)

ELIOMYS SARDUS Barrett-Hamilton.

1901. EUomys sardics Barrett-Hamilton, Ann. and Mag. Nat. Hist.,

7th ser., vii, p. 3-40, April, 1901.
1910. Eliomijs sardus Trouessart, Paune Mamm. d'Europe, p. 138.

Type locality. — Tricoli, Cagliostra, Sardinia.

Geographical clistribiition. — Sardinia and Corsica.

Diagnosis. — Similar to Elio)ai/s pall idiis 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
Eliomijs pallidus.

Skull and teeth. — The skull does not attain the large size of
that of E. pallidus. In form it is peculiar in the relatively
greater depth of brain-case, the ratio of depth through bulla to
condylobasal length ranging in four specimens from 41 -|- 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 sjaecimens : 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.
Corsica : Asco, 5.

i juv., ?. Tricoli, Sardinia. O. Thomas (p). 1. 3. 8. 1-2.

(G. Meloni.) (1. 8. 8. 1. Type of species.)

6, 9. Lanusei, Sardinia. 0. Thomas (p). 1. 3. 8. 3-4.

[O. Meloni.)

2 (5, 2 9. Asco, Corsica. {Dr. O. Thomas (p). 8. 11. 30. 1-4.

F. Major.)

ELIOMYS LUSITANICUS Reuvens.

1890. El{io'mys] nitela var. lusitanica Eeuvens, Die Myosidae oder

Schlaefer, p. 28, footnote (Lisbon, Portugal).
1897. Mioxus nitela var. amori Graels, Mem. Real Acad. Sci., Madrid,

XVII, p. 481 (Cordova, Spain).
1901. Elliomysl amori Thomas, Proc. Zool. Sec, London, p. 41.
1904. EUomys amori Cabrera, Bol. Real Soc. Espan. Hist. Nat., xvii,

p. 187, Api-il, 1904.
1908. EUomys lusitanicus Cabrera, Ann. and Mag. Nat. Hist., 8th ser., i,

p. 189, February, 1908.
1910. EUomys quercinus lusitanicus Trouessart, Paune Mamm. d'Europe,

p. 137.

Type locality. — Lisbon, Portugal.

Geographical distribution. — Southern portion of the Iberian
Peninsula ; exact limits of range not known.

Diagnosis. — Largest European member of the genus EUomys
(hind foot in adults usually more than 30 mm.) ; skull con-

ELIOMVS 561

spieuously larger and heavier than in E. qnerriniin, the corulylo-
basal length of skull, ^G-^D mm. ; colour sliglitly darker and
more brighty russet than in E. quercinus, the tail usually with
black sub-terminal area forming a complete ring.

Colour. — Paratype (No. 4G. 11. 21. 1): hair.s of back with
three distinct colour bands: (1) slate-grey (8 mm.) ; (2) light
bufly grey (2 mm.) ; (3) russet (2 mm.), the extreme tips of hairs
whitish. Longer hairs black except for the slate-grey bases.
The general effect is a peculiar dull reddish brown, faintly
" lined " by the longer black hairs and a little "powdered "by
the whitish tips to the ordinary hairs. On sides the bufly 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. qKcrrinun. Tail as in E. qncrcinus,
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 difierence
that might readily be accounted for by their fi'esher condition,
the paratype having been in the Museum collection moi'e than
fifty years longer than the others. In five 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 Donana, 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.

SlcuJl and teeth. — Except for its noticeably larger size ,the
skull resembles that of Eliomys quercinas. 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. pallidus and E. saidii,s. Teeth as in E. quercinus, but appre-
ciably larger, the difference more noticeable to the eye than
might be supposed from the actual measurements.

Meafturements. — Paratype (sex not known) from Lisbon,
Portugal : hind foot, 30. Adult male from Goto Donana, Huelva,
Spain : head and body, 166 ; tail, 120 ; hind foot, 31 ; ear from
meatus, 27. Three adult males from 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.

lieninrJcs. — 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). 4G. 11. 21. 1.

^ (Paratype of species.)

2 6, ?. Seville, Spain. {Dr. A. Lord Lilford (p). 95. 3. 3. 19-21.

Euiz.)

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RODENTIA

6 st Seville. {Dr. A. Buiz) Lord Lilford (p). 95. 3. 3. 22.

6, ?. Seville, Spain. Lord Lilford (p). 11. 1. 1. 108-109.

1. Goto Douaiia, Huelva. A. Chapman (c & p). 0. 5. 10. 1.

Jerez, Cadiz. A. Chapman (c & p). S. 3. 26. 1-2.

■> ,'

Genus DYROMYS Thomas.

1780. Myoxiis Zimmermann, Geogr. Gesch., ii, p. 351 (part).

1857. Eliomys Blasius, Saugethiere Deutschlands, p. 288 (Sab-genus of

Myoxus) part.
1890. Myoxus Reuvens, Die Myoxidae oder Schlaefer, p. 24 (Sub-genus of

Myoxus) part.

1906. Dvyomys Thomas, Proc. Zool. Soc, London, 1905. ii, p. 345,

April 5, 1906 (Sub-genus of Eliomys). Not of Philippi, An. Mus.
Nac. de Chile, xiv, p. 20, 1900.

1907. Dyromys Thomas, Ann. and Mag. Nat. Hist., 7th scr., xx, p. 40G,

November, 1907 (Substitute for Dryomys).

Type. — Mug niteduJa Pallas.

GeoijrapJiiral di.sfribvtion. — From central Asia through Asia
Minor to Hungary and eastern Switzerland.

Characfcrti. — Teeth resembling those of Elknitiis but relatively
smaller than in any of the other European members of the
family, the ci'owns 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 m^, 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 Eliomyi: 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 differing
from Eliomys in the uniformly haired, distichous tail.

HemarJcs. — 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 laces of one species)
occur in Eurojie.

KEY TO THE EUROPEAN FORMS OF DYIWMYS.

Skull broad and robust, the zygomatic
breadth about 17 mm. ; auditory bullae
decidedly enlarged (Rustschuk, Bul-
garia) D. robustiis, p. 572.

Skull slender, the zygomatic breadth about
15 mm. ; auditory bullse not specially
enlarged D. nitedula, p. 567.

567

Teeth woak ; crown area of upper pre-
molar decidedly more than hali that

of first molar (Greece) V. nitcdula wiiujei, 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) I). nUi'diUa nUedida,^. 568.

General colour of adults greyish brown

above (Tirol and eastern Alps) D. niiedula inlcrmedim, p. 569.

DYROMVS NITEDULA Pallas.

(Synonymy under subspecies.)

Geographical distribution. — South-ea.stern Euroiae from eastern
Switzerland and north-eastern Italy into Asia Minor ; north to
northern Hungary. 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 bullae not
sjDecially enlarged.

External characters. — Externally as in EUoniys 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 margin 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 between
tubercles less papillose, scarcely more than reticulate, and
postero-internal tubercle not much longer than that at base of
thumb, though narrower. Mammaj : 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 whitish hairs at
edges. Underparts and feet whitish or very pale bufF, this
colour usually sufi'using 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.

SJcuU. — In general ajipearance the skull resembles that of
Eliomys quercinus, 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

zygomata. The form of the auditory bulla is also, as in Elioniys,
much more inflated anteriorly tlian posteriorly, and readily dis-
tinguishable from the rounder, more evenly inflated bulla of Glis
and Muacardimix.

Teetli. — The detailed structure of the teeth has already been
sufliciently described. As compared with those of Eliomi/s
qu('rrinii.'< the molar rows are more nearly parallel, and there is

FlC. 111.
Diirom.'jn iiltedula. Nat. size.

i-'IG. 112.
Dtiromys niteihtla. Cheek-teeth, x 10.

usually less contrast in size both above and Ijelow between the
premolar and last molar with the two middle teeth.

Rcmarliii. — Among the European Mnscardinidse this species is
immediately recognizable by the combination of a dark face
streak and uniformly bushy tail. In size it is intermediate
lietween Muscardinus avellanarius and Eliomys quercinus, but
nearer the former. In difl'erent parts of its range Dyrowyx
nifednJn has become modified into four local races, three of
which occur in Europe.

Dykomys nitedula nitedula Pallas.

1778. Mils nitedula Pallas, Nov. Sp. Quadr. Glir. Ord., p. 88, part, des-
cription, uot synonymy (Region of the lower Volga, Russia).

1782. Myoxus dnjas Schreber, Saugthiere, pi. ccxxv b. ; description, iv,
p. 831, 1787 (Region of the lower Volga, Russia).

1857. Mijoxiis dnjas Blasius, Siiugethiere Deutschlands, x^- 29.5.

1906. Eliom:/-^ nitcdultis Thomas, Proc. Zool. Soc, London, 1905, ii, p. 348,
April 5, 1906.

1910. Dyromys nitedula Trouessart, Faune Mamm. d'Europe, p. 133.

Tijpe localifi/. — Region of the lower Volga, Russia.

Geographical distriltHtlon. — South-eastern Europe north of
the Balkan Peninsula, west to Hungary. Exact limits of distri-
bution not known.

DYROMYS 569

DiiitjiKisis. - Skull with I'ostrum rather long, the distance from
anterior root of zygoma to tip of nasal more than 8 mm. ; teeth
robust ; crown ai-ea of upper premolar scarcely half that of first
molar ; general colour of adults yellowish brown above.

Colour. — Adult : upper parts a j'ellowish lirown intermediate
between the russet of llidgsvay and yellowish wood-brown, the
back inconspicuously "lined" with black, the sides lightening
almost to ochraceous-buff ; underparts pale cream-bulT; 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 gi'ey, this colour continuing to ear as a boi'der to black
stripe about 2 mm. wide. An obscurely delined 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 Zuberec,
northern Hungary, 20 '5, 21 and 21 mm. For cranial measure-
ments see Table, p. 571.

Spcci77iens examined. — Twelve, from the following localities : —
Austria-Hungary : Zuberef-, northern Hungary, 6 ; Herkulesbad, 3 ;
Hungary, no exact locality, 1.
Servia : Belgrade, 2.

4, 1 juv. Zubereo, Hungary. Budapest Museum (e). 94. 3. 1. 37-41.

1. Zuberer. G. Barrett-Hamilton 11. 1. 2. 59.

(P)-
2 6,19. Herkulesbad. {F. Cox.) Hon. N. C. Eothschild 7. 9. 16. 12-14.

2 9. Belgrade, Servia. Dr. B. B. Sharpe (p). 75. 8. 24. 3-4.

Dyromys nitedula intebmedius Nehring.

1902. Myoxus intermedins Nehring, Sitz.-Ber. Gesellsch. Naturforsch.

Freunde, Berlin, p. 155.
1910. Dyromijs nitedula inicrmedius Trouessart, Faune Mamm. d'Europe,

p. 134.

Type localit)/. — Near Lienz, Tirol, Austria-Hungary.

Geographical distrihiition. — Tirol and eastern portion of the
Alps (Engadine and Cadore).

Diagnosis. — Similar to Dyronnjs nitidula nitedula, but general
colour of upper parts greyish brown without tinge of yellow.

Colonr. — 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 creamy white in strong contrast ; underparts
and feet buffy white, this colour suffusing whole underside of
tail. Face markings as in true nitedula, but grey edging along
upper side of black lines less e-wdent.

Measurements. — Two adult males from Vulpera-Tarasp,

570

RODENTIA

Gri.sons, Switzerland : head and body, 02 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, 86;
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.).

Bemarls. — At first sight this animal might be supposed to be
the young of Dyromys nitedtda nitedula, but none of the three
specimens examined shows any indication of immaturity.

1 al. Cadore, Venetian Alps, Dr. E. Festa (r). 9. 1. IS. 1.
Italy.

Dyromys nitedula wingei Nehring.

1S81. Elioinys dnjas Winge, Vidensk. Meddel. fra den naturh. Foreu. i
Kjobenhavn, 1881, p. 50 (near Athens). Not Mus dryas Pallas.

1902. Myoxus icincjei Nehring, Sitz.-Ber. CTCseUsch. Naturforsch. Freunde,
Berlin, p. 5 (Parnassus region, Greece).

1910. Dyromys nitedula ivingei Trouessart, Faune Mamm. d'Europe, p. 134.

Type locality. — Parnassus region, Greece.

Geograpliical distrihution. — Known only from the t\'pe locality
and from the vicinity of Athens.

Diaijiioslii. — 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 S mm., and teeth weak ; upper pre-
molar with crown area decidedly more than half that of fii"st molar.

Colour. — The colour appears to be indistinguishable from that
of D. nitedula nitedula.

Skull. — The only skulls examined are broken. While the
bram-case does not appear to differ from that of true D. nitedtda,
the rostrum is shorter and weaker than in the allied species,
though not different in form. Auditory bulke apparently larger
than in D. n. nitedida, but the material is not sufficient to show
whether this character is constant.

Teetlt. — 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 witli less broadening at middle.

Measurements. — 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 meatu.s, — and 12 "4.

Specimens examined. — Two, both from Tatoi, north of Athens, Greece.

Remarlis. — Though at present very imperfectly known the
Grecian Dyromys appears to be a well -characterized local form.

2. Tatoi, Greece. (C. Mottaz.) Hon. N. C. Roths- 8. 10. 2. 24-25.

child (p).

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572 RODENTIA

DYROMYS ROBUSTUS Miller.

1910. Dijyomijs rohusiiis Miller, Ann. and Mag. Nat. Hist., Stb ser., vi,
p. 459, November, 1910. Type in British Museum.

TjjjM- locaJifij. — Rustschuk, Bulgaria.

Gev(jrapltical dlxiribntion. — Known from the type locality only.

Diagnosis. — Like D;/romys nitedula but skull broader and
more robust, with relatively shorter ro.struin and much enlarged
auditory bulla;.

External characters and colour. — Not appreciably diflerent
from those of D. nitedula (no skins examined).

Shdl and teeth. — Skull of essentially the same length as in
the largest specimens of Dyromi/s nitedula, Imt conspicuously
broader and more robust ; brain-case relatively shorter and
broaderj its width above zygomatic roots nearly equal to distance
from posterior border of interparietal 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 bulhe, the brain-case appears deeper than in the related
species ; auditory bulke of the same form as in D. nitedula, 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. nitedula ; relative size of upper
premolar about as in D. n. iinncjei.

Measurements. — Type (adult female in alcohol) : head and
body, 95 ; tail, S7 ; hind foot, 22 ; ear from meatus, 15. For
cranial measurements see Table, p. 571.

Specimen crcrniincd .—The type.

9 al. Rustschuk, Bulgaria. K. Andersen (c). 11. 10. 8. 1.

{Type of species.)

Genus GLIS ISrisson.

1762. Glis Brissou, Regn. Anim. in Classis ix distrib., 2nd cd., p. 13(6'/)*).
1780. Myoxus Zimmermanu, Geogr. Gesch., ii, p. 351 (Type by tautonymy

Sciurus glis Linnseus).
1843. Myoxus Wagner, Abhandl. kais Bayer. Ak. Wissensch., Miincben,

Math.-Phys. Classe in, p. 185 (Sub-genus of Myoxus).
1857. Glis Blasius, Saugethiere Dcutschlands, p. 288 (Sub-genus of

Myoxus).
1890. Myoxus Reuvens, Die Myoxidae oder Schlaefcr, p. 56 (Sub-genus of

Mjjoxus).
1895. Glis Merriam, Science, N.S., i, p. 376, April 5, 1895 (Genus).
lEOO. EUks Schulze, Zeitschr. fiir Natnrwiss., Stuttgart, Lxxiii, p. 200

(Sub-genus of Myoxus) part, included glis and nitedula.

Type species. — Glis Brisson = Sciurus glis Linna?us.
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.

Character>i. — Skull flattened, slightly angvdar, the interorbital
region with lateral ridges which unite to form a median crest in
old age ; brain-case rather broad and low, the auditory bulhe
moderate ; ectopterygoid reduced to a low ridge ; jugal very long
in front, extending almost or quite to laclu-ymal ; mandible with
angular portion entire ; dental formula as in Eliomyx ; crowns of
cheek-teeth very slightly concave, the inner and outer margins
scarcely elevated, the outer side of jh^ and m' with five low
cusps ; crowns of m^ and wi- alike in form, each crossed by four
complete transverse ridges, in the spaces between which lie three
incomplete ridges ; external appearance sijuirrel-like, the tail
conspicuously distichous.

RemarJcs. — 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 3Iitscar(Unus.

GLIS GLIS Lin mens.
(Synonymy under subspecies.)

GeograpMcal fh'sfribution. — From the northern portion of the
Iberian Peninsula to Asia Minor ; north to the Baltic coast of
(Germany ; south to Sardinia and Sicily ; west to the Atlantic.

Diagnofiis. — General characters as in the genus ; size largest
of the European 3Iiiscardiriidge (head and body about IGO to
190; hind foot about 27 to 34; condylobasal length of skull
about 3G to 44) ; colour greyish above, whitish below, the tail
usually darker or more slaty than body ; no blackish markings
anywhere.

External charactersi. — General form and appearance squirrel-
like. Details of structure much as in Eliomys quercinus except
in the following particulars. Ear much smaller, extending barely
to eye M'hen laid forward ; scarcely a trace of ridge on inner
surface of conch, but meatus Mdth a small though evident anti-
tragus-like lobe. Eeet more robust than in the other European
members of the family, but relative lengths of digits as in
Eliomi/x ; 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 jiosterior third,
its tubercles not ditiering noticeably among themselves in form,
the postero-internal tubercle not elongated, but the three posterior

574

KODENTIA

larger than the three anterior. Claw of inner toe relatively
more I'educed, closely appressed to pad. Tail easily broken as in
Eliomys 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 : p'2 — '2, a 2 — 2, i2 — 2 = 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 j underfur slate-grey at base, the general
body colour appearing at extreme tips only ; hairs unusually
glossy throughout, producing a noticeable metallic silvery lustre,
especially on posterior half (»f back ; head faintly lighter than
body ; a narrow dusky eye-ring ; ears like surrounding parts or
somewhat dusky ; under parts 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, buff'y white below, tlie light area on
under side of tail usually better defined than in adult.

Fig. 113.
Giix giis. Nat. size.

Sli-ull. — In general the skull differs from that of Eliomys
querclnuH chiefly in its broader, lower form, a peculiarity that is
e.specially noticeable in posterior view. Profiles essentially as in

GLIS 575

E. querclnus except that nasals ai-e abruptly bent downward at
tip and ventral 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 inflated than in Eliomys, particularly
in its posterior portion, which is not visible when skull is viewed
from above, and is not closely appressed 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 mesojjterygoid space essentially as in
Eliomys, but hamular not in contact with bulla, and ectopterygoid
reduced to a low ridge. Mandible much more I'obust than that
of Eliomys, especially in its anterior portion ; coronoid process
more 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
squarish, 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 posterior of which is least developed ;
between these three incomplete ridges extend inwai'd to or a
little beyond middle, the second reaching outer side. These ridges
are best developed in m^ and wt^, 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 ni- 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 cro\^n, while in ni^ the structure
of all the ridges becomes indefinite near periphery of crown.
Lower molars essentially li^e the maxillary teeth, but with

576

EODENTIA

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 only

Fig. 114.
aiiis yll!<. Cheek-teeth. X 10.

two incomplete ridges (the second and third) ; nin with crown
longer and all the ridges better developed than in )»''.

IteinarJi's. — Glis glifi is immediately recognizable among
European memVjers 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. ; con-
dylobasal length of skull usually 3.5 to 39 mm.

(Central Europe, south to northern Italy) G. ijUn gliji, 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. <j. ilalicus, 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 gus glis Liunseus.

17GG. [Sciurits] glis Linnaeus, Syst. Nat., i, 12th cd., p. 87 (Central

Europe).
1779. lOlis] csculentns Blumeubach, Haudb. der Naturgescb., p. 7'.) (Central

Europe).
181G. Glis vulgaris Okeu, Lehrbuch der Naturgescb., in, pt. 2, p. 868

(Germany).
1832. ? M\_iioxus] giglis F. Cuvier, N. Ann. Mus. d'H. N. Paris, i, p. 444.

Nomeu nudum, " le loir proprement dit.''
1840-45. Myoxiis avcllanus Owen, Odontography, ii, p. 25, pi. 105.
1857. Myoxus glis Biasius, Saugetbiere Deutschlands, p. 292.
1890. Myoxus glis Reuvens, Die Myoxidae oder Schlaefer, p. 61.
1895. Gtis (sic) glis Merriam, Science, N.S., i, p. 376, April 5, 1895.
1910. Glis glis Trouessart, Faune ;Mamm. d'Europe, p. 181.

Ty/2^e locality. — Germany.

Geo(jrapliical 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 enlarged ; general colour of upper parts a yellowish broccoli-
brown scarcely 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, Huuyad,
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-HuNGAHY : Steiermark, 1 ; Hollos, 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.) ; Toggeu-
burg, St. Gallen, 1 (U.S.N.M.). ; Mels, St. Gallen, 2 (B.M. and U.S.N.M.) ;
Wolfbalden, 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, Ti^inn, 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, Turin, 10 (Turin); Monca-
lieri, Turin, 7 (Turin).

2 (5, 2 ?. Nimes, Gard, France. O. Tlaomas (p). 8. 8. 10. 53-56.

(C. Mottaz.)

2, 2 juv. France. Purchased 46. 1. 2. 1-4.

(Lefebvre).

1. France. A. Baillon (c). 46. a.

2 9. Marxheim, Bavaria. Lord Lilford (p). 11. 1. 1. 102-103.

{Schuchardt.)

?. Ekernkrug, Germany. Lord Lilford (p). 11. 1. 1. 104.

1. Trieste, Austria-Hungary. Lord Lilford (p). 11. 1. 1. 152.

(J. O. Haggard.)

i. Horkulesbad, Austria- Hon. N. C. Rotlis- 7. 9. 16. 11.
Hungary. [F.J. Cox.) child (p).

4 (5, 2 ?. Hatszeg, Hunyad, Tran- C. G. Danford (c). 3. 11. 8. 21-26.
sylvania.

1. Miirren, Bern, Switzer- W. Gurtner (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.)

? juv. Thayngen, Schaffhausen. 0. Thomas (p). 2. 8. 4. 35.

{E. H. Zollikofcr.)

9. Mels, St. Gallen. {E. H. 0. Thomas (p). 2. 8. 4. 34.

Zolliliofer.)

(J St. Switzerland. E. R. Alston (p). 79.9.12.1.

Glis glis italicus Barrett-Hamilton.

1898. Glis italicus Barrett-Hamilton, Ann. and IMag. 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 ? GUs insularis Trouessart, Paune Mamm.
d'Europe, p. 132.

Type locality.- — Siena, Italy.

Geographical distribution. — Italy, from region of Genoa,
southward ; Sicily ?

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, 15'^ ; hind foot, 3^ ; 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;

GLis 579

hiud foot, 34; ear from meatus, 21. Young female from near
Palermo, Sicily (type of insularis) : 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,
Pinalborgo, Genoa, 6 (Genoa) ; near Vittoria, western Liguria, 2 (Genoa) ;
Caluzzano, 1 (Genoa) ; Malasana, 2 Genoa) ; near Moutariolo, 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.

jRcmarlcs. — Throughout southern and central Italy, north
to about the region of Genoa, the large dormouse appears to be
very comstant in its characters, dift'ering 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 piobably 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 for 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 9. Genoa. Marquis G. Doria (p). 89. 12. 11. 1-3.

?. Verona. [Conie delle O. Thomas (p). 99. 11. 10. 4.

Oddi.)
9. Florence. A. Savage Laudor 97. 3. 7. 2.

(c & p).
<5, 5 9. Siena. (Brogi.) Dr. E. Hamilton (p). 98. 10. 2. 11-lG.

(98. 10. 2. 14. Ti/2)e of subspecies.)
2. Siena. {Brogi.) G. Barrett-Hamilton 11. 1. 2. 60-61.

(!-)•
9. Monte Sirino, Lagonegro. O. 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. Tyjye of G. insularis Barrett-Hamilton.)

Glis glis melonii Thomas.

1907. Glis melonii Thomas, Ann. and Mag. Nat. Hist., 7th ser., six,

p. 445, May, 1907. Type in British Museum.
1910. Glis melonii Trouessart, Faune Mamm. d'Europe, p. 132.

Type locality. — Marcurighe, Urzulei, Ogliastra, Sardinia.

Geogrnpliical clistrihution. — Sardinia and probably Corsica also.

Diagnosis. — Similar to Glis glis italicus, l)ut not attaining so
large a size (skull rarely more than 41 mm. in condylobasal
length), and dark terminal area on tail not so extensive, seldom
involving more than distal tlfird.

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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. — Twenty-four, all from Sardinia (B.M. and
Genoa).

Roiinrlcs. — 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 c5, 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. Qlia qlis pyrenaicus Cabrera, Ann. and IMag. Nat. Hist., 8th ser., i,
p. 19:3, February, 1900.

Type localitij. — Alio, Navarra, Spain.

Geograplu'cal 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-detined zone of pure yellowish buff extending from
the cheek to the hip. Tail glosssy brownish grey, with the usual
whitish line along under side. Ears and orbital rings brown.
Hands and feet white ; a broad l)rown metatarsal patch.

»SVc«7/.- — Like that of tyjjical (t. i/lis, but larger, approaching
that of Glis (jlis italicus.

Measurements. — Type (in flesh): head and body, 169; tail-
vertebrfe, 137; hind foot, 28; ear, 16. Skull: greate.st 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. Miiscardinus Kaup, Entw.-Gescb. u. Natiirl. Syst. Europ. Thierwelt,

I, p. 134.
1843. Muscardinus Wagner, Abhandl. k. Baj'er. Akad. Wissensoh., Miiu-

chen, Math.-Phys. 01., in, p. 185 (Sub-genus of Mijoxus).

1857. Muscardinus Blasius, Saugefcbiere Deutschlands, p. 289 (Sub-genus

of My ox us).
1890. Muscardinus Beuvens, Die Myosidae oder Scblaefer, p. 25 (Sub-genus
of Myoxiis).

Type species. — 3Ius avellanarius Linnseu-s.

Geographical disirihution. — 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 Eliomi/s and Di/romys, 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 meml)ers 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 nt} and wr
conspicuously unlike in size and form, the live 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 gi'oup.

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 OF MUSCARDINUS.

Upper parts dull yellowish brown ; no sbarp line
of demarcation along sides ; no white stripe
between ear and ej'e (Distribution general)... M. avellanarius, p. 583.

Upper parts brigbt bufi ; a sharp line of demarca-
tion aloug sides ; a vertical wbite stripe
between ear and eye (Central Italy) M. pulcher, p. 590.

MUSCARDINUS AVELLANARIUS LinnaJUS.

1758. [il/Hs] avellanarius Linnaeus, Syst. Nat., i, 10th ed., p. 62 (Sweden).
1782. Myoxus muscardijius Schreber, Siiugtbiere, pi. ccxxvii ; text, iv,
p. 835, 1788 (Germany).

1858. Myoxus avellanarius Blasius, Saugethiere Deutschlauds, p. 297.

584 KODENTIA

1869. Mils corilinum Fatio, Faune Vert. Suisse, i, p. 183 (name attributed
to Schreber).

1890. Muscardinus avellanariiis Reuvens, Die Myosidae oder Schlaefer,
p. 69.

1900. M[uscardiniis] avellanariiis anglicus Barrett-Hamilton, Proc. Zool.
Soc, London, p. 86 (Bedford Purlieus, Thornhaugb, Northampton-
shire, England). Type in British Museum.

1910. Muscardinus avellanariiis and J/, avellanariiis anglicus Trouessart,
Faune ]Mamm. d'Europe, p. 135.

Tyjie locality. — Central Sweden.

Geographical distribufion. — 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 stripe in front of ear.

External characters. — General form essentially as in Eliomijs
quercinus, but eye relatively larger 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 Di/romys, 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 corre.s2)onding
tubercle, its claw obsolete ; other digits relatively longer than
in the r'elated genera. Tail partly prehensile, slightly
flattened, extending about to ears when laid forward, uniformly
haired and loosely short-busliv throughout. Mamma? :
pl_l, al_l 42-2 = 8.

Colour. — Entire upper j^^rts and outer surface of legs
yellowish brown, the exact shade between the wood-brown or
clay-colour and buft' of Ridgway, sometimes with a slight greyish
or ecru-drab cast across shoulders, the middle of back incon-
spicuously s])rinkled with blackish hairs ; sides, flanks, cheeks and
ears tinged with ochraceous-bufl" ; tail concolor with back above,
faintly lighter below, the tip with a dusky tinge ; underparts
and inner surface of legs light ochraceous-bufl', slightly contrasted
\\ith 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 jiure white (to base of hairs), this occasionally

MUSCAEDINUS

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-riug ; feet ochraceous-buff, the toes sprinkled with silvery
hair.

SJiuIl. — In general form the skull differs from that of the
other European Muscardinidee 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 Glis,
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
angulai'-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 ;
infraorbital foramen relatively smaller
and more nearly circular in outline than
in any of the other Eui'opean members
of the family. Nasals and nasal branches
of premaxillaries 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 1H3 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 avellanariug.

Nat. size.

586

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 emargination 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 ; %\hen viewed from the
side the outer terminations of the
five ridges have the appearance of
five low cusps, decreasing regularly
in height from befoi'e 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, scjuarely
transverse, equally spaced ridges, all* extending from border to
border, the outer extremities appearing as five or six low cusps.
Third u])per molar like second, l>ut 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 on" (except in m^),
their terminations producing the usual effect of low cusps along
outer border ; crown area diminishing I'egularly from m^ to ?%
(which is only a little more than half that of anterior
tooth) ; Hi., and m^ squarish in outline, slightly longer than wide ;
Wj decidedly longer than wide, narrower anteriorly than
posteriorly.

* Except occasionally the third.

Musranlinus arellaiiarins.
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 (Juines, 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 Padoia, 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 ; Chalfout, Buckingham, 1 ; Bury St. Edmunds, Suffolk, 2 ;
Colcliester, Essex, 3; London (purchased alive in), 2; Norwood, Headley,
Surrey, 1; Eatham, 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-j\Ioselle, 4 ;
Cranves-Sales, Haute-Savoie, 2; Lucinges, Haute-Savoie, 2; Montauban,
Haute-Savoie, 1 ; Scientriers, Haute-Savoie, 1 (Jlottaz) ; 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; Zuberec,
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. GaUen, 2; Ziiberwangeu,
St. Galleu, 1 (U.S.N.M.); Vulpera-Tarasp, Grisons, 5 (Rothschild);
Ticino, 1 (U.S.N.M.); Lugano, Ticino, 1 (U.S.N.M.).

Italy-: Padoia, 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.

BcmarJcs. — Skins from different parts of the range of 3Jms-
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

5. Halton, Buckingham- Hon. N. C. Roths- 11. 1. 3. 490.

shire, England. child (p).

9. 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.)
i, ? al. Burv St. Edmunds, Suf- Dr. Giinther (c & p). 90. 5. 27. 1.
folk.

* Type of anglicus Barrett-Hamilton.

t The type of angliczis is an adult in full, rich winter pelage.

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EODENTIA

,ij C V c^

,^c

Purchased in London.

Ratham, Sussex.

(TF. Jeffrey:,
Sussex.

Honiton, Devonshire.

G.Barrett-Hamilton 11. 1. 2. 113.

E. K Alston (p). 79. 9. 12. 2.

E. Cavendish Taylor 11.1.3.403.

G. C. Shortridge 11. 1. 3. 402.
(c & P).
England. Yarrell Collection. 56. 12. 10. 682.

<J, 9. Guines, Pas - de - Calais, O. Thomas (c & p). 94.6.6.12-13.
France.
3 9, juv. Manonville, Meurthe-et- Lord Lilford (p). 11.1.1.105-107.

Moselle. {Lcynwnt.)
2 6. Cranves - Sales, Haute- 0. Thomas (p). 5.11.18.9-10.

Savoie, 1200 m.

(.4. Bobert.)
S, 9. Lucinges, Haute-Savoie. O. Thomas (p). 6. 4. 2. 4-5.

{A. Robert.)
6. Moutauban, Haute-Savoie, 0. Thomas (p). 6. 4. 7. 6.

900 m. (.4. Bobert.)
1 St. France. {F. Petit.) E. Hargett (p). 97. 11. 25. 4.

6. Oberwald, Silesia, 400 m. Lord Lilford (p). 99. 1. 9. 16.

Germany. (17. Baer.)
9 al. Silesia. Dr. Kaup (p).

3. Csallokoz-Soniorja, Hun- Budai^est Mus. (e). 94. 3. 1. 43-45.

gary.
1. Zuberec, Hungary. Budapest Mus. (e.) . 94. 3. 1. 42.

9 al. Brasso, Hungary. Prof. L. von IMehely 95.8.12.3.

l,4]uv. Hatszeg, Hunyad, Trau- C. G. Danford (c). 3. 11. 8. 27-31.

sylvania.
9, 9 juv. St. Gallen, Switzerland. 0. Thomas (p). 2. 8. 4. 36-37.

{E. H. ZoUikofer.)
9. Siena, Italy. (Brogi.) G.Barrett-Hamilton 11. 1. 2. 112.

MUSCARDINUS PULCHER Barrett-Hamilton.

1855. '? Mijoxus speciosxis Dehne, Allgem. deutsche Naturbist. Zeitung,

neue Folge, i, p. ISO, Tursi, Basilicata, Italy.
1898. Muscardinus •puldier Barrett-Hamilton, Ann. and Mag. Nat. Hist.,

7th ser., ii, p. 423, November, 1898, Perugia, Italy.
1900. M{uscardinus] avellanarius speciosiis Barrett-Hamilton, Proc. Zool.

See, London, p. 87.

1910. Muscardinus avellanarius speciosus Trouessart, Fauna Mamm.
d'Europe, p. 136.

Tn2]e locality. — Perugia, pi-ovince of Perugia, Italy.

Geograpliical distribution. — At present known from Perugia,
Rome, and the vicinity of Naples, Itah'.

Diagnosis. — Like Museardinus avellanarius but general colour
brighter, the flanks nearly raw-sienna; white of throat extending
upward as a conspicuous whiti.sh stripe between eye and ear.
Teeth somewhat larger than in 31. avellanarius.

External characters. — In external form I can detect no difler-
ence between this species and 3Iu.sca7-dinus avellanarius : mamma? 8,
arranged as in the more northern animal.

MUSCARDINUS 591

Colour. — Upper pai'ts 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 M. avcUanarius, this area when
present not forming noticeable contrast with rest of underparts.

Shull and teeth. — The skull and teeth do not differ appre-
ciably from those of Museardimis avellanar'mti, though the teeth
are usually a trifle larger.

Measurements. — Type (adult male) : head and body, 90 ;
tail, 68 ; hind foot, 10 ; 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. — Fiftj'-seven, from the following localities in Italy :
Perugia, 1 (type) ; Monte Cimiuo, Rome, 1 (Genoa) ; Sorrento, 54
(U.S.N.M.) ; Palermo, Sicily, 1 (too yonng to be positively identified).

Remarks. — This species is well differentiated from Muscar-
dinus aveUanarius by its bright, 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. Whitaljer (p). 8. 9. 28. 1.

Family MURID^.
1821. Miiridx Gray, London IMed. Repos., xv, p. 303, April 1, 1823.

Geographical distribution. — As in the order Rodentia.

Characters. — Anterior portion of zygomatic arch not formed
chiefly by jugal ; infraorbital foramen large, wider above than
below, without accessory canal ; jugal bone splint-like, sujjported
by long zygomatic processes of maxillary and squamosal, not
in contact with lachrymal anteriorly ; mandible with angular

592 EODENTIA

portion arising from under side of alveolus of incisor, its lower
border not angulated ; tibia and fibula joined ; cheek-teeth never
more than |^, rooted or rootless, the crowns tuberculate or
prismatic.

BrmarJi's. — 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 MUBID^E.

Molars prismatic, hypsodonfc or rootless, their crowns

flat (Voles and Lemmings) Micivtinx, p. 610.

Molars tuberculate, brachyodont, rooted.

Tubercles of maxillary molars arranged in two primary

longitudinal series Cricet'mx, p. 592.

Tubercles of maxillary molars arranged in three

primary series ^Iurin;i',^.l'dl.

Sub-Family CRICETIN^.
1866. Cricetinx Murray, Geogr. Distrib. Mamm., p. 358.

Geof/rcqMcal distribution. — Entire American Continent ; cen-
tral region of Asia from eastern China west into south-eastern
and central Europe ; southern Africa.

Characters. — As in the sub-family Murinse, but tubercles of
maxillary teeth arranged in two primary longitudinal rows.

BemarJcs. — 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 OP CRICETIN^.

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

foot ; maramEe 8 (Central Euro^^e, west into

Belgium and France) Cricetus, 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 ; mammas 16 (Eastern Europe, west into

Roumania and Bulgaria) Mesocricctus, p. 605.

CRICETULDS 593

Genus CRICETULUS Milne-Edwards.

1867. Cricetulus Milne-Edwards, Ann. des Sci. Nat., Paris, 5th ser., Zool.,
vir, p. 375.

Ti/pc species. — Cricetulus (jriscus Milne-Edwards.

Geographical distrihution. — Central region of Asia from
eastern China west through Asia Minor to the eastern portion
of the Balkan Peninsula.

CJiaractirs. — Like Criceius, but less highly modified : form
murine ; feet with normally developed tubercles ; tail in
European si^ecie? 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 Apodemus ; 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
Criceius.

Itemarlcs. — The genus Cricetulus contains about a dozen
described species, whose distribution extends across the entire
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. Gricetus arenarius Winge, Vidensk. Middel. fra den naturh. Foren.

i Kjebenhavn, 4th ser., iii (1881), p. 31.
1902. Cricetulus attlcus Nehring, Sitz.-Ber. Gesellsch. Naturforsch.

Freuude, Berlin, p. 3, January, 1902.
1910. Cricetulus atticus Trouessart, Faune Mamm. d'Europe, p. 165.

Tijpe locality. — Pentelikon, Attica, Greece.

Geographical distribution. — Greece ; limits of range nob
known, bub 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. instead
of about 27 mm., hind foot about 13 '6 mm. instead of 15 mm.
Skull with brain-case somewhat narrower than in C. phieus. 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 ibericns 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 Q

594

RODENTIA

t:^

blackish. The individual haii's are slate-grey at base, those of
the uuderfur 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 grey is replaced by light buff. 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 and over base of whiskers to sides of muzzle, the lips thus
entirely white and sharply defined against the narrow grey
muzzle. A very 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 bufly white, including hairy portion of sole. Tail light
cream-bufi', the upper surface sprinkled with blackish hairs along
median region.

Shill. — Apart from its much smaller size (approximately as
in Apodemiis flaviroUis) the skull
differs from that of Cricefus cricctus
in its generally less modified, more
murine appearance. Dorsal profile
sligiitly and evenly convex through-
out. Brain-case relatively narrower
and rostrum relatively broader than
in Apodemus flavicollis and A. sylva-
ticus, but these peculiarities not
carried to the same extreme as in
Criceius 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 Apodeimis sylva-
ticus but nari-ower and deeper, and
with condj'les more projecting. Interparietal about as in
A. sylvaficus, 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 lai'ge, 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.

CRICETULUS

595

2 Q

596 RODENTIA

with posterior border of m^, the hamulars not diverging pos-
teriorly. Incisive foramina moderately long and narrow, much
as in (Jriccfiis 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 Ajjodemus. 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. For cranial measurements see Table, p. 595.

Specimens exaviiiied.— Three, all from the vicinity of Athens.

Bemarlcs. — Though nearly related to the Cricetulus phseus of
Asia Minor, the Grecian animal appears to be sufficiently distin-
guished by its smaller size.

2 ?. Athens, Greece. (C. Mottaz.) Hon. N. C. Koths- 8. 10. 2. 47-48.

child (p).

Genus CRICETUS Leske.

1779. Cricetus Leske, Anfaugsgruiide der Naturgesch., i, p. 168.

1799. Hamster Lac^pede, Tabl. des divisions ordres et genres des Mamm.,

p. 10 {Hamster nigricans Lac^pede = Mas cricetus Linnaeus).
1869. Cricetus Blasius, Saugethiere Deutschlands, p. 305.
1873. Heliomys Gray, Ann. and INIag. Nat. Hist., 4th ser., xii, p. 417,

November, 1873 (H.jeudii = Cricetus cricetus).

Type species. — Mus cricetus Linna?us (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 ; mamma?,
c^ ; 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 moi'e than half as broad as
brain-case ; ectopterygoid fossa short, deep, moderately pit-like ;
infraorbital foramen with well developed extei'nal plate ; enamel
pattern characterized by presence of noticeable supplemental
median loops or islands in maxillary teeth.

CRICETUS

597

Remarks. — The genus Cricetus 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 Linnajus.

(Synonymy under subspecies.) i

Geor/rapMeal flisfrihution. — Central Europe from Belgium and
noi'thern 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.

E.vternal characters. — Form I'obust 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 ; sui'face 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 gi'oove 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.
Mamma3 •.p2 — 2,i2 — 2 = 8.

Colour. — Upper parts a uniform light yellowish brown or
brownish buff", the exact shade usually something between the
wood-brown and ochraceous-bufi" 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 ai'ea 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.

SJcull. — 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 lachrymal 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 general
outline when viewed from above (when of latter form with
ends truncate at lambdal and interorbital regions) ; most
of the surface of brain-case is formed by the broad temporal
areas of muscle attachment, between which lies the narrow flat
median region continued 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 jn-ocess, 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 ;

I'ICi. 118.
Cricetus cricetus. Nat. size.

behind angles the ridges and furrow pass gradually into the
narrowest anterior portion of Hat 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

"00 _ 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 branches 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, posteriorly 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 l>roadly rounded anterior extremity scarcely extending
to level of alveolus of m^, the plate-like, ill-defined hamulars in
contact with auditory IjulliB ; ectopterygoids well developed,
lying, except at their anterior extremity, at a higher (more
dorsal) 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. no.

Criceliis cricehts. Teeth X 5. (i, unworn ; h, sliyUtly wcjiii.

shaft sliglitly and somewhat obli(|uely compressed, the antero-
posterior diaineter 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 towai'd 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
donl)le transvei-se 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 wj\ 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 in- 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, antei'o-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
disappeai-ed, 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 nij relatively smaller and m^ 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 m.,,
its posterior pair of cusps very little reduced ; anterior border of
each tooth with low but evident ridge on outer side.

lieinarks. — Cn'ceius cricetus diflfers conspicuously from all
other European rodents in its iinusual colour pattern with
underparts entirely black. Four local forms have been described.

602

EODENTIA

Three of these occur west of Russia. Their status is at present
by no means clearly understood.

KEY TO THE EUROPEAN EACES OP 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 uim.

Occipital portion of brain-case high, its truncation

moderately oblique as in typical race (Belgium

and western Germany) C. c. caneacens, p. 603.

Occipital portion of brain-case low, its truncation

unusually oblique (Roumania) C. c. nehringl, p. 605.

Cricetus cricetus cricetus Linmvus.

1758. [Mils'] cricetus Linnaeus, Syst. Nat., i, 10th ed., p. 60, Germany.
1792. lf[Ms] Cricetus germanicus Kerr, Anim. King., p. 243 (Cricetus

gervianicus in Syst. Catal., inserted between pp. 32 and 38)

Germany.
1799. Hamster nigricans Lac^pede, Tabl. des division, ordres et genres des

Mamm., p. 10 (Renaming of Mus cricetus).
1801. Mlus] clricetus] fidvus Bechstein, Gemeinn. Naturgesch. Deutsch-

lauds, I, 2nd ed., p. 1010 (Thiiringen, Germany).
1803. Cricetus vulgaris Geoffrey, Catal. Mammif. du Mus. Nat. d'Hist.

Nat., p. 196, northern and eastern Europe (Renaming of

Mus cricetus).
1811. Cricetus frumentarius Pallas, Zoogr. Rosso. -Asiat., p. 161 (Renaming

of Mus cricetus).
1857. Cricetus frumentarius Blasius, Siiugethiere Deutschlands, 'p. 306.
1867. [Cricetus vulgaris] varius Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lvi, pt. i, p. 98

(Europe).
1867. [Cricetus vulgarus] albus Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lvi, pt. i, p. 98

(Germany).
1867. [Cricetus vulgaris] niger Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lvi, pt. I, p. 98

(Austria, Hungary, Germany).
1873. Heliomys jcudii Gray, Ann. and Mag. Nat. Hist., 4th ser., xii,p. 417,

November, 1873 (locality imknown).
1894. Cricetus cricetus Dahl, Die Heimat, iv, p. 130, June, 1894.
1906. Cr[icctus] vulgaris niger Simroth, Biol. Centralblatt, xxvr, p. 337,

June 1, 1906. Valley of the Saale, Germany.
1910. Cricetus cricetus and C. cricetus niger Trouessart, Fauue Mamm.

d'Europe, p. 159.

Tjipe locality. — Germany.

Geographical distrihution. — 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.

CRICETDS 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 examiiud. — Nineteen, from the foUowiag localities : —

History unknown, 1 skull (type of Heliomijs jeudii Gray).

Gebmany: No exact locality, 1; Ingelbeim, Kheinhesseu, 6; Ober-

lausitz. Saxony, 1 ; Lotzen, Saxony, 1 (U.S.N.M.) ; near ]\Iagdeburg,

Saxony, 6; Saxony, no exact locality, 1 (U.S.N.M.); Strassburg, 1

(Lataste).

Austeia-Hdngaey : Molna Szecsod, Hungary, 1 (U.S.N.M.), an

immature individual, perbaps not true cricetus.

1. Germany. (Muschler.) E. li. Alston (p). 79. 9. 25. 35.

6 S. Ingelbeim, Ebeinbessen. C. Hilgert (c). 8. 11. 2. 38-43.

S. Oberlausitz, Saxony. Dr. E. Hamilton (p). 97. 12. 4. 33.

3, 1 juv. Magdeburg. Dr. W. WolterstorS 92.12.1.24-27.

skull. Lidtb de Jeude Col- 67. 4. 12. 314.

lection.
(Type of Heliomys jeudii Gray.)

Cricetus cricetus canescens Nehring.

1899. Cricetus vulgaris var. canescens Nehring, Sitz.-Ber., Gesellsch.

Naturforscb. Ereunde, Berlin, p. 1.
1910. Cricetus cricetus canescens Trouessart, Fauue Mamm. d'Europe,

p. 160.

T^pe localifi/.—Neav Fexhe-Slins, banks of the Maas,
Belgium.

Geographical (listribution.— 'Eastern Belgium and north-
western Germany ; probably north-eastern France also.*

Diagnosis. — Similar to C. 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 c.ramined. — Four, from tbe following localities : —
Belgium : Tirlemont, Liege, 3.
Germany : Brunswick, 1 (Eield).

Remarks. — The status of this race is not satisfactory. The
Belgian specimens which I have seen show none of the

* I have seen no Frencb specimens, but the form will probably prove
to be canescens. For account of recent cbanges in distribution in France
see Trouessart, Faune Mamm. d'Europe, p. 159.

604

RODENTIA

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605

peculiarities in colour described by Nehrinj;^, but agree perfectly
ill this respect 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
crici'tun.

2 (5, 1 al. Tirleinont, Belgium. Rev. L. Wamau (p). 8. 11. 23. 1-3.

Cricetus cricetus nehringi Matschie.

1901. Cricetus nehrincji Matschie, Sitz.-Ber., Gesellsch. Naturforscli.

Freunde, Berlin, p. 232.
1910. Cricetus cricetus nehringi Trouessart, Faune IMamm. d'Eurox3e,

p. 160.

Tjipe locality. — Described from specimens from Slobosia,
Cernavoda, and Barza, Roumania. The first may be assumed to
be the type locality.

Gcograpliical 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 posteriorly 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, Roumania (U.S.N.M.).

Genus MESOCRICETUS Nehrmg.

1898. Mcsocricetus Nehring, Zool. Anzeiger, xxi, p. 49-1, September 5,
1898 (Sub-genus of Cricetus).

1898. Semicricetus Nehring, Zool. Anzeiger, xxi, p. 494, footnote (Alterna-
tive for Mcsocricetus).

1898. Mediocricetus Nehring, Zool. Anzeiger, xxi, p. 494, footnote (Alter-
native for Mesocricetus).

1900. Mcsocricetus 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 RODENTIA

hind foot, nearly concealed in fur, and mammse 14 to 16 * instead
of 8 ; skull resembling that of Cncetiis in general features, but
anteorbital foramen without external plate, and flattened area on
dorsal surface of ))rain-caso broad, its outline somewhat truncate-
diamond shaped ; ectoi^terygoid fossa short, deep, and con-
spicuously pit-like ; teeth not essentially diflerent from those of
Gricetus, 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 the Old World group of Crlcetinse.

MESOCRICETUS NEWTONI Nehring.

1898. Cricctus ncwtoni Nehring, Zool. Anzeiger, xxi, -g. 329, May 16, 1898.
1898. Gricetus (Mcsocricetus) ncwtoni Nehring, Wiegmann's Archiv fiir
Naturgeschichte, 1898, i, p. 386, December, 1898.

1900. M[c$ocricctus] ncxvtoni Satunin, Zool. Anzeiger, xxiii, p. 301, May 28,

1900.
1910. Mcsocricetus nciutoni Trouessart, Faune Mamm. d'Burope, p. 162.

Type locality. — Schumla, eastern Bulgaria.

Geographical distribution. — Eastern portions of Bulgaria and
Roumania. Exact limits of range not known.

Diagnosis. — Size smaller than in Gricetus 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 backward 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 Gricetus 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 Gricetus, extending to eye when laid forward. Feet as in
Gricetus, 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.
Mamma;: 7-7 or 8-8 (in skin No. 122097, U.S.N.M.,
apparently j^ 4 — 4, i 4 — 4).

Golour. — 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 from 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. Freundo, Berlin,

1901, p. 155.

MESOCRICETCS

607

region in Cricefas cricetus ; chin dull whitish ; upper lio 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
markings 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

Fia. 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 lambdal
suture, the two borders together producing a diamond-shaped
figure. Interparietal with antero-posterior diameter relatively
greater and tranverse diameter relatively 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
infraorl)ital 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 vei-y 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
l)oth 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.

Specimens examined. — Nine, from the following localities : —
Roumania: Malcoci, Dobrudscha, 7 (B.M. and U.S.N.M.).
BuLGAEiA : Melschka, near Rustchuk, 2 (Andersen).

Remarks. — Aj^art from the generic characters, its peculiar

MKSOCRICKTUS

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EODENTIA

colour pattern will at once distinguish this animal from all other
European rodents.

1. Malooci, Dobrudsclia, Purchased (Pruliere) (p). 86. 4. 2. 5.
Roumania.

1. Rustcliuk, Bulgaria. Dr. Knud Anderseu (p). 6. 5. 17. 1.

Sub-Family MICROTIN.E.
1906. Microtinee Miller, North Amer. Fauna, No. 12, p. 8.

Geographical distribution. — Northern jaortion of both hemi-
spheres, south to jNIexico, northern India, and the Mediterranean
coast of Europe

Characters. — Murine rodents with definitely prismatic,
hvpsodont or rootless, flat-crowned molars.

Bemarlcs. — The sub-family Microtinee contains about thirty
gi'oups, 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 3Iicroti, containing the true voles, and the
Ellohii, the last highly modified for undergi'ound 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 members of the subfamily are found throughout
this region, some of them representing genera now extinct.

KEY TO THE EUROPEAN GENERA OF MICROTIN.^.
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 Mjjopus, 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 Evoiomijs, 'p. 623.

Bony palate terminating posteriorly in a sloping
median ridge and two lateral pits.
Sole with six well developed tubercles ; mammae, 8. Microtns, p. 658.
Third upper molar with three re-entrant angles

on inner side Mici-otu),; p. 659.

Third upper molar with two re-entrant angles on

inner side Chionomijs, p. 712.

Sole with five well developed tubercles ; mammte,
8 or 4.
First lower molar with first outer and first
inner triangle communicating ; mammfe, 4 ;
general form modified for underground life... Pitijmys, p. 752.
First lower molar with first outer and first inner
triangle not communicating ; mammfe, 8 ;
general form modified for aquatic (rarely
subterranean) life Arvicola,"^. 723.

Gil

Genus MYOPUS Miller.

1910. Mi/ojmn Miller, Smithsonian lliscell. Coll., lii, p. 497, January 12,
"1910.

Tv/jje species. — Mi/odes schist icolor Lilljeborg.

Geo(jraphical disfrihiition. — Southern Norway and central
Sweden ; eastward into Finland and perhaps much further.
Limits of range not known.

Characters. — Skull and teeth as in Levimus ; general form
vole like, 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
manus normal, much shorter than first
and second phalanges combined, the
claws not enlarged ; ear well developed
though small, with distinct meatal valve.

Hchiat-ks. — The genus Mi/ojms is ,,, ,.

characterized by the combination of the c , ^ ,, , ^ ,
'' -11 bkeleton oi fore foot of

skull and teeth OI Lcmnms with the Lemmus (a) and Myojms (b).

general body-form and foot structure of

the voles. It is, therefore, less specialized than the other Old

World lemnnngs, representing a stage of development equivalent

to that of the American Synaptomys. Only one species is

known.

MYOPUS SCHISTICOLOR Lilljeborg.

1844. Myodes schisticolor Lilljeborg, Ofversigt af Kongl. Vetenskaps-Akad.
Forhandl., Stockholm, i, p. 33, March 20, 1844.

1910. Leimnus schisticolor Trouessart, Faune Mamm. d'Europe, p. 199.

1911. Ml/opus schisticolor Oollett, Norges Pattedyr, Hefte 3, n. 130

March, 1911.

Type locality. — Near Lillehammer at north end of Mj6.sen,
(iudbrandsdal, Norway.

Geographical distribution. — Fir forests of southern Norway
and central Sweden ; eastward into Finland and perhaps much
further.* Exact limits of range not known.

Diagnosis. — General characters as in the genus ; size
essentially as in Microtns agresfis (head and bod\' 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, Siingothiere, 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

KODENTIA

External characters. — Form vole-like, slightly more robust
than in Microtus agrestis, the legs not shortened and feet not
enlarged as in Lemmus lenimus. 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, tijl 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 chgit
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 lai-ger 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 (jlareoJus, its
five tubercles relatively larger and more crowded, occupying
umch more than half the surface of the region in which they
occur, its digits with no special jjeculiarities 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 hindermost are
somewhat smaller and more terete 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 vertebne. Mammae : j^ 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 1 5 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.

Sliull and teeth. — Apart from their noticeably smaller size the
skull and teeth do not dift'er essentially from those of Lemmus
lemmus. The general outline of the skull when viewed from

613

1

Ridges joined, high.
,, low.
,, moderate.

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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 j)r, instead of to level
of first prism of m^, anteorbital foramen with slit-like lower
portion so shortened as to be practically absent.

In the few specimens examined the inner border of cutting
edge of upper incisor is less folded backward than in Lemmus
lem.mnft, and the outer re-entrant angle of vi.^ 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.

Spi'ciniciis examined. — Fifteen, from the following localities: —
Norway : Hurdalen, Gudbrandsdal, 1 ; Fluberg, Hadeland, 2 (B.M. and

U.S.N. M.) ; Eidsvold, Akershus, 2 ; Heen, Christiania, 1.

Sweden: Ncrike, Wermland, 2 (U.S.N.M.) ; Upsala, 1 (U.S.N.M.) ;

Medolpad, \V. Norrland, 4 ; no exact locality, 1 (U.S.N.]M.) ; Dalarne, 1.

1. Hurdalen, Gudbrandsdal, Dr. R. Collett (p). 84.10.81.3.

Norway.
6 juv. Fluberg, Hadeland. Christiania Museum 93. 3. 1. 18.

(E).

2 9 al. Eid:svold, Akersbus. Dr. R. Collett (p). 84. 10. 31. 1-2.

6 juv. Heen, Christiania. Christiania Museum 93. 8. 1. 19.

(E).

2 i juv. Medelpad, W. Norrland, Lord Lilford (p). 11. 1. 1. 111-114.

2 9 juv. Sweden.
1 al. Dalarne. • Stockholm Museum 90. 8. 1. 18.

(E).

Genus LEMMUS Link.

1795. Lenmms Link. Zool. Beytriige, i, pt. ii, p. 75. Type by tautonymy

Mas Icvimns Linnteus.
1811. Myodcs Pallas, Zoogr. Rosso-Asiat., i, p. 172 (part).
1877. Myodes Coues, Monogr. N. Amer. Rodeutia, p. 237.
1896. Lcminm Miller, N. Amer. Fauna, No. 12, p. 36, July 23, 1896.

Type species. — Mns Jehimus Linnasus.

Geographical distribution. — Arctic region of both hemispheres ;
in Europe south to southern Norway.

Characters. — General form 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 phalange^ ; ungual phalanges of manus greatly enlai'ged,
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 broad and low, the zygoma
cons^jicuously 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 ; m^ with four transverse
loops and no closed triangles ; m^ with three transverse loops and
one closed triangle.

Memarhs. — 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. \_Mus] lemmus Linnseus, Syst. Nat., i, lOth ed., p. 59 (Sweden).

1808. L[einmus'] lemmus Tiedemann, Zoologie, i, p. 474.

1820. Lemmus borealis Nilsson, Skand. Faun., i, p. 185 (Substitute for

lemmus).
1822. Lemmus norivcgicus Desmarest, Mammalogie, pt. ii, p. 287 (Norway).
1895. Myodes lemmus Collett, Christiania Vetenskabs, Selskabs Forhand-

liuger, 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 bo 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 bullaj very large,
their substance conspicuously spongy ; a large foramen below and behind
alveolus of m.^ ; tooth-rows noticeably converging anteriorly ; upper molars
with outer re-entrant angles conspicuously deeper than inner ; m-, m^, and
m^ with one or more transverse loops in addition to the terminal loop.

Dicrostouy.r.— 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 bullcB not unusually large, their substance scarcely
spongy ; no noticeable foramen below and behind alveolus of m^ ; tooth-
rows nearly parallel ; upper molars with outer and inner re-entrant angles
approximately equal ; m", m^ and m.^ without transverse loops in addition
to the terminal loop. Externally Dicwstonij.v is distinguished from
Lemmus by the reduction of tlie 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 rod?:ntia

1896. L[emmus'] Icnimus Miller, North American Fauna, No. 12, p. 37,

July 23, 1896.
1910. Lcmvius Icvimus Trouessart, Faune Mamm. d'Europe, p. 198.

Type locality. — Mountains of Lappmark, Sweden.

Geograpliicaldistrihution. — 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.| During seasons of abnormal increase the animals wander
to the extreme south of Norway.

Diagnomfi. — 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 widtli 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 jiointed 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, nearly as long as vertebrae at tip where they

* CoUett, Norges Pattedyr, p. 138, 1911.

t Lilljeborg, Sveriges ooh Norges Eyggradsdjur, i, p. 32G, 1874.

I

LEMMUS G 1 7

form a dense pencil ; anniilations irregular but rather well
defined, about O'O 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. Mammse :
p-2-2, i 2-2 = 8.

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 apj^earing 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 strij^e 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 butfy 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 j^ale
buflf; some trace of a buffy 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 usualty sprinkled
with blackish hairs. There is not much seasonal change in
colour. Winter specimens tend to be more j-ellowish, 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.

SJiull. — 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 alDOut 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 bullse ;

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-({uadrate, 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
l^rojecting more backward and outward than downward, their
lowest point slightly below level of condyles ; tloor of brain-case
with no sjiecial features, the basioccipital with a moderately
defined median longitudinal ridge, its width along basal suture

Via. 122.
Lemmu,!, Icmmiis. Nat. size.

contained about 2^ times in median length ; auditory bulla large
but not very liighly 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 anterioi'ly, 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

I

LEMMUS 619

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 i/t', 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 foi- 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 compiessed ; 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.

Teeth. — As compared with those of the typical voles, such as
the members of the genera Microtus and Arvicula, 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 flattened border
directed inward, the anterior margin pale j,,jq ^.,3

yellow in colour, slightly wider than posterior Lemmus lemmus.
margin, the greatest transverse diameter a Teeth, x 5.

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 internalh',
following course of enamel, its posterior border forming a distinct

620 RODEXTIA

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 hke 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 outer re-entrant angle, a second loop essentially
like first loop of m^, and two closed triangles like the posterior
pair in m^ but slightly smaller ; re-entrant angles three, all well
develojied and regularly alternating, two outer and one inner.
Third upper molar witla 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 m- 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 I'e-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 angles 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 extremity of loop
as a closed triangle not quite equal in size to the outer closed
triangle of nt' and m" ; terminal loop similar in form to that of
the other mandibular teeth, but slightly larger and less narrowed.

I

LEMMUS 621

Meusurciiients. — 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, j). 622.

Specimens examined. — Seven ty-sevcu, from the following localities : —
Norway : Troudhjem, 4 ; Roraas, Trondhjem, 5 (B.M. and U.S.N.M.) ;
Brekkebygdon, Troudhjem, 6 ; Hyllingeu, Trondbjom, 3 ; Vigel Fjeld,
Trondhjem, 4; Kvikne, Trondhjem, 2; Dovre Mts., 1 (U.S.N.M.); Hjerkin,
Dovre, 5; Fille Fjeld 3; jMolmen, northern Gudbrandsdal, 2; Tonsfet,
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.IM.).

2. Roraas, Trondhjem, Nor- Ohristiania Museum 93. 3. 1. 16-17.

way. (e).

Z 6,b9. Trondhjem. G.Barrett-Hamilton 8. lU. 5. 1-S.

2(5. Kvikue, Trondhjem. G.Barrett-Hamilton 8.10.5.12-13.

<5, 2 9. Hjerkin, Dovre. G.Barrett-Hamilton 8. 10. 5. 9-11.
{J. L. Bonhotc.) (p).

(5, 9, Fille Fjeld. [Brown and E. R. Alston (p). 79. 9.25.63-65.

(J juv. Alston.)

2 6. Molmen, Gudbrandsdal. R. J. Cuninghame 98. 5. 2. 10-11.

(c & P).

2. Norway. F. Du Cane Godman 73. 3. 21. 2-3.

(P).
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. KoWioff.)

Note on Lemmus lemmus crassidens Nehring.

On March 3, 1896, Major G. E. H. Barrett-Hamilton exhibited
before tlie Zoological Society of London " several fresh-looking
skeletons of the Norway Lemming (My odes lemTiuis), obtained by
Dr. H. Gadow in caves near Athouguia, in Portugal." Commenting
on these specinrens 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 — Microfus. 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, My odes lemniHS. "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 tlie original skin, which had become so dried and
hardened tliat in order to enable myself to examine the skeletons I
had to get it removed. 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

Observations.

Ridges joined, low.

, , , , moderate.

,, moderate.
,, ,, low.

,, moderate,
nearly joined.
,, joined, low.

moderate.

,, low.

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Number.

85. 5. 2. 10
8. 10. 5. 2

1.54541

8. 10. 5. 9

154542
8. 10. 5. 10
8. 10. 19. 25

153134

8. 10. 19. 23

153138

153133

8. 10. 19. 24

15313G
8. 10. 19. 2G
8. 10. 19. 27

8GGr, 2 L*

',', iL

Is

5

' — ' — ■ - . . . 1

L. lemmus lemmus.

Norway : Tilulmen .

Brekkebygden
Tonsset, Hede

marken
Hjerkiu, Dovre

Sweden: Jemtland

L. lemmus "crassidens,

Portugal ?

EVOTOMYS 623

regret to say, not one of these most interesting specimens has been
preserved in the condition in which I received it. Some of the
vertebras 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
nature, and to come to tlie belief that they were of recent origin and
perhaps unimportant. . .The present skulls resemble those of recent
Lemmings very closely indeed ... I cannot find any characters
sufficiently important to enable me to separate the two specifically "
(I'roc. Zool. Soc. London, 1896, pp. 304-305). Three years later,
however, they were examined by Nehring and distinguished from the
Norway Lenuning as Mijodcs lemmtis 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, jd- 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 maximimi (see 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 Icmmus; in the type they equal those of the
largest-toothed Norwegian si^ecimen in the series (from jMolmen,
northern Gudbrandsdal) ; in one complete skull, one skull without
lower jaw, and one odd lower jaw they are slightly above the
maximum of lemvius ; 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 perfectlj' matched
among the seventeen Scandinavian specimens, while five are not.
This makes it impossible at present to synonymize crassidens with
lemmus. 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. Mijodcs do Selys-Longchamps, Etudes do Micromamm., p. 87

(section).

1840. Hijpiuhi'US Keyserliug and Blasius, Wirbelth. Europas, p. 34 (sub-

genus : type Mus glareohis Schreber), not Hypiuhcus Illiger, 1811.
1858. Hijpuda'us Blasius, Siiugethiere Deutschlands, p. 336 (sub-genus).
1874. Evolomys Coues, Proc. Acad. Nat. Sci. Philadelphia, p. 186 (genus,

type Mus rutilns Pallas).
1896. Evotomi's Miller, North Amer. Fauna, No. 12, p. 42, July 23, 1896.

624 RODENTIA

1900. Craseoniys Miller, Proc. Washington Acad. Sci., ii, p. 87 (sub-genus
of Evotomijs : type Hupudivus rufocanus Sundevall).

1907. Craseoniys Thomas, Proc. Zool. Soc, London, 1906, p. 863, April 11,
1907 (genus).

Tjipe species. — Mus rutilus Pallas.

Geographical distrihuiion. — 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.

BemarJcs. — In the shortness of the mandibular incisor root, a
peculiarity which it shares with the American Phenaronnjs, the
genus Evotomijs 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, j

KEY TO THE EUROPEAN FORJ^IS OP EVOTOMYS.

Tail densely haired, the pencil at least one-fourth

as long as vertebrffl (Arctic Europe) E. rutilus, p. 646.

Tail moderately haired, the pencil much loss than
one-fourth as long as vertebra?.
Skull of adult massive, its size large, the condylo-
basal length frequently more than 26 mm.
(25 to 27 '6 mm.); teeth heavy, length of
tooth-rows 6 ram. or more.
Red of dorsal area conspicuously contrasted
with grey of sides ; ear noticeably over-
topping fur ; vv' normally without third
re-entrant angle on inner side (Northern
and mountainous portions of Scandinavia) E. rufocanus, p. 648.

* True Evotomys, Craseoniys Miller, and Phaulomys Thomas (Ann.
and Mag. Nat. Hist., 7th ser., xv, p. 493, May, 1905. Type Evotomys
sviithii Thomas, from Kobe, Hondo, Japan).

t For observations on the lack of distinctions between Phaulomys and
Craseoniys see Anderson, Ann. and Mag. Nat. Hist., 8th ser., iv, p. 318,
October, 1909. For notes on the status of Craseoniys see account of
Evotomys rufocanus, below, p. 658.

EVOTOMYS

G25

Red of dorsal area not couspicuously contrasted
with yellowish brown of sides ; ear scarcely
overtopping fur ; m^ normally with third
re-entrant angle on inner side (Channel

Islands) E. avsarms, p. G45.

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. C44.

Nasal about equal to diastema, rarely much

longer E. glareolus, p. 6£G.

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. y. istericiis, 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. succiais, p. 636.

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. (J. iilareolus,-^. 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) K. y. hritannicus, p. 634.
Size larger, hind foot usually more than
18 mm. (17 to 20 mm.) ; condylobasal
length of skull usuaUy 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) E. g. norvegicus, p. 638.

Red dorsal area narrow, well defined from

dull grey sides (Pyrenees) E. g. vasconiee, p. 689.

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. helvetieus, p. 640.

•2 S

626

Size larger (hind foot 18 '6 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. uagen', p. 611.

Teeth heavy ; incisive foramina wide,
the width of the two together more
than J length (Southern Italy) E. <j. hallucalis, p. 04.3.

EVOTOMYS GLAREOLUS Schreber.

(Synonymy under subspecies.)

Geographical distrihxdion. — 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
somewliat more, moderately haired, the jjencil always much less
than one-fourth as long as vertebrfe ; 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 heavj^, the length of tooth-rows rarely
6 mm. (4 '8 to 6 mm.) ; colour variable but never so bright as in
Evotoxnjs rufilua, that of the larger local foi-ms 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 liairs 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 linely
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 j^ubescent 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 (about half as
large as postero-interual). Tail about half as long as head
and body or slightly more, its anuulatious ill defined and
completely concealed by the dense covering of hair ; pencil
well developed, usually about one-sixth as long as vertebne.
Mammaj : p2 — 2, i 2 — 2 = 8.

C'o/o«r.— 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 Evotomj/s rufocanus ; under-
parts light grey with or without a yellowish brown wash ; feet
greyish ; tail obscurely bicolor.

SJiull. — The skull of Evotonij/s glareolm is small and lightly
built, the surface smooth and rounded, without noticeable ridges
or depressions for muscular attachment. Dorsal profile in
general slightly convex throughout,
though flattened or even a little con-
cave in interorbital region, the nasals
sloping I'ather abruptly forward, their
surface nearly flat ; there is usually
no sharp distinction between dorsal
and posterior outlines, the profile
merely bending abruptly downward
jiosteriorly over somewhat obliquely
truncate occipital region ; ventral
protile 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 betw^een 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 arched, 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.
Evotomys glareolus.

Nat. size.

628

EODENTIA

from bulltie, 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 bullre 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 essentiallj^
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 bullie. 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-
detined. 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
anterior portion of inner surface ; cutting sui'face of tooth
distinctly hollowed but not suiBciently to produce any suggestion

EVOTOMYS 629

of the weakened appearance characteristic of Lenimus and
Myopus, its posterior border fading gradually into surface of
tooth or marked off by a slight angle. Lower incisor noticeably
more slender than upper tooth, its root extending beneath that
of wis, 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 Microius, the prisms extending uninterruptedly to
base. At a slightly later stage the extreme base is seen to be
smooth, foi'ming 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

Fig. 125.
Emtomijs glareolus. Enamel pattern in two individuals, x 5.

prong of each tooth somewhat larger than the posterior ; with
advancing age the prongs become i-elatively 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 Eurojaean 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 ?»'', which in turn is usually
somewhat longer than nr, the discrepancy in neither case very

630

RODENTIA

noticeable ; m^ nearly equal to m.-, and in.f combined, m.^ slightly
shorter than m^. 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 acute, 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 mai'ked in youth and earlier disappearing with age.
Third upper molar with antei'ior 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 ; posteroexternal
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-enti'ant 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 cluriiig tlie 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

lower molar 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 h?^ than in ??Z;„ in which the anterior re-entrant
angle is sometimes obsolete. The first and second loo2:)s of m.,
and the second loop of m^ 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.

Itemarlis. — 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
movmtains of southern Italy are noticeably larger. There seems
little doubt, however, that intergradation between the small
and lai'ge forms takes place in every instance, though as yet it
can be satisfactorily demonstrated in the case of cjlarcolus and
nageri only.

its presence or absence is also largely a racial character, as may be seen

from 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 britannicus.... 43 .. 11 .. 32 .. 25'5

E. glareolus ^uecicus 37 .. 12 .. 25 .. 32'4

E . glareolus istericHs 56 .. 36 .. 20 .. 64-3

E . glareolus norvegicus. 20 5 .. 15 .. 25 -0

E. glareolus vasconLr 29 .. 8 .. 21 .. 27-6

E. glareolus helveticus 100 .. 68 .. 32 .. 68-0

E. glareolus nageri 100 .. 85 .. 15 .. 85-0

E . glareolus hallucalis 1 .. 1 .. — .. —

E. skomerensis 15 .. 14 .. 1 .. 93-3

E. cxsarius 22 22 .. 0 .. 100

E.rutilus 16 .. 16 .. 0 .. 100

E. rufocanus 16 .. 1 15 6-2

632

RODENTIA

EvoTOMYS GLAEEOLUS GLAREOLUS Schreber.

1780. Mas qlarcolus Schreber, Saugthiere, iv, p. 680, pi. cxc b (LoUand,

Denraark).
1792. Mus rutilus minor Kerr, Anim. Kingd., p. 237 (Casan and Goettingen).

Based on Pallas, Glires, p. 247. Not Mus rattus tninor Kerr,

Anim. Kingd., p. 229.
1792. \_Mas rutilus] ;8 minor Donndorff, Zool. Beytrago, i, p. 452. Based

on Pallas, Glires, p. 247.
1803. Lcmnius arvalis Geofiroy, Catal. Mammif. du Mus. Nat. d'Hist. Nat.,

p. 185 (Meudon,. Seine, France). Not Mus arvalis Pallas.
1828. Arvicola fulvus jMillet, Faune de Maine-et-Loire, ii, p. 40 (Angers,

Maine-et-Loire, France). Not Lanmus fulvus Geofiroy, 1803.
1881. Hypudxus liercynicus Mehlis, Isis, p. 876 (Harz Mountains, Germany).
1834. Lcnimus rubidus Baillon, il^rn. Soc. Royale d'Eniulation d'Abbeville,

1833, p. 54 (Abbeville, Somme, France).
1834. H[jji?ud!i'us] glarcolus Melchior, Den Danske Stats og Norges

Pattedyr, p. 116.
1836. Arvicola rufcsccns de S61ys-Longchamps, Essai Monogr. sur les

Campagnols des Environs de Iji6ge, p. 13 (Longchamps-sur-Ger,

Belgium).
1842. Arvicola i^ratcnsis F. Guvier, Hist. Nat. des Mammif., vii, Tabl.

Gen. et M6th. Described and figured in Livr. 68 of same work,

1834 (Abbeville, Somme, France).
1857. Arvicola cjlareolus a. Blasius, Siiugethiere Deutschlands, p. 337 (part).
1896. E\votomys'] glarcolus Miller, North American Fauna, No. 12, p. 44,

July 23, 1896.
1900. Evotomys hcrci/nieiis hcrcynicus Miller, Proc. Washington Acad.

Sci., II, p. 100, July 26, 1900 (part).
1900. Evotomys hcrcynicus rubidus Miller, Proc. Washington Acad. Sci.,

II, p.'l02, July 26, 1900.

1909. Evotomys glarcolus Miller, Ann. and Mag. Nat. Hist., 8th ser., iii,

pp. 419-420, May, 1909.

1910. Evotomys glarcolus Trouessart, Faune ilamm. d'Europo, p. 170.

Tjipc lorality. — Island of Lolland, Denmark.

Geor/raphical distribution. — AVest-central Europe nortli of the
Alps and Pyrenees and south of the Baltic ; from the Atlantic
coast east to Silesia ; north-eastern limit of range not shown.

Diaijnosis. — 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., condyloba.sal
length of skull, 23 to 24-6.

Colour. — Winter pelage : red dorsal area diffuse and ill-
defined, extending from eyes nearly to ba.se of tail and tending to
.spread slightly over sides. In general colour it is very nearly
the mars-brown or prouts-brown of RidgWcxy, usually with a
tinge of russet, the exact shade resulting from various 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
abi-uptly into the strongly buff-tinged grey of belly. The buff
wash on ventral surface is variable, but usually conspicuous.

r

EVOTOMYS 63."5

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 browi* above, soiled white below.

Skull and teeth. — Skull small, nearly the minimum for the
species (see Table of measurements, p. 650) ; auditory bullie not
abruptly inflated on inner side, a character scarcely apj^reciable,
however, except on comparison with E. glareolus istericiis. Third
upper molar with third inner re-entrant angle present in about
one-half the specimens examined (see p. 631).

Mensurements. — 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 Miiuseklippe, 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, p. 650.

Specimens examined. — One hundred and eighty-sis, from the following
localities : —

Denmark: Hillerod, Zealand, 7; Nj'stad, Lolland, 7 (U.S.N.M.) ;
Odense, Fyu, 2.

Holland: Oosterbeek, Guelderland, 1; near Leiden, 2 (U.S.N.M.).

Belgium: Waremme, Li6ge, 28 (U.S.N.M.); Maredsous, Li6ge, 15; no
exact locality, 1.

France : Guines, Pas-de-Calais, 2 ; Pont de Briques, Pas-de-Calais, 5 ;
Dinan, C6tes-du-Nord, 4 ; Barbizon, Seine-et-Marne, 9 ; Mont-Dore, Puy-
de - Dome, 3 ; Liguieres - Bonneville, Charente, 16 (Mottaz) ; Cadillac,
Gironde, 1 (U.S.N.M.) ; Foret de Bouconne, Gers, 1 ; Lyons, Rhone, 1.

Germany: Brunswick, 44 (B.JM. and U.S.N.M.); Babrenberg, Harz
:Mts., 2 (U.S.N.M.); Mauseklippe, Bodetbal, Harz Mts., 6 (U.S.N.M.)
Ingelheim, Rbeinhessen, 6; Strassburg, 4; near Magdeburg, Saxony, 5
Tbarandt, Saxony, 1 ; Saxony, no exact locality, 1 ; Niesky, Silesia, 6
Gross Hennersdorf, Silesia, 2; Wolfshau, Silesia, 4 (U.S.N.M.).

Austria-Hungary : Haida, Arva, Bohemia, 1 (not typical).

2 S, 5 9. Hillerod, Zealand, Den- O. Thomas (c & p). 98. 6. 7. 13-19.
mark.

1 .St. Odense, Fyn. Prof. Sundeval (p). 49. 11. 1. 5.

?. Oosterbeek, Guelderland, O. Thomas (c & r). 98. 2. 1. 15.
Holland.

4:6,4:9. Maredsous, Liege, Bel- Hev. L. Warnau (p). 8. 10. 18. 1-8.
gium.

1. Belgium. E. de Selys Long- 45. 7. 58.

champs (p).

<J, 9. Guines, Pas - de - Calais, 0. Thomas (c & p). 94. G. 6. 16-17.

France.

6, 4 9. Boulogne, Pas-de-Calais. 0. Thomas (c & p). 98. 1. 9. 16-20.

634 RODENTIA

i 6. Dinan, Brittany. G. Barrett-Hamilton 8.9.29.4-5.

(c & p). 11. 1. 2. 43-44.

3 (5, 3 ?. Barbizon,Seine-et-Marne. G. S. Miller (c). 8. 8. 4. 242-247.

1. Lyons, Rhone. Purchased. 43. 10. 14. 9.
6. Foret de Bouconne, Gers. O. Thomas (p). 6. 4. 1. 86.

{A. Bobert.)

2 6,19. Brunswick, Germany. G.Barrett-Hamilton 8.9.29.6-7.

(c & p). 11. 1. 2. 45.

2(5,4 9. Ingelheim, Rheinhessen. C. Hilgert (c). 8.11.2.46-51.

2 ,5, 2 9. Strassburg, Alsace. O. Thomas (p). 8. 8. 10. 122-

(C. Mottaz.) 127.

3. IMagdeburg, Saxony. Dr. W. Wolterstorff 92. 12. 1. 18-20.

(c & P).

2. Hcrrenkrug, Magdeburg. Lord Lilford (p). 8. 10. 19. 4-5.
9 al. Tharandt, Saxony. Berlin Museum (e). 93. 1. 1. 21.

(Nitsclic.)

1. Saxony. Dr. Thienemann (p). 45. 11. 15. 18.

2 6, 9. Niesky, Silesia, Germany. Dr. E. Hamilton (p). 97. 12. 4. 24-2G.

{W. Baer.)

2 6, 9. Niesky, Silesia. {W.Bacr.) Lord Lilford (p). 99. 1. 9. 23, 24,

26.

6, 9. Gross Hennersdorf, Si- Lord Lilford (p). 99. 1. 9. 22, 25.
lesia. (W. Bacr.)

6. Haida, Bohemia. Lord Lilford (p). 8. 10. 19. 1.

EVOTOMYS GLAREOLUS BEITANNICUS Miller.

1832. Arvicola riparius Yarrell, Proc. Zool. Soc, London, p. 109 (England,

no exact locality). Not of Ord, 1825.
1900. Evotomys hcrcynicus britannicus Miller, Proc. Washington Acad.

Sci., II, p. 103, July 26, 1900 (Basingstoke, Hampshire, England).

Type in British Museum.
1910. Evotomys glarcolus biitannicns Troucssart, Faune Mamm. d'Europe,

p. 170.

Tfipe localiti/. — Basingstoke, Hampshire, England.

Geographical distrihtition. — 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.).

Measurt'hients. — 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 tyjje 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 Dumpliail, Elgin,
Scotland : head and body, 102 and 110 ; tail vertebrae, 55 and 51 ;
hind foot, 16 and 16 "8. For cranial measurements see Table,
p. 652.

Sjx'cimcns examined. — One hundred and eighty - three, from the
following localities : —

Scotland: Birnie, ilorayshire, 2; Dunphail, Elgin, 5; Fochabers,
Elgin (Edinburgh), 1 ; Lhanbride, Elgin, 1 ; Uortachy, 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 ; Kirtle
Bridge, Dumfriesshire, 1 ; Dumfriesshire, no exact locality, 1 ; Craigmere,
Galloway, 3 (Edinburgh).

England: Filey, Yorkshire, 1 ; Lanfair, Anglesej', 3 ; Llansligo, Angle-
sey, 2 ; Plas Maclog, Anglesey, 1 ; Parkgate West, Cheshire, 1 ; Higher
Sutton, Macclesfiold, 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;iLilford, Northamptonshire, 3; Oundlo, Northamptonshire, 1;
Histon, Cambridgeshire, 1 ; Worcestershire, 1; Wormsley, Oxfordshire, 4;
Luton, Bedfordshire, 2; Toddington, Bedfordshire, 1; Welwyn, Hertford-
shire,!'; 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,!; Bridport, Dorset, 9 ; Northlow,
Devonshire, 6.

6.

?al.

4 c5, 1 ?.

9.
2?.

2(5.
<5, 2?.

2 6.

2.

i.

6, 1.

6.
1.

L^, 9 al.

'2 i, 3 9.

4 9.

9.

6.

9.

1 al.

lal.

6.

B i r n i e, Morayshire

Scotland.
Birnie, Morayshire.

E. R. Alston (p).

Dunphail, Elgin.

Llanbride, Elgin.
Dunblane, Perthshire.

Dalmeuy. Edinburgh.
Cromlix, Stirlingshire

Stockbriggs,
shire.

Drumlaurig Woods,
Dumfriesshire.

Kirtle Bridge, Dum-
friesshire.

L a n f a i r, Anglesey,
Wales.

Plas Maclog, Anglesey.

Northenden, Cheshire,
England.

Aberia, Merionethshire,
Wales.

Aberia, Merionethshire.

Langharne, Carmarthen-
shire.

Bridgend, Glamorgan-
shire.

Swansea, Glamorgan-
shire.

Waltham, Lincolnshire,
England.

Herefordshire.

Bishopstone, Hereford-
shire.
Worcestershire.

Rev. G. Gordon

(c & p).
W. R. Ogilvie-Grant

(c & p).
W. Taylor (c & p).
W. R. Ogilvie-Grant

(c & p).
W. Evans (c & p).
W. R. Ogilvie-Grant

(c & p).
Lanark- E. R. Alston (c & p).

Jardine Collection.

Miss D. Bate (c & p).

C. Oldham (c & p).

T. A. Coward (c& p).
G. Barrett-Hamilton

G. H. Caton Haigh

(c & p).
G. Caton Haigh

(c & p).
W. E. de Winton

(c & p).
R. I. Pocock (c & p).

C. Oldham (c & r).

G. H. Caton Haigh

(c & p).
Rev. S. O. Ridley

(c & p).
Rev. S. 0. Ridley

(c & p).
J. S. Elliott (c & p).

79. 9. 25. 58.

88. 11. 22. 1.

11.1.3.209-210.
11.1.3.239-241.
11. 1. 3. 238.
11.1.3.211-212.

11.1.3.235-236.
11.1.3.243-245.

79. 9. 25, 56-57.

86. 7. 2. 9-10.

11. 1. 3. 242.

11.1.8.229-230.

11. 1. 3. 234.
11. 1. 3. 237.

94. 6. 5. 1-2.

11. 1. 3. 231.

253-256.
11.1.3.257-260.

11. 1. 3. 232.

11. 1. 3. 233.

11. 1. 3. 213.

82. 1. 28. 1.

81. 1. 1. 3.

11. 1. 3. 214.

636 RODENTIA

6,9. Wormsley, Oxfordshire. W. R. Ogilvie-Grant 11.1.3.215-216

(c & p).
9. Toddington, Bedford- Joseph Scrivner 98.2.27.1.

(albino) shire. (c & p).

6. Welwyn, Hertfordshire. W. R. Ogilvie-Grant 11. 1. 3. 217.

(c & p).
2 juv. al. Hillingdon, Middlesex. O. 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. 401.

9. Sussex. 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. Ti/jic of subspecies.)
2 6,0 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: 6, 4:9. Bversloy, Hampshire. G. S. Miller (c). 7. 7. 7. 2939-40,

2956-7,2972-
3, 2978-9.
2 (5, 2 9. Bembridgc, I. of Wight. W. R. Ogilvie-Grant 11. 1. 3. 221-224.

(c & pj.

9. St. Helens, I. of Wight. O. 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.

Sd, 3 9. Northlew, Devonshire. R. C. Wroughton (p). 11. 1. 3. 227

228, 261-264.

EvoTOMYS GLAREOLUS suEcicus Miller.

1900. Evotoniys hercynicus succiciis Miller, Proc. Washington Acad. Sci.,
II, p. 101, July 26, 1900 (Upsala, Sweden). Type in U.S. National
Museum.

1910. Evotomi/s glarcolus sueciciis Trouessart, Faune Mamm. d'Europe,
p. 171.

Tjjpr hicnlitij. — Up.sala, Sweden.

Geoyrapliical distribution. — Lowlands of Sweden, Finbind and
south-eastern watershed of Norway.

Diagnosis. — Similar to Evotonijis glareohis glareolus, but red
dorsal area narrower and less diffuse, and sides, face, ;ind 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
vi'-^ appears to be considerably less often present (see Table,
p. 6.31).

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

63<

( 16' 4-17' 6). Average and extremes of eight specimens from
Eggedal, Buskerud, Norway : head and body, 97-5 (92-102-5) ;
tail, 54-6 (52-59); hind foot, lG-6 (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.

<5, 9. Medelpad, W. Norrland, Lord Lilford (i>). 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 6. 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).

(5, ?. Asker, Christiania. Christiania IMuseum 93. 3. 1. 14-15.

(E).

EvOTOMYS GLAREOLUS 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., in, p. 419, May, 1909. Type in British Musevmi.

1910. Evotomi/s glareolus istericus Trouessart, Fauna Mamm. d'Europe

p. 172.

Type locality. — Bustenari,' Prahova, Roumania (in the Car-
pathians, north-west of Bucharest ; altitude 480 m.).

Geographical 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 bulhe 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 3'ellowish rufous.

Colour. — Summer pelage : dorsal stripe narrow and well
defined, not tending to spread over sides. It is rufous slightly
Naried with yellowish wood-brown, and rather thickly sprinkled
with black-tipped hairs. Face, cheeks, and sides pale yellowish
\\()od-brown, tinged with grey ; underparts varying from creamy
white to a yellowLsh cream-bufl:". Rump like sides, rather
strt)ngly contrasted with dorsal stripe. Feet greyish white. A
I lark 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

EODENTIA

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.

Skull and teeth. — The skull resembles that of the typical race
except that the auditory bullfe are larger, their inner margins
more nearly approaching each other, and rising more abruptly
above level of basioccipital. Teeth as in E. glareolus glareolus ;
third upper molar with third re-entrant angle on inner side
present in about 64 per cent of fifty-six skulls examined.

3Teasuretnents. — Tyj^e (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 live 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 measurements see Table, p. 651.

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

Germany: Strass, near Burgheim, Bavaria, 5 ; Marxheim, Bavaria, 48
(B.M. and U.S.N.M.).

Austria-Hungahy: Csall6koz-Somorja,Pressburg, Hungary, 1; Hatszeg,
Hnnyad, Hungary, 11.

Roumania: Bustenari, Prahova, 1.

3(5. Strass, Burgheim, Lord Lilford (r). 8.10.19.6-8.

Bavaria. [Kiirbiiz.)
5 c5, 5 ?. Marxheim, Bavaria. Lord Lilford (p). 8. 10. 19. 9-18.

1. Csalloljoz-Somorja, Press- Budapest Mus. (e). 91. 3. 1. 59.

burg, Hungary.
8 (5, 2 ?. 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.
i. Bustenari, Prahova, Rou- Lord Lilford (p). 4. 4. G. 72.

mania. {W. Dodson.) [T'jpe of subspecies.)

EvoTOMYS rjLARHOLUS NORVEGicus Miller.

1900. Evotomiis norvegicus 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 localif I/. —Bergen, Norway.

Geographical distribution. — Western Nor-way, north at least
to Nordland ; apparently confined to the western watershed.

Diagnosis. — Larger than Evotomys glareolus glareolus (hind
foot in adults, IS -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 G39

side ; red dorsal area broad, not well defined from buffy grey
sides of body.

Colour. — Summer pelage : dorsal stripe dull ferruginous
slightly varied with light wood-brown and much darkened bj' a
uniform sprinkling of black-tipped hairs, fading rather abruptly
into the light wood-brown of face, cheeks and sides ; rum]^ wood-
jjrown tinged with red in median line and forming no noticeable
contrast with colour of back ; underpai'ts 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.

SJciiU. — The skull is decidedly larger and more heavily built
than that of Evotomys glareolus glareolus and the other small
races. Zygomata heavy, abruptly spreading anteriorly, 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.

Tectli. — The teeth are larger than those of true Evoiomys
(/Jareolus, 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.

Spechnens examined. — Twenty-six, from the following localities in
western Norway: Mo, Eanen, Nordland, 2 (U.S.N.M.) ; Saltdalen, Bodil,
Nordland, 3; Sondfjord, 1; Bergen, 4 (U.S.N.M.); Opheim, Bergen, 2;
Graven, Hardanger, 14.

3. Saltdalen, Bodo, Norway. Dr. R. Collett (p). 84. 10. 31. 8 -10.

Skull. Sondfjord. " Dr. R. Collett (p). 84. 10. 13. 12.

6, 9. Graven, Hardanger. U.S. Nat. Mus. (e). 5. 8. 5. 1-2.
(Miss Stejnegcr.)

EvoTOMYs GLAREOLUS VASC0NL4: Miller.

1900. Evotoinys vasconix Miller, Proc. Washington Acad. Sci., it, 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

640

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 bufFy.

Measurements. — 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 '6 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.

Sjxciinens examined. — Thirty-four, from the following localities in
France: Port6, Pyr^n6es-0rientales, 11; I'Hospitalet, Ariege, 5; Ax-les-
Thermes, Ariege, 4 ; Caterille, Haute-Garonne, G ; Luchon, Haute-Garonne, 1 ;
Moutr^jeau, Haute-Garonne, 2 (U.S.N.M.); Bareges, Hautes-Pyr6n6es, 5.

(5,2 9.

Port6, Pyrenees-Orien-
tales, Prance.

G

S. Miller (c).

8.

8.

4. 248-250

2 6, ?.

Port6, Pyrenees-Orien-
tales. (.4. Robert.)

0

Thomas (p).

8

9.

1. 59-61.

6, 9.

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

0.

Thomas (p).

8.

9.

1. G2-63.

2 i.

Ax-les-Therines, Ariege.

V.

Builles (p).

8.

3.

27. 12-13.

6, 9.

Ax-les-Thermes, Ariege.

G.

S. Miller (c).

S.

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).

C.

4.

1. 93.

9.

Bareges, Haute s-
Pyr6ness.

G.

S. Miller (c).

8.

8.

i. 251.

EVOTOMYS GLAREOLUS HELVETICUS Miller.

1900. Evotomijs liercynicus helveticus Miller, Proc. Washington Acad. Sci.,
II, p. 98, July 26, 1900. Typo in British Museum.

1910. Evotonnjs glareoliis helveticus Trouessart, Fauna Mamra. d'Europe,
p. 171.

Tyije lociditji. — Montauban, Haute-Savoie, France (near
Geneva, Switzerland). Altitude, 900 m.

Geogra^jhical distribution. — Jura Mountains ; south through
the non-Alpine parts of Switzerland and along the lower western
portion of the French Alps.

Diagnosis. — Intermediate between Evotomys glareolus glareolus
and E. (J. nageri in size (hind foot, 17 to 19 ram., condylobasal
length of skull, 23 to 25*4 mm.), but colour somewhat paler and
more buffy greyish than in either.

Pleasure merits. — 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 Ceneva, 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 IMottaz) ; Geneva,
23 (U.S.N.M. and Mottaz).

Fbance : 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.

s.

Montauban, Haute-

0. Thomas (p). 2. 6. 7. 1.

Savoie, 900m. France.

{Type of subspecies.)

[A. Robert.)

6(5,5?.

Cranves - Sales, Haute-
Savoie. (A. Robert.)

0. Thomas (p). 5. 11. 18. 33-43.

6,9.

Barcelonnette, Basses-
Alpes.

U.S. Nat. Mus. (E). 5. 8. 5. 3-4.

4 <J, 2 9.

St. Paul, Basses-Alpes.
{C. Mottaz.)

0. Thomas (p). • 8.8.10.116-121.

EvOTOMYS GLAREOLUS NAGERI Schinz.

1845. Hypludxus] nageri Schinz, Synops. Mamm., ii, p. 237 (Oberalpsee,

Urserenthal, Uri, Switzerland).
1857. Arvicola glareolus b. Blasius, Saugethiere Deutschlands, p. 337.
1862. Myodes bicolor Fatio, Rev. 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.

Type locality. — Oberalpsee, near Andermatt, Uri, Switzerland.

Geographical distribution. — -Alps (except westernmost jjortion)
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 ai'ea narrow and dark, the sides of body dull greyish in
evident though not conspicuous contrast.

Colour. — Dorsal stripe rather well defined, 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 brow n 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.

SlcuU and teeth. — Skull largest among the Continental Euro-
pean members of the group, £. glareolus hallucalis perhaps excepted.
In form it shows no special peculiarities. Brain-case relatively
large, a peculiarity more 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, 6-5; 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 (6-2-68); 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 examined. — One hundred aud seventy-five, from the following
localities : —

France: 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, 6
(U.S.N.INI. and Mottaz) ; Gordola, Locarno, Ticino, 3.

EVOTOMYS 643

Italy : Padola, Cadore, 1 (Turin) ; Casamazzagno, Cadore, 1 (Turin) ;
Ceresole d'Alba, Turin, 2 ; Stupinigi, Turin, 1 (Turin) ; Moncalieri, Turin,
9 (U.S.N.M. and Turin) ; Ambonasco, in mountains above Chiavari,
Liguria, 2 (Genoa) ; no exact locality, 1.

i, 4 9. Vitznau, Lucerne, Swit- 0. Thomas (c & i>). 5. 8. 3. 22-2G.
zerland.

3 S. Untervatz, Grisons. Lord Lilford (p). 1. 11. 7. 15-17.

2 6, 9. Locarno, Ticino. 0. Thomas (c & p). 5. 8. 2. 15-17.

c5, 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 glareolus Cavanna, Bull. Soc. Entomol. Ital., xiv, p. 87
(Serra Crispo, Monte PoUino, Basilicata, Italy).

1906. Evotomijs nageri haUucalis Thomas, Ann. and Mag. Nat. Hist
7th ser., xviii,p. 221, September, 1906. Type in British Museum

1910. Evotomijs nageri haUucalis Trouessart, Faune Mamm. d'Europe,
p. 168.

Type localitjj. — Santa Eufemia d'Aspromonte, Calabria, Italy.

Geographical distribution. — High mountains of southern Italy.

Diagnosis. — Externally similar to Evotornys glareolus nageri,
but underparts less washed with yellowish brown ; skull with
longer, narrower brain-ca.se 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-buff the colour is similar to that of Evotornys
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 Evotornys
glareolus nageri and the other members of the group in the
decided elongation of the brain-case and a slight shortening of
tlie 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 slightly more slender than usual in

i Evotornys glareolus nageri, though a few specimens from the Alps

are essentially the same. jNIolars 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 excmiined. — The type.

Bemarks. — 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 skomerensis Barrett-Hamilton, Proc. Royal Irish Acad.,

XXIV, p. 316. Type in British jNIuseum.
1910. Evotomys skomerensis Trouessart, Faune Mamm. d'Europe, p. 167.

Tyjpe locality. — Skomer Island, off coast of Pembrokeshire,
Wales.

Geographical distrihufton. — 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 underjaarts ; 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 : upjaer parts a bright light reddish
brown approaching the orange-rufous of Eidgway, 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.

Shill. — The skull differs notably from that of Ecotoviys
glareolus nageri and E. g. norvegicus in the short, broad, rather
strongly ridged and angled brain-case, the con.spicuous 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
anteriorly than in E. glareolus nageri and E. g. norvegicus.
Otherwise the dorsal j^rofile 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. csesarius. 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 E. glareolus norveglcus. Auditory bullae relatively
larger than in the large forms of Evotoini/s glareolus.

Teeth. — The teeth are essentially like those of Evotomys
glareolus nageri ; third 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.M. and
U.S.N.M.).

5 (J, 2 ?. 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. R. Drane (c & p). 11. 1. 3. 265-266.

EVOTOMYS CjESARIUS 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 ]\Iuseum.
1910. Evotomys cxsarius Trouessart, Faune Mamm. d'Europe, p. 169.

Type 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

i more massive than in any of the Continental members of the

I genus, E. rufocanus not excepted.

Colour. — Winter pelage : upper parts a rich, dark, reddish
j 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 ; underparts 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 KODENTIA

duller than in winter, the exact shade nearly hazel ; sides broccoli-
brown, in rather strong contrast with back, but fading insensibly
into the dull buif of underparts. Tail and feet as in winter.

Sl-iiII. — 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. skomerensis ; but the dorsal
surface of brain-case is not flattened as in the Skomer vole.
Interorbital region w^ide, not tending to assume a cylindrical
form as in E. rufocanus, 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. skomerensis. Postorbital processes rather large, but less
3Iicrotus-like than in E. rufocanus and with no trace of a ridge
extending obliquely backward toward parietal such as occurs in
E. skomerensis. 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 bullas
large, not peculiar in form. Zygomata not very abruptly spread-
ing, the median portion of the two arches parallel.

Teeth. — Except for their larger size the teeth are essentially
as in Ecotomys ginreolus. Third upper molar with third inner
re-entrant angle normally present (see Table, p. 631). Anterior
loop of Hij very short.

Measurements. — External measurements of type (not fully
adult male): head and body, 96; tail, 49; hind foot, 18; ear
from meatus, 11. 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.

Specmieyis examined. — Eighteen, all from the island of Jersey.

2 6. St. Helier, Jersev. G. Barrett-Hamilton 3. 2. 11. 1-2.

(c & p). (3. 2. 11. 2. Typeoi species.)

5(5,2 9. Trinity, Jersey. 0. Thomas (p). 8.9.2.12-17.

(R. H. Bnntiufj.)

EVOTOMYS RUTILUS Pallas.
1778. Mils riitilus Pallas, Nov. Spec. Quadr. Glir. Ord., p. 246.
1874. Evotomys rutilus Cones, Proc. Acad. Nat. Sci., Philadelphia, p. 187.
1900. Emtomys rutilus ^Miller, Proc. Washington Acad. Sci., ii, p. 91,

July 26, 1900.
1910. Evotomys riitilus Trouessart, Faune Mamm. d'Europe, p. 169.

Type locality. — Siberia east of the Obi.

Geoijrapltical distribution. — Arctic Asia and Europe, south in
Norway to Tromso, and in Sweden to Norbotten.

* With the possible exception of E. glarcolus hallucalis, adults of which
have not been examined.

EVOTOMYS 647

Diagnosis. — Size medium (hind foot about IS 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-(|uarter as long as vertebraj ; dorsal area clear bright chest-
nut, sharply but not conspicuously contrasted with colour of
sides.

Colour. — Dor,sal .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.

SkiiU. — In general the skull resembles that of Evotomi/s
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 j^alate
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 Evotomys
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 iir^ fully as
deep as second, the anterior portion of the
tooth therefore never assuming the Alticola-l\\ie
aspect characteristic of that region in E. glareo-
lus ; inner side of vv' 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 ; -pm. vie.

Wg showing a decided tendency to assume a Evotom,,s mtihis.
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.il. and U.S.N.M.) ; Tornei^,
Lappmark, 3 (B.M. and U.S.N.M.; Karesuando, Norbotten, 2 (U.S.N.M.).

648 EODENTIA

Bemarlcs. — Externally Evotonii/s rntilus is distingviisliable 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. Hyimd/>'its rufocanus Siindevall, Ofversigt af Kongl. Vetensk. Akad.

Porhandl., iii, p. 122.
1897. Evotomys rufocanus Bailey, Proc. Biol. Soc., Washington, xi, p. 122,

IMay 13, 1897.
1900. Evotomys rufocanus Miller, Proc. Washington Acad. Sci., ii, p. 89, ll

July 26, 1900. j'

1910. Evotomys (Craseomys) rufocanus Trouessart, Fauna Mamm. d'Europe,

p. 173.

Tijpe locality. — Lappmark, Sweden.

GeorjrapMcal distribution. — Northern Europe, south in the
mountains of Norway to Do\Te, east into Asia.

Diagnosis. — Form essentially as in Evotomys glareolus, neither
tail nor ear shortened ; size large, as in E. ceesarius (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, con.spicuously 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 bufiy
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 massivenessthat of any
of the forms of Evotomys glareolus, in this respect essentially
agreeing with that of E. ceesarius. 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 rufocanus. Nat. size.

along anterior suture equal to about one-third length from suture
to foramen magnum. Auditory bull?e rather large, slightly more
narrowed antero-internally than in JEJ. 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. INTandible 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. rutihts. 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 ?h, become completelj' closed.
Anterior loop of )n^ with inner re entrant angle relatively deeper

Efotomim rvfocan »*■.
Enamel pattern, x 5.

650

KODENTIA

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EVOTOMYS

651

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652

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653

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654

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Molar roots moderate.

,, short.
,, moderate.

„ long.
,, moderate.
,, short.
,, moderate,

,, short.
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■ M.OJ-HIOO'i

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O) 1

Number.

86994*
8. 8. 4. 252
153111
153114
172196
172198

1943 Mottaz
2771 „
1507 „
1518
1472
1521
566 „
124443
124431

Q) ..._,_..

§

E. glareolus vasconise.

France : Montr6jeau,Haute-Garoun
Ax-les-Thermes, Arifege
I'Hospitalet ,,
Port6, Pyren^os-Orientales

Bareges, Ilautes-Pyrfin^'cs

E. glareolus helveticus.

Switzerland : Chesieres, Yaud
Bioux ,,

St. Cergues, Vaud

EVOTOMYS

655

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a :;oao -pg

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t-tOOt-COt-COCDt-CDOt-COt-COt-CDC-t-OOt-t-t-t-t-

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658 RODENTIA

than in E. (jlareolus, 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. For 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.) ; Miibxien, Kristiansamt, 1 ; Sundal Fjeld, 1 ; Hyllingen,

Trondhjem, 1 ; Tanen, Finnmarken, 2 (U.S.N.M.).

Sweden: Karesuando, Norbotteu, .3 (B.M. and U.S.N.M.); Lappmark

(no exact locality), 1 ; Tornea, Lappmark, 2 (U.S.N.M.) ; Kvickjoclf , 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
Evotomys csesarms, intermediate in character between true
Evotomys and Craseomys, and the fuller comprehension of the
dental peculiarities of Evotomys rntilus, 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.

i. Stabursnaes, Porsanger. Dr. R. Collett (p). 84. 10. 31. 7.

6 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).

?. Hyllingen, Trondhjem. G. Barrett-Hamilton 11. 1. 2. 114.

(P)-

5. Karesuando, Norbotten, Stockholm Museum 90. 8. 1. 15.

Sweden. (e).

1. Kvickjock, Norbotten. Prof. Sundevall (p). 49. 11. 1. 11.

6. 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 (listrihiition. — Northern portions of northern
hemisphere south in America to southern Mexico and in Europe
to the Mediterranean coast.

MICROTDS 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 ; mamm;e, 8 ; plantar tubercles, 6 ; neither fur,
external form, nor skull modified for aquatic or subterranean life.

Memarlcs. — The genus Microtus as here understood, restricted
to the species with normal skull, palate and enamel folding,
8 mammif, 6 plantar tubercles, and no special modifications of
external form, is the largest and most widely distributed group
in the sub-family. About 100 sj^ecies 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 Eui-ope. Among them are
represented two sub-genera.

Sub-Genus MICROTUS Schrank.

1798. Microtus Schrank, Fauna Boica, i, p. 72 (Microtus terrestris Schrank

= Mus arvalis Pallas).
1817. Mynomes Kafinesque, Amer. Monthly Magazine, ii, p. 45 (Mynomes

pratensis Rafinesque = Arvicola pcnnsylvanicus Ord).
1836. Hnniotomys de S61ys-Longchamps, Essai monographique sur les

Campagnols des environs de Liege, p. 7. Part (arvalis and

terrestris = scherman).
1857. Paludicola Blasius, Saugethiere Deutschlands, p. 33.S. Part (amphi-

bius = schcrman, nivalis and ratticeps). Not of Wagler, 1830.
1857. Agi'icola Blasius, Saugethiere Deutschlands, p. 334 (agrestis).
1867. Praticola Patio, Les Campagnols du Bassin du L6man, p. 34

(included amipliihius = scherman, nivalis, arvalis, ratticeps and

campcstris = agrestis). Not of Swainson, 1837.
1867. Sylvicola Fatio, Les Campagnols du Bassin du L6man, p. 63

(agrestis). Not of Harris, 1782.
1883. Microtus Lataste, Le Naturaliste, ii, p. 348.
1890. Campicola Schulze, Schrifteu Naturwiss. Viereins d. Harzes in

Wernigerode, v, p. 24. Part (arvalis, szibterraneus and campcstris

= 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, Paune 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 portion of the Balkan Penin-
sula, Asia Minor and northern India.

Diai/uosis. — 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 detined.

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 OF THE SUB-GENUS 2IICR0TUS.

Second upper molar with two closed triangles on

inner side M. agresiis, p. 662.

Condylobasal length of skull in largest individuals
27 to 28-4 mm.
First upper molar with third inner triangle
usually present; miderparts 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. levernedii, 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. hirtus, p. 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) ^[. a. neijlectus, p. 675.

General colour of upper parts lighter,

usually with no decided tinge of russet

(West-central Continental Europe).... M. a. baillom, p. 672.

Second upper 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 ou outer side.
Nasals moderatelj' 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. cabrer.i', p. 701.

Condylobasal length of skull about 30 mm. ;
upper tooth-row about 8 mm., the teeth

unusually robust (South-eastern Spain) M. dentatus, ]i. 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.
Rostrum 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. angidaris, p. 706.

Rostrum normal in form (distance from
front of zj'goma 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 bullffi unusually large ; upper
molars with osteodentine spaces re-
duced (Thessaly) M. hartingi, p. 704.

Auditory bullae not unusually large;
upper molars with osteodentine
spaces of normal width.
General form of skull narrow and
agrcstis-hke ; 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. sarniiis, p. 700.
General form of skull broad, not
agrestis-Uke ; 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 buSy grey (Ork-
ney Islands).
Brain-case not flattened, the ratio
of occipital depth to width
about 57 ; condylobasal length
of largest skulls 28 to 2'J'6 mm.

(South Orkney Islands) M. oixadensis, p. C94.

Brain-case flattened, the ratio of
occipital depth to width about
53 ; condylobasal length of

662 RODENTIA

largest skulls 26-6 to Si7"6 mm.

(North Orkney Islands) M. sandayensis, p. 696.

First lower molar with anterior
outer re-entrant angle obso-
lete ; back decidedly greyish ;
underparts scarcely tinged
with yellowish brown (Sanday

Island) M. s. sandayensis, 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. uTst);v, 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 (.41ps) 31. incertus, 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. asfiii-iauus, 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. lems, 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. dujAicatits, 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. arralis, p. 683.

Size slightly larger and colour a
little more yellowish (Pyre-
nees) M. a. meridianus, J). 686.

MICROTUS AGRESTIS Liimjeus.

(Synonymy under subspecies.)

Geographical distribution. — Northern and central Europe fi'om
northern Scandinavia to the Alps, Pyrenees, Galicia and Portugal,
and from England, Scotland and the Hebrides eastward.

MICROTDS

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
measured 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 3| 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-thii;d 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 i-egion 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 sufficiently 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

RODENTIA

distribution or texture, its quality somewhat harsh and loose.
Mamrn* : p 2-2, « 2-2 = 8.

SA-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
fiattish, 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 jjarietals marked in old age by well developed
longitudinal ridges which are continuous po.steriorly 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 basioccijjital 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. Z3-gomata 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

I'lfJ. 129.
Microtus agrestis. Nat. size.

MicKoxns 665

wall of canal rudimentary below, absent above. Rostrum slender,
scarcely wider than interorbital region, rather deep proximally
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 j^remaxillaries ; incisive foi-amina 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 with
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 fr-om 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 ai-ea rather deeply but simply hol-
lowed, the anterior (enamel) edge entire and
nearly straight when viewed from in front,
t>bli(juely flattened and with rather abruptly ^^^ ^^^

hack ward-curved extremities when viewed mtrotiisagrestis.
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 antcro-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 defined, with sharp
salient and re-entrant angles ; upjjer 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 !«', but wider externally than internally, its long axis
nearly transversfe to tooth-row, its length nearly equal to greatest
width 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 7ii^, 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 limlj 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 Hehridean
form, M. agresiis cxsiil, 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 667

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 re-entrant 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
Ttiy 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 w*, and nu, 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.

Bemarks. — 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

merabei' of the sub-genus is Microtus arvalis, fully adult
individuals may be recognized by their larger size, more hair}'
ears, larger tubercles on sole, and by the browner, less buify
coloration, the dark and light elements of which are more
coarsely mixed. As might be anticipated from the animal's wide
range, numerous local forms have been developed. Seven of
these are now known.

MiCBOTUS AGRKSTIS AGRESTIS LinUfCUS.

1761. Mils agrestis Linnaeus, Fauna Suecica, 2nd ed., p. 11 (Upsala,

Sweden).
1766. \JiItis] gregarius Linnaeus, Syst. Nat., i, 12fcli ed., p. Si (Germany

and Sweden).
1792. Mils arvalis nigricans Kerr, Anim. Kingd., p. 239 (Renaming of Mus

agrestis).
1820. Lemmus arvalis Nilsson, Skand. Fauna, i, p. 189 (not Mas arvalis

PaUas).
1841. Arvicola agrestis de Selys-Longchamps, Bull, de I'Acad. Royale des

Sci. des Arts et Belles-Lettres de Bruxelles, viii, pt. 2, p. 236.
1844. L[(:vimus] insiilaris Nilsson, Ofversigt af Kougl. Vetensk.-Akad.

Forhandl., Stockholm, i, p. 34, March 20, 1844. (Ostgotha

Skarg&rd, Sweden.)
1857. Arvicola agrestis a. Blasius, Siiugethiere Deutschlands, p. 369 (part).
1884. Microtus agrestis Lataste, Ann. jMus. Civ. Stor. Nat. Genova, xx,

p. 255, March, 1884.
1896. Microtus agrestis Barrett-Hamilton, Proc. Zool. Soc, London, p. 602.
1910. Microtus agrestis Trouessart, Faune Mamm. d'Europe, p. 175 (part).

Tyjje locality. — -Upsala, Sweden.

Geographical distribution. — Scandinavian Peninsula ; eastward
into Finland. Exact limits of range not known.

Diagnosis. — Size lai'ge (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 bistei', 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.

Sktill 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 eijual to zygomatic breadth.
Teeth with no special peculiarities ; m^ 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.

R. J. Cuninghame 98. 5. 2. 6-9.

(c & p).
R. J. Cuninghame 98. 2. 28. 6-7.

(c & p).
G. Barrett-Hamilton 11. 1. 2. 46-47.

(c & p).
E. Lort Phillips 94. 10. 22. 1.

(c & p).
Ohristiania Museum 93. 3. 1. 12.

(E).

Miller Collection. 7. 7. 7. 45.37.

R. J. Cuninghame 8. 8. 9. 80-37.

(c & p).

Baron E. de S61ys- 45. 7. 5. 12.

Lougchamps (p).

Lord Lilford (p). 0. 5. 15. 2-5.

Prof. Sundevall (c). 45. 10. 25. 5-6.

MiCROTus AGRESTis ExsuL Miller.

1908. Microtus agrcstis cxsul 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.

Ty2}e locality. — North Uist, Hebrides, Scotland.
Geograplnccd distribution. — North and South Uist, Hebrides.

* In a little more than five per cent, of the specimens examined.

4 6.
2 6.

Molmen, Kristiansamt,

Norway.
Holaaker, Kristiansamt.

2 6.
1.

Hjerkin, Kristiansamt.

(N. F. Ticehurst.)
Sundal Fjeld.

1. .

Gausdal.

1.
3 6,5 9.

Gausdal.
Mandal.

1.

Jemtland, Sweden.

4 6.
6, 9.

Upsala. {G. Kolthoff.)
Stockholm.

670

Diagnosis. — Size and cranial characters as in the large
Microttis 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 Mrtus 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
1.36 skulls.

Measurements. — External measurements of

type (adult female) : head and body, 1 23 ;

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.1\I. and Edinburgh) ; Loch Boisdale, South
Uist, 3 (Edinburgh).

Bemarlcs. — 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

6, 3 ?. N. Uist, Hebrides. J. F. Davison (c & p). 6. 3. 1. 1-4.

(6. 3. 1. 3. Ti/pe of subspecies.)
1 al. N. Uist. J. A. Harvie Brown (p). 79. 9. 18. 2.

Microtus mjVeMs exsitl.
Enamel pattern, x 5.

* 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. 4.58-512, May 4, 1907, and Naturen (Bergen), sxxii,
pp. 193-202, 269-277, July-September, 1908.

671

MlCROTUS AGRESTIS LEVERNEDII Crespon.

1844. A[rvicola] levernedii Crespon, Paune M6ridionale, i, p. 73 (Marshes

between St. Gilles and Aigues-iNIortes, Gard, France). Type in

Nimes Museum.
1857. Arvicola agrestis a. Blasius, Saugethiere Deutschlands, p. 3G9 (part).
1869. [Arvicola agrestis] var. 7iigra Fatio, Fauna Vert. Suisse, i, p. 241

(Engstlen, Berne, Switzerland. Alt. 1750 m.). Type in Geneva

Museum.
1900. [Arvicola ag^-estis] riifa 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 IMuseum.
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).

Ty2}e locality. — Marshes between St. Gilles and Aigues-Mortes,
Gard, France.

Geo(jraphic(d dlstiihution. — 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 : —
Feance: 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.T*!.) ; Etupes, Doubs, 1.

Switzerland : Geneva, 9 (Mottaz and Geneva ; types of latifrons Fatio
and rufa Fatio 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 7iiger Fatio) ;
Degersheim, St. Gallen, 1 (U.S.N.M.) ; Chur, Grisons, 1 (U.S.N.M.) ;
Oberholsen Valley, 1.

Eemarlis. — 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

RODENTIA

matei'ial 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 specimens 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 (J. St. Gilles, Gard, France. G. S. Miller (c). S. 8. 4.233-2.3G.

6. ]\Iontauban, Haute-Savoie, O. Thomas (p). 6. 4. 2. 7.

900 m. {A. Robert.)

6,9. C r an ves - Sales, Haute- A. Robert (c & p). 5.4.9.8-9.
Savoie, 900 m.

5(5,9. Cranves-Sales, Haute-Savoie, O. Thomas (p). 5.11.8.24-27,

1200 m. (.4. Robert.) 29, 32.

?. Lucinges, Haute-Savoie, O. Thomas (p). 6.4.2.7-8.

900 m. (A. Robert.)

9. Etupes, Doubs. (C. Mottaz.) O. 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 dc Sclys-Longchamps.

1841. Arvlicolci] baiUonide Selys-Longchamps, Atti della seconda Riuuione

degli Scienziati Italiani, Torino, 1840, p. 225 (Northern France;

Abbeville, Somme, from context).
1845. '? A[rvicola] intermedia Bonaparte, Atti della sesta Riunione degli

Scienziati Italiani, Torino, 1844, p. .350 (cited as doubtful synonym

of bailloni, with de S61ys-Longchamps as authority). Nomen

nudum.
1857. Arvicola agrestis a. Blasius, Siiugethiere Deutscblands, p. 369 (part).
1896. Microtiis agrestis negledus Barrett-Hamilton, Proc. Zool. Soc,

London, p. 602 (part).
1910. Microtus agrestis neglectus Trouessart, Faune Mamm. d'Europe,

p. 176 (part).

Type locality. — Abbeville, Somme, France.

Geographical didribution. — 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.

Diagiwui.'i. — Similar to Microtus agrestis agrestis but smaller
(hind foot, 17 to 18 "6 mm.; condylobasal length of skull in
largest individuals, 24 '5 to 26 '6 mm.); skull with brain-case
tending to assume the narrower form characteristic of 31. agrestis
levernedii.

Measurements. — Adult male from Hillerod, Zealand, Denmark:
head and body, 113; tail, 35; 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, 8eine-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 : —

Denmabk: Hillerod, Zealand, 3; Skansen, LoUand, 1 (U.S.N.M.).

Gebmany : Brunswick, 28 (U.S.N.M.).

Phance : Guines, Pas-de-Calais, 1 ; Dinan, C6tes-du-Nord, 1 ; Manon-
ville, Meurthe-et-Moselle, 1 ; Barbizon, Seine-et-Marne, 3 ; Mont Dore,
Puy-de-D6me, 1 ; Solf6rino, Landes, 1 ; Poret-de-Bouoonne, Gers, 1 ;
Port6, Pyr6n6es-Orientales, 2; I'Hospitalet, Ariege, 1; Pyrenees, no exact
locality, 1.

6, 2 9. Hillerod, Zealand, Den- 0. Thomas (c & p). 98. 6. 7. 20-22.
mark.
6. Guines, Pas - de - Calais, O. Thomas (c & p). 94. 6. 6. 18.

Prance.
6. Manonville, Meurthe-et- Lord Lilford (p). 1. 11. 7. 12.

Moselle. (Lomont.)
6, ?. Barbizon, Seiue-et-Marno. G. S. Miller (c). 8. 8. 4. 231-232.

6. Solf6rino, Landes. O. Thomas (p). 6. 4. 1. 75.

(A.Bobert.)
9. Foret de Bouconne, Gers. O. Thomas (p). 6. 4. 1. 76.

(A. Bobert.)
(5,9. Port6, Pyr6nees-0rientales. G. S. Miller (c). 8.8.4.229-230.

9. L'Hospitalet, Ariege. G. S. Miller (c). 8. 8. 4. 228.

(5 juv. Mt. Dore, Pay -de -Dome, G. S. Miller (c). 8.8.4.227.

4500 ft.

MiCROTUS AGRESTis HiRTUS Bellamy.

1839. Arvicola hirta Bellamy, Nat. Hist. South Devon, p. 373 (Devonshire,
England).

1817. A{rvicola] britannicus de Selys-Longchamps, Revue Zoologique,

1847, p. 307, October, 1847 (England).
1847. \_Arvicola\ britatitiicus de S61ys-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 neglcctus Barrett-Hamilton, Proc. Zool. Soc,
London, p. 602 (part).

1910. Microtus agrestis neglcctus Trouessart, Pauue ]Mamm. d'Europe,
p. 176 (part).

Type locality. — Yealmpton, Devonshire, England.

GeofjrapMcal distribution. — England and the lowlands of
southern Scotland.

Diaijnosis. — Size as in Microtus agrestis bailloni 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

674

KODENTIA

I'IG. 132.
Microtus agrestis liirtus. Nat. size.

tending to assume the short squarish form characteristic of
true agrestis ; vipper 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-
bvift* though not so bright ; sides paler
and with a noticeable huffy tinge ;
underparts light ochraceous-buff irre-
gularly darkened by the slaty bases of
the hairs ; feet and under surface of
tail concolor with belly ; ujaper 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,
1.10 (108-112); tail, 29-2 (27-32); hind foot, 16-5 (16-18).
For cranial measurements see Table, p. 678.

Specimens examined. — One hundred find thirty-four, from the following
localities : —

Scotland : Windygates, Fife, 4 ; Stockbriggs, Lanarkshire, 2 ; Black-
wood, Lanark, 7 ; Crieff, 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,
Lincolnsliire, 5 ; Cheadle, Staffordshire, 1 ; Anglesey, Carnarvonshire, 1 ;
St. Bride's, Pembrokeshire, 1 ; Sandringham, Norfolk, 1 ; l\Iethwold Fen,
Norfolk, 3; Lowestoft, Suffolk, 1; Kugby, 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 ; Southcrndo wn, near Bridgend, Glamorgan-
shire, 1 ; 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.
2 6, ?. Wyseby, Dumfriesshire, Miss D. Bate (c & r). 11. 1. 3. 267-269.

Wyseby, Dumfriesshire,
Scotland.

1. Drumlaurig, Dumfries-
shire.

?. Crieff, Perthshire.

6, 6 juv. Stockbriggs, Lanarkshire.
?. Hawick, Roxburghshire.
? imm. St. Brides, Pembroke-
(albino) shire, Wales.

Sir W. Jardine (c). 86. 7. 2. 11.

W. R. Ogilvie-Grant

(c & p).
E. R. Alston (c & p).
J. E. Harting (c & p).
Hon. C. Edwards

(c & p).

88. 5. 30. 1.

79. 9. 25. 54-55.
11. 1. 3. 289.
90. 12. 5. 2.

MICROTUS

G75

9 al. Mars ham, Yorkshire, W. B. Tegetmeicr (p). 93.8.31.1.

England. (J. Carter.)

2 S. Bowdon, Cheshire. G. Barrett-Hamilton 11. 1. 2. 48-49.

(T. A. Coward.) (p).

3 S, 9. Grimsby, Lincolnshire. G. Barrett-Hamilton 11. 1. 2. 50-53.

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

2.5,2 9. Grimsby. (G. E. Caton W. E. de Winton (p). 11.1.3.246-249.
Haigh.)

6, 9. Cheadlo, Staffordshire. E. W. H. Blogg (p). 11. 1. 3. 287-288.

6. Anglesey. T. A. Coward (c & p). 11. 1. 3. 270.

2.5,19. Oundle, Northampton- Lord Lilford (c & p). 11.1.3.271-273.

shire.

2(5,2 9. Graftonbury, Hereford- W. E. de Winton 11.1.3.274-277.

shire. (c & p).

S. Shelford, Cambridge- Dr. Hans Gadow 5. 1. 21. 1.

shire. (c & p).

2 St. Cambridgeshire. J. Baker (c & p). 39. 9. 29. 4-5.

3(5,4 9. Earlsfield, Surrey. C. H. B. Grant (c (fc p). 11.1.3.278-284.

i6. Alum Bay, Isle of Wight. O. Thomas (c i& p). 11.1.3.285-286.

1. Bonohurch, Isle of Wight. Rev. C. A. Bury 44.9.2.5.

(c & p).

19. New Forest, Hampshire. G. S. Miller (c). 7. 7. 7. 2900-

2924.

MiCROTUS AGRESTis NEGLECTUS Jenyns.

1841. Arvicola negledus Jenyns, Ann. and Mag. Nat. Hist., 1st ser., vii,

p. 270, June, 1841.
1857. Arvicola agrcstis b. Blasius, Saugethiere Deutschlands, p. 369 (part).
1896. Microtus agrcstis neglectus Barrett-Hamilton, Proc. Zool. Soc.,

London, p. 602 (part).
1910. Microtus agrcstis ncglectiis Trouessart, Faune Mamm. d'Europe,

p. 176 (part).

Type locality. — Moors near Megarnie Castle, Pertlishire,
Scotland.

Geographical distribution. — -Highlands of Scotland.

Diagnosis. — Like Microtus agrcstis 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 bailloni. 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.
Uiiderparts 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 3T. agrestis hirtus, though it shows no peculiarities of form.
Some of the larger skulls might readily be confused with those
of small individuals of 31. agrestis agrestis or M. agrestis exsul ;
but their true identity is shown by the small auditory bullae,
which I'ctain the small size characteristic of the British races
and M. agrcstis bailloni. The teeth show no peculiarities ; m^

2x2

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,, joined, high.

,, ,, low.

to to to CO CO i to to lO CO to CO to o

.<.IB([!XBJiI

Cq-*CNTHCN-*Ttl (NO(M<MC<l^C>1

totococotototo cotototocoto to

■aiqipiiuK

otMOoiooco oooqocooo oq

CO to o CO to to lo to to to to in lo m

tHt— It— ItHt— ItHtH tHtHtHt— li-lrH T-H

■BlUa^SBIQ

^tOtO-*tD-*0 (MCOOOCM^-^ O
t- t^ t- L~ t^ t- L^ t* t^ t^ t- t^ t^ t-

'IBSBil

COtOThOCM^CM 00^-*.O<MCXl
t^t^t-t-t^L^t- tOt*t-'t*t-tO

•(treipsm)
inilap [BCiidtaao

O 1 00 to ^ '^ <M CD O CM oq o ^ o

to 1 to to CO CO to ci> cb CO CO cjo to to

•qipBajq
IB^uIiooo

tOO-lCMCOO'*^ ^tOO<M_uO-*

TH<yililAlC--lCN>H ,M,H(MrH.-HCM tH
rHiHi-lT-liHiHT-l i-HrHiHi-l^rH i-l

•IIOJIOU^SUOO

[Bijq.i'oja^ui

(MtOtOtOCO^-* <MOCN<MO'*<M

comcocooocOOT cocococococo co

•qipissjq
.ii^biuoSjCz

(NCOCMOOOO O-^CNOOOO
rHi— trHT-iiHT-Hi-l tHtHt— ItHtHtH r-i

■mSiiai
[BSBqo[^puoo

00"Hc000tOTj(CM '*tD_j^tOOCO-*

in-in»b>ncb>n io>ocD-*»oici'*

<MiOeM<MC<l(M<M <MCq<M(MCM(MCq
CM

Sex.

*0*0*0*0*0^*0 O'O'^O* *T3

^

98. 6. 7. 19
85270
85275
85652
85654
85658
85659

51 Kinnear

133

169

11. 1. 3. 269
11. 1. 2. 48
11. 1. 2. 49

11. 1. 3. 270

Locality.

M. agrestis bailloni.

Denmark: Hillerud, Zealand .
Germany : Brunswick .

M. agrestis hirtus.

Scotland : Lanarkshire .
Pifeshire

Dumfriesshire
England : Bowdon, Cheshire .

Anglesey, Carnar- 1
vonshire . . j

679

:33

■* CO CD CD O •* 0_U
CM cq eqcNtN <N (M(N

>~,r0 p~, ro

00 o ooo oq oo ■* oq ■* , , -^ o

ICCDCOCDCOCOCOCO COCOCD' ' CDCD

O cq o 00 <M ■* cq oq -* CO co , , cx) •*
cococoiOcocDcDco cococo' ' coco

CO -* CD 00 CD o CD 00 ■* o cq oo
lOioiOiOiocDioio iccocdI I coco

rHr-li-HiHiHiHiHTH iHiHrH i-HiH

(NCNO'^CTfMoioo ooqcq. , <o ^

t- t- t- t- t- t- t- CO t- t- t- ' ' t- t-

O o 00 oq 00 o oq CO • oo o oo , , ■* co

t- t- CO t- CO t- t- CO CO t- CD ' ' l>- t-

cDOcD-*oooo-«Ji Ooqcq. . .co

CDCDCDCOCOCOCOCO CDcOco' ' 'cD

CO o 00 Tj< ^ cq cq ■* cq oo cD -*

iHTHiHiHrHT-ICqiH t-iAht-iI I lAl
t-lrHT-l>Hi-lTHiHrH iHtHtH iH

cqcqcqcq^cqcqrjf -^oq-*, . cooo
cocococococococo cococo ' coco

O O cocDn CD ooo ■* -* ooo

THT-liHiHTHi-lrHrH iHrH iHtH

O -* ■* CD "tl

... -CO

lO lO lO CO

cq cq CT cq CO

CM- "o 'O

.-I t-00CT5OC0G0Cq

t- t- t- t- 00 o th 00

ci cq cqcqcq o ct>c^

cq cq ci

CO CO CO CO CO . . .

t- t~ t-

tH iH rH tH .H . . .
t- t:~ t-

rH rH r-l T-l iH . . .

tH 1-1 T-lrtiH t- t-t~

cq CO

^^O

_j _; M
t-i i-i ^-j

00 00 ""*

■^ CO

'K

„

rthamp
aftonbu
ord .
rlsfield,

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.^2

O w •" c8

OJ CD

IZhO H

:zi

ho ^

2 3

iJ C8

680 RODENTIA

with postero-internal loop present in the usual small jiercentage
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, 46 ; 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, Rannoch, Perthshire, -3.

cJ, 2 ?. Beauly, Inverness-shire, Hon. Margaret Eraser 8.1.4.1-3.

Scotland.

(c & P).

2al.

Ben Nevis, Inverness.

W. R. Ogilvie-Grant
(c & P).

82. 10. 6. 1-2

1 al.

Stonehaven, Kincardine-

E. Lort Phillips

85. 10. 5. 1.

shire.

(c & P).

2 6, ?.

Rannoch, Perthshire.

Hon. N. C. Roths-

8. 9. 14. 1-3.

{F. J. Cox.)

child (p).

?.

Dunphail, Elgin.

W. R. Ogilvie-Grant
(c & P).

11. 1. 3. 405.

9.

Elgin.

W. R. Ogilvie-Grant
(c & P).

11. 1. 3. 404.

MiCROTUS AGRESTI.S ROZIANUS Bocage.

1865. Arvicola roiianns Bocage, Mem. Acad. Real das Sciencias de Lishoa,

N.S., III, pt. 2, p. 7 (articles separately paged).
1910. Microtus agrestis rosiamis Trouessart, Fauue JIannn. d'Europe,

p. 177.

Tijpe locallfij. — CJeria, near Coimbra, Portugal.

(leograjjliical dislr'dmtion. — Portugal and north-western Spain.
Limits of range unknown.

Diagnosis. — Size and general appeai'ance, so far as is known,*
as in Microtus agrestis hailloni ; skull with auditory bulhe notice-
ably flattened, and so reduced in size that the greatest diameter
(from paroccipital process to tip of anterior spine) is contained
31 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 form the skull agrees with that of the
other small races of Microtus ngrestis, but it is immediately
recognizable by the small, slightly inflated auditory bullaj, 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 been altered by the action
of alcohol.

MICROTUS 681

Teeth. — While in no way essentially different from those of
the other small races of Microtus agrestis 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.

3Teasurements. — 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 the following localities in Spain :
La Coruna, 1 ; Villalba, Lugo, 2.

2al. Villalba, Spain. Dr. V. L. Seoane (c & p). 94.1.1.14-15..
1 al. Coruna. Dr. V. L. Seoane (c & p). 95. 4. 29. 3.

MICROTUS ARVALIS Pallas.

(Synonymy under subspecies.)

Geographical distrihution. — Continental Europe from the
Baltic to the Pyrenees and northern Italy, and from the Atlantic
coast eastward.

Diagnosis.— 8ize 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 i"anging 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 rounded 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 j^rocess low, not distinctly angular ; inter-
parietal with antero-posterior diameter, exclusive of median
projection, obviously more than half transverse diameter, the

682

^is/

Fig. 133.
Microtus arvalis. Xat. .size.

exact proportion of length to breadth varying in the different
races ; auditory buUie frequeiitly though not always larger than in
31. agrestis (proportionately to size
of skull) ; nasals narrowing 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 aj^pearance. In details of enamel folding the
only imjiortant difference between the two
animals is the comj^lete absence in M. arvalis
of a postero-internal loop to m^. The third
upper molar has exactly the same elements
as that of 31. agrestis, and is subject to
similar variations in form. First lower
molar with re-entrant angle on inner side of
anterior loop usually less developed than in
31. agrestis, that on outer side somewhat
deeper, so that the two are approximately
equal ; this causes the loop to apj^ear to
project forward, or to turn outward instead
of inward.

Semarl's. — 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
3t. agrestis, dry sj^ecimens, 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 Boruer, Arbeiten aus der Kaiserlichen Biolog-
ischen Anstalt fiir Land- und Forstwirtschaft, v, Heft ii, i^p. 37-89,
pis. iv-vi, 1905.

FIG. 134.

ifierottis arvalis.

Enamel pattern, x S.

G83

MiCROTUS ARVALIS ARVALIS Pallas.

1778. Mus. arvalis Pallas, Nov. Sp. Quadr. Glir. Ord., p. 78 (German}').
1801. ilf[?/s] arv[alis] albtis Bcchstein, Gemeinu. Naturgesch. Deutschlands,

I, 2nd ed., p. 998 (Thiiringen, Germany).
1803. LeDWins fiilvus Geoffrey, Catal. Mammif. du Mus. Nat. d'Hist. Nat.,

Paris, "p. 187 (France).
1822. Arvicola vulgaris Desmarest, llammalogie, pt. ii, p. 282.

1840. Arvicola arvensis Schinz, Europ. Fauna, i, p. 60 (Substitute for

arvalis) .

1841. [Arvicola] arvalis de S61ys-Longchamps, Bull, do I'Acad. Royale 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. ater de S^lys-Longchamps, Atti della sesta
Riunione degli Scienziati Italiaui, Torino, 1844, p. 321 (Nomen
nudum).

1847. ?? Arvicola cunicularius Ray, Rev. Zool., p. 312, October, 1847
(Riceys, near Troyes, Aube, France).

1853. Arvicola campcstris Blasius, Gelehrte Anzeigen, Miinchen, xxxvii,
p. 106, July 29, 1853 (Brunswick, Germany). For identification
with arvalis see Rorig and Bonier, Arbeiteu aus der Kaiserlichen
Biologischen Anstalt fiir Land- und Forstwirtschaft, v. Heft ii,
pp. 74-75, 1905.

1857. Arvicola campestris Blasius, Siiugethiere Deutschlands, p. 375.

1857. Arvicola arvalis Blasius, Saugethiere Deutschlands, p. 379.

1884. Microtus arvalis Lataste, Ann. Mus. Civ. Stor. Nat., Genova, xx,
p. 259, March, 1884.

1905. Arvicola arvalis, galliarcli Fatio, Arch. Sci. Phys. et Nat., Geneve,
4th ser., xix, p. 197, February 15, 1905 (Bulle, 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 ccmtigiia Rorig and Borner, Arbeiten aus der
Kaiserlichen Biologischen Anstalt fiir Land- und Forstwirtschaft,
V, Heft II, p. 76 (Rotheuburg, 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 dcprcssa Riirig and Borner, Arbeiten aus der
Kaiserlichen Biologischen Anstalt fiir Land- und 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 principalis 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 ]\Iamm. d'Europe, p. 173 (part).

1910. Microtus agrestis camp)cstris Trouessart, Faune Mamm. d'Europe,
p. 176.

Ti/jie locality. — Germany. (Under the name arvalis Pallas
included the small ^'oles of the arvalis and agrestis type occurring
in the rei^ion extending from England into western Siberia.

684 EObE^TlA

Germany is one of the localities mentioned in the text (p. 78),
and to a form occurring in Germany the name arvalis 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
suQused 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 bufF, 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.

SJiuU and teeth. — The skull of typical Microtus arvalis may
):)est be described as lacking any special peculiarities. It is
moderately long and nai'row, 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 slojjing forward at an
angle of about 23" ; interorbital ridges uniting to form an evident
crest in fully adult individuals ; auditory bullaj rather large,
smootlily inflated. Teeth small, the enamel pattern normal,
tliough 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 andl2.
For cranial measurements see Table, p. 688.

Specimens examined. — One hundred and seventj'-nine, from the following
localities : —

Belgium : Li6ge, 1 ; Waremme, Ijii-ge, 6 (U.S.N.M.) ; Maredsous,
Namur, :3.

France: l':tupes, 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 (ilottaz) ; Foret de Bouconne,
Gers, 1.

Germany : 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) ; Valine de Joux, Vaud, 1 (IMottaz) ;
Chesi^res, Vaud, G (Mottaz) ; Lausanne, Vaud, 4 (U.S.N.M.) ; Neuchatel, 1
(U.S.N.M.); Bulle, Fribourg, 1 (Geneva; type of (jalliardi Patio); Mei-
ringen, Berne, 1 (U.S.N.M.); Vitznau, Lucerne, 1; Ziibervyangen, St.
Gallen, 11 (U.S.N.M. and Mottaz); Mels, St. GaUen, 3 (U.S.N.M.) ; Val
Tours, Grisons, 1 (U.S.N.M.).

Italy : Milan, 1 ; Fiorentina, Bologna, 3 ; Ferrara, 1.

1. Li^ge, Belgium.

2 6, 9. Maredsous, Namur.
5 (5, 9. Etupes, Doubs, France.
(C. Mottaz.)
2 6. Cranves - Sales, Haute-
Savoie.
6, 2 ?. Cranves - Sales, Haute-
Savoie. (.4. Robert.)
9. Foret de Bouconne, Gers.
(.4. Robert.)
2 6. Schwarzburg, Thiiringen,
Germany. (Schuchardt.)

2 9. Ummerstadt, Thiiringen.

[Schuclmrdt.)
i. Rudolstadt, Thiiringen.

(Schiichardt.)
6. Kalbe, Saxonj'.
9, 2 juv. Magdeburg, Saxony.

al.
2(5,3 9. Magdeburg. {Dr. Woltcr-
siorff.)
1 al. Tharandt, Saxony.

{Nitschc.)
6, 2 9. Niesky, Silesia.

2. Niesky, Silesia. (W. Bacr.)

6. Marxheim, Bavaria. (Dr.

Woltcrstorff.)

3 6. Strassburg, Alsace.

(C. Mottaz.)
2al. Colpin,JIark Brandenburg.

7. CsaUokoz-Somorja, Press-

burg, Austria-Hungary.
<J, 2 9 al. Prag, Bohemia.

6. Geneva, Switzerland.

6. Vitznau, Lucerne.
6, 2 9. Fiorentina, Bologna, Italy.

1. Milan.

Baron E. de S61ys-
Lougchamps (p).
Piev. G. Fournier(p).
O. Thomas (p).

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 IMuseum (e).

Dr. E.Hamilton (p).
Lord Lilford (p).
Lord Lilford (p).

O. Thomas (p).

Dr.H. Gadow(c&p).
Budapest Museum

(E).

V. Fric (p).
E. R. Alston (p).
O. Thomas (c & p).
Genoa Museum (k).

Baron E. de S61ys-
Longchamps (p).

37. 1. 3. 173.

3. 3. 30. 3-5.
8.8.10.106-111.

5. 4. 9. 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. 8-4.

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

MiCROTUS ARVALIS MERIDIANUS Miller.

1908. Microtus arvalis meridianus Miller, Ann. and INIag. Nat. Hist.

8tb ser., i, p. 197, February, 1908. Tyi^e in British Museum.
1910. Microtus arvalis meiidianus Trouessart, Paune Mamm. d'Europe,

p. 174.

Type locality. — Near Biarritz, Basses-Pyrenees, France.

Oeograpliical distrihution. — South-western France, in and near
the Pyrenees.

Diagnosis. — Like Microtus arcalis 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.

Slciill 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, 115; tail, 32 ; hind foot, 16-6; ear, 1 2. 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' 8-16 -6) ; ear, 11 -2 (10-12). For cranial measurements
see Table, p. 689.

Specimens examined. — Eleven, from the following localities in south-
western France: Porte, Pyren<-es-0rientale5, 1; Pic du Midi, Hautes-
Pyr6n6es, 1 skull (Lalaste) ; Biarritz, Basses-Pyr6n6es, 8 ; Pyrenees, no
exact locality, 1.

9. Port6,Pyron6es-Orientales, 0. Thomas (p). 8.9.1.70.
France. (A. Robert.)

2 6, 4 ?. Biarritz, Basses-Pyr6nees. J. F. Davison (c & r). 6. 6. 4. 23-28.

(6. 6. 4. 2G. Ty2x of subspecies.)

1. Pyrenees. Baron E. de S61ys- 45. 7. 5. 6.

Longchamps (p).

1 Tomes Collection. 7. 1. 1. 137.

Microtus arvalis duplicatus Rorig and Biirner.

1905. [Arvicola arvalis~\ forma duplicata Eorig and Burner, 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 buffy and less uniform ; 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 underfur 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.

Slull 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 bulhe large and well inflated. Teeth showing no
special peculiarities.

Measurements. — Avei'age 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 Konigsberg, 1 (U.S.N.M.) ; Ten-
kitten, 40 (U.S.N.M.); Frische Haft, 2 (U.S.N.M.); Sanglienen, 4
(U.S.N.M.); Legehueu, 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 foi^ms of Microtus arvalis by its large size, grizzled
greyish colour, and robust, strongly angled skull. So far as at
present Ifnown it is confined to the coast region of extreme
eastern Germany.

Microtus arvalis levis Miller.

1908. Microtus levis Miller, Ann. and Mag. Nat. Hist., 8th ser., i, p. 197,

February, 1908. Type in British Museum.
1910. Microtus levis Trouessart, Faune ]\Iamm. d'Europc, p. 182.

Type localitij.- — Gageni, Prahova, Roumania (at foot of
Carpathians, north-west of Bucharest).

Geograpliical distribution. — Roumania, southern Hungary and
north-eastern Italy ; limits of range not known.

Diacjnosis. — Size and external appearance essentially as in
3T. arvalis duplicatus ; skull narrow and rounded, the interorbital
region not developing a distinct ridge until late in life ; brain-case
lung and narrow, its length measured from interorbital constric-
tion to condyle greater than zygomatic breadth.

688

RODENTIA

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EODENTIA

Colour. — The colour is less yellowish than in M. arvalis
arvalis, the light element a pale cream-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, huffy white below. Underparts in most specimens
rather strongly suffused with pale cream-liuff'.

SJcull 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 nrralis, in this respect
resembling dujtlicatus. 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 usually 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 Wy, 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 : —
AusTBiA-HuNGABY : Cepiii, near Eszek, Hungary, 8.
RouMANiA : Gageni, Prahova, 17 ; Bustenari, Prahova, 1 ; near
Bucharest, 3 (Mottaz) ; Dobrudsoha, 1.
Bulgaria : Bustschuk, 1 (Andersen).
Italy: Caorle, Venice, 1.

2&Gal. Cepin, Eszek, Hungary. Budapest Mnseum(E). 94.7.23.1.3-20.

6.i,6 9. Gageni, Prahova, Kou- Lord Lilford (p). 4.4.6.53-63.

mania. (17. Dodson.) (4. 4. 6. 55. Type of subspecies.)

6. Bustenari, Prahova. Lord Lilford (p). 4, 4. 6. 64.

(T7. Dodson.)

1. Dobrudscha, Roumania. Purchased (Pruliere). 86. 4. 2. 6.

MICROTUS INCERTUS de Selys-Longchamps.

1811. Arvicola incertus de Selys-Longchamps, Atti della Seconda Riunione

degli Scienziati Italiani, Torino, 1840, p. 225 (near summit of

St. Gothard, Switzerland).
1809. [Arvicola arvalis'] var. fulva Patio, Faune Vert. Suisse, I, p. 236

(near summit of Purka, 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 ot fulva).
1905. [Microtus incertus] Major, Ann. and Mag. Nat. Hist., 7th ser., xv,

p. 511, May, 1905.

691

1907. Microtus arvalis incertiis Mottaz, M6m. Soc. Zool. de France, xx,

p. 32, September, 1907.
1910. Pitymys inccrtus Trouessart, Faune Mamm. d'Europe, p. 188.

Type locality. — 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 occipital 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 bulla?
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 bufTy
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
dupliratns.

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, IG (Rothschild); Campfer, Grisons, 7 (Roths-
child) ; Eugadine, 1.

Austria-Hungary: Meran, Tirol, 1; Paneveggio, Tirol, 1.

BemarJis. — 3[icrotus incertus is readily distinguishable from
M. arvalis by its flattened brain-case, small auditory bullae and

2 Y 2

FIG. 135.
Slicrotus incertus. Nat.

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MICROTUS 693

protrudiug upper incisors. In immature specimens the skull is
strikingly like that of Pitymys suhterraneus, but when fully adult
the interorbital region becomes narrow and ridged, so that the
resemblance to Pitymys is lost.

6, ?. Furka Pass, Switzer- Lord Lilford (p). 1. 11. 7. 13-14.
land.

2. St. Gothard Pass. Purchased. 46. 6. 15. 44, 114.

6. Engadiue (Fatio). E. R. Alston (p). 79. 9. 25. 51.

6 al. Meran, Tirol, Aus- Dr. V. Fric (p). 90. 1. 30. 1.

tria-Hungary.

9 al. Paneveggio, Tirol. IMarquis G. Doria (p). 90. 3. 5. 15.

MICROTUS ASTURIANUS Miller.

1908. Microtus asturianns Miller, Ann. and Mag. Nat. Hist., 8th ser., i,

p. 198, February, 1908.
1910. Microtus asturianus Trouessart, Faune Mamm. d'Europe, p. 181.

Tyi)e locality. — Pajares, Leon, Spain.

Geographical distrihution. — 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 bulke very large. Colour about as
in M. arvalis meridianus.

Colour. — Upper parts buiiy 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 bufly
grey tinged with drab, not conspicuously different from colour of
back ; tail obscurely bicolor, buffy grey below, brownish mixed
with grey above.

Skull. — The skull is larger than in M. arvalis, 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 31. cahrerse and M. liartingi ; this peculiarity is due
chiefiy 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 bullas 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 31. cahrerse, 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 molai-s, 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.

Specimens examined. — Seven, from the following localities in Spain : —
Pajares, Leon, 3 ; La Granja, Segovia, 4.

6, 2 9. Pajares, Leon, Spain. 0. Thomas (p). 8. 2. 9. 205-207.

{N. Gonzalez.) (8. 2. 9. 206. Type of species.)

2 (5,2 juv. al. La Granja, Segovia. Purchased (Escalera). 8.7.30.15-18.

MICROTUS ORCADENSIS Millais.

1904. Microtus orcadcnsis Millais, The Zoologist, 4th ser., viii, p. 244,

July, 1904.

1905. Microtus orcadensis ISIajor, Ann. and Mag. Nat. Hist., 7th ser., xv,

p. 324, March, 1905.
1909. Microtus orcadcnsis Trouessart, Faune Mamm. d'Europe, p. 177.

Tyjye localitij. — 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 31. 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 ochraceous-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.

Skull. — Except for its larger size and relatively shorter, more

095

Fig. 136.
Microtus oreadensis.

widely .spreading zygomatic arche.s, the .skidl differs 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 different 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 3[. arvalis, the distance be-
tween them the same as in the smaller

species. Auditory bullte rather small

relatively to size of skull, their actual

measurements not much greater than

in M. arralis.

Teeth. — In structure the teeth do not differ appreciably from

those of Microtus 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 ti'iangles. 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

Enamel pattern, x 5. ™ales 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.

Fig. 137.
Microtus oreadensis.

SjJccimcns 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, 6 (Edinburgh); South
Ronaldshay, 3.

BemarJis. — 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
Microius sarnhis of Guernsey and the extinct Microtus corneri
Hinton,* from the Late Pleistocene deposits of the Ightham
Fissure, Kent, and Langwith Cave, Derbyshire.

6, 1 al. Rousay Island, Orkneys. W. Cowan (c & p). 4. 8. 22. 1-2.

i. Sandwick, Pomona J. G. Millais (c & p). 4.6.21.1.

Island. {Type of species.)

6 c?. 6 9. Loch Stennes, Pomona N. B. Kinnear (c & p). ) ^ , i,' „' i _a'

i^i*^*^^- je'. ir7.i-3.'

S, 2 9. S. Ronaldshay Island. Edinburgh Museum (p). 7. 11. IC. 1-3.
6, 9. Orkneys. J. G. Millais (c & p). 5. 11. 22. 1-2.

MICROTUS SANDAYENSIS Millais.

(Synonymy under subspecies.)

Geographical distribution. — 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.

BemarJcs. — 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.

697

MiCROTUS SANDAYENSIS SANDAYENSIS Millais.

1905. Microtus orcadensis sandayensis Millais, Mamm. Great Brit, and

Ireland, ii, p. 280,
1908. Microtus sandayensis Miller, Ann. and Mag. Nat. Hist., 8th ser., i,

p. 199, February, 1908.
1910. Microtus sandayensis Trouessart, Paune Mamm. d'Europe, p. 177.

Ti/jie locality. — Sanday Island, North Orkney Islands.

Geograi)liical dintrihution. — -At present known from Sanday
Island only.

Diaynosls. — 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 wliite throughout but rather
strongly darkened by a sprinkling of blackish hairs along median
dorsal surface.

Sicull. — 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,

Microtus sandayensis (a) aucl (a') ;
il. orcadensis (b). Nat. size.

their form in this respect resembling that of If. arvalis rather
than that of M. orcadensis. They are, however, as short and as
widely spreading as in the more neai'ly related species. Inter-
parietal as in M, orcadensis. Auditory bulke slightly smaller
than in orcadensis.

698 RODENTIA

Teeth. — The teeth resemble those of 31. arvalk and M. orca-

densis except in the form of the anterior loop of the first lower

molar. In both arvalis and orcadensis this loop

is deeply indented by two re-entrant angles,

one on each side, the two angles approximately

equal in depth. In M. .HandayeriHis sandayensis

the inner angle remains as in the related

species, but the outer, 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 different from the ordinary type

to j^ermit of certain identification of the animal

by this character alone.

iiG. 139. Measurements. — External measurements of

^ smidaycnsis.'^^''Tl\'' 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).

9. Sanday Island, Orkneys. J. G. ]Millais (c & p). 5. 11. 22. 3.

{Type of species.)
2 J. Sandav. N. B. Kinnear (c & p). 6. 11. 18. 7-8.

4d4?. Sanday. W. R. Ogilvie - Grant 11.11.2.1-8.

(c & p).

MiCROTUS SANDAYEN'SIS WESTR.E Miller.

1908. Microtus sandayensis tvestr.-v Miller, Ann. and Mag. Nat. Hist.,

8th ser., i, p. 199, February, 1908.
1910. Microtus sandayensis wcstnt- Trouessart, Faune I\Iamm. d'Europe,

p. 178.

Type locality. — Westray Island, North Orkney Islands.

Geographical distribution. — At present known from Westray
Island only.

Diarjnosis. — First lower molar with anterior outer re-entrant
angle occasionally as deep as in M. orcadensis ; colour not so pale
as in the Sanday vole, the under parts strongly washed with
yellowish brown.

Colour. — General effect 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-buft', nearly as in M. orcadensis, 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

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700 RODENTIA

SJiulI. — The skull so exactly resembles tliat of Microtus son-
day ensis isandayensis 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 first 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 this 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, 1 1 and 1 1 . 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 Tilag. Nat. Hist., 8th ser., iii

p. 420, May, 1909.

1910. Microtus sarnius Trouessart, Faune !Mamm. d'Europe, p. 178.

Tyj)''' locality. — St. Martins, Guernsey, Channel Islands.

Geofirapliical distribution. — Island of Guernsey.

Diagnosis. — Size about as in 3licrotus orcadensis or slightly
less (hind foot about 18*5 mm., condylobasal length of skuil
about 28 ram.) ; skull narrower than that of orcadensis, its
appearance in old individuals essentially as in the larger races of
M. agrestis ; colour agrestis-\\\ie, 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 I'eddish. 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.

Skull. — 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

MICEOTUS 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. orcadcnsis at the same
stage, though the zygomata are longer and less abruptly
spreading, and the auditory bullse are smaller.

Teeth. — In all respects except for their slightly smaller size
tlie teeth resemble those of Mierotus 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.

Hemarlis. — Misled by the extreme narrowness of the very
aged type skull, I at first supposed this animal to be a member of
the afjrestis group with aberrant dentition. Further material
shows that it is related to the voles of the Orkney Islands and to
the extinct Mierotus corneri of the British mainland, the three
living species and their fossil relative ajaparently belonging to an
older fauna than that now inhabiting Great Britain and the
mainland of Europe.*

4:6,19. St. Martins, Guernsey. 0. Thomas (p). 8. 9. 2. 24-28.
(B. H. Bunting.) (8. 9. 2. 27. Type of species.)

MICROTUS CABRERjE Thomas.

1906. Mierotus cabrene Thomas, Ann. and Mag. Nat. Hist., 7th sor., xvii,

p. 57G, June, 1906.
1910. Mierotus cabrerai Trouessart, Paune Mamm. d'Europe, p. 181.

Type localit]]. — Rascafria, south side of Sierra de Guadarrama,
Province of Madrid, Spain.

Geograpliical 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 tijjs 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. 36, July, 1910.

702

RODENTIA

the underfur a dull, buflfy wood-brown (2 mm.) at tip, the longer
hairs black, those of sides and rump with conspicuous bulFy 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-bufF 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
butfy below, dull brownish above.

Skull. — The skull differs from that
of all other known European species
of Microtus 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
longitvTdinal 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 bulL-e not peculiar, their
size relatively about as in M. ari-alis.

Teeth.— The teeth are strictly of the M.
arvalis type. Terminal loop of both m'^ and
n*! 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 tlie tooth is a nearly straight
loop, almost as wide -as long, and evenly con-
stricted from both sides proximally, while in
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.

Fio. 14".
Mii-rotun cahrene. Nat. size.

MICROTUS 703

as a normal character. Upper incisors short, directed downward
and slightly backward, concealed by nasals when skull is viewed
from above.

Mea.surements. — 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 near Rascafria, in the Sierra
de Guadarrama, Madrid, Spain, and a second specimen without definite
locality.

6. Rascafria, Sierra do 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 cahrenv Cabrera, Asoc. Espafi. Progr. Cien., Congr. Zara-

gosa, 1908, p. 49, June, 1910 (part).
1910. Microtus dentahis Miller, Ann. and Mag. Nat. Hist., 8th ser., vi,

p. 459, November, 1910. Type in Madrid Museum.

Tyjje locality. — Molinicos, Sierra de Segura, Albacete, Spain.

Geograjjhical distribution. — Known only from the Sierra de
Segura.

Diagnosis. — Similar to Microtus cahrerse 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. ; m.^ with a completely closed triangle on outer side.

Colour. — Like that of 31. cahrerse, 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 cahrerse ; underparts light grey
scarcely tinged with buff ; no evident line of demarcation along
sides.

Sliull. — The only known skull is imperfect, lacking the auditory
bullcB and base of brain-case. It is that of an individual slightly
older than the type of Microtus cahrerse, 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. cahrerse, the incisive foramina fully 1^- 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. cahrerse ; interorbital region longer and narrower, but lateral
ridges and median groove not peculiar. Mandible as in M.
cahrerse, 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 Mici'otus cahrerse,

704

EODENTIA

but are immediately distinguishable by the unusually large size
of the molars. Posterior loop of vi'-^ and anterior loop of vi^ rather
short

Fia. 142.
Microtux dentatuK. X ■'>.

tendency to flattening characteristic of cahrerse ; m. 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 (Madrid).

MICROTUS HARTINGI Barrett-Hamilton.

1903. Microtiis {Microtus) hartingi Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., xi, p. 307, March, 1903. Type in British Museum.
1910. Microtus hartingi Trouessart, Paune Mamm. d'Europe, p. 174.

Type locality. — Larissa, Thessaly, Greece.

Geographical distrihution. — Known only from the vicinity of
Larissa.

Diagnosis. — Size large, slightly exceeding that of Microtus
orcadensis and M. cahrerse (hind foot about IS, condylobasal
length of skull about 29 mm.) ; tail relatively short ; colour more
yellowish than in any of the European forms of M. 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 white, the dorsal surface of tail slightly more yellow

705

(about concolor with flanks), tlie belly irregularly clouded by

the slaty bases of the hairs.

ShuU. — 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. cab-

rcrse). 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. arvali.'i, their termination a little

emai'ginate and slightly exceeded

by nasal bi'anches of pre-maxil-

laries. Palatal foramina narrow, as

in 31. arvalis. Auditory bulhe 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 31. 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.

Fi". 144. 3Ieasurements. — External measurements of

Micriiius hartimrji. X 10. type (adult male): head and body, 107; tail,

26 ; hind foot, IS. A second adult male from

the type locality: head and body, 115 ; tail, 27; hind foot, 18 ;

ear from meatus, 10. For cranial measurements see Table, p. 707.

Specimens examined. — Eight, all from the type locality.

Remarks. — Though very difi"erent from the other European
species of 3Iicrotus, this animal is nearly related to 31. guentheri

2 z

Fig. 143.
-Uici-otus hartingi: Nat. size.

706 EODENTIA

Danford and Alston, of Asia Minor. It diflfers merely in its
slightly paler, more yellowish colour, somewhat more robust
skull, and more inflated auditory buUse.

S, 2 al. Larissa, Thessaly, J.. E. Harting (p). 93. 4. 5. 1-3.

Greece. (93. 4. 5. 1. Type of species.)

i al. Larissa, Thessaly. P. H. Mavrogordato 93. 3. 28. 1.

(c & p).
6, 9, 2 juv. al. Thessaly. Dr. F. Loeffler (c & p). 92. 12. 7. 1-4.

MICROTUS ANGULARIS Miller.

1908. Microtiis angularis Miller, Ann. and Mag. Nat. Hist., 8th ser., i,

p. 198, February, 1908. Type in British Museum.
1910. Microtus angularis Trouessart, Fauno Mamm. d'Europe, p. 182.

Type locality. — Transylvania (probably near Hatszeg, Hun-
yad, Hungary). / Vt- ^'f.' v. i<^<

G('ogra])lucal distribution. — Known only from the type locality.

Diar/nosis. — A large member of the 3Iicrotus arvalis group,
about equal to the Grecian 31. 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 3Iicrotus arvalis arvalis in
colour.

SJcull and tccili. — The skull diiTers from that of all other

known European species of 3Iicrutus 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^^^Zj^ 7~^S\, (about 34° instead of about 18 ' to 22 as in

)'^k'\ X^fv N ^^' '^'■^■^^''^j ^- liartinrji and 31. orcadensis)

yC^^^^^X-^ajrac^ — *^u ^^ which the nasals slope downward.

V Auditory bidlse relatively larger than in

Fio. 145. 3/ arvalis, but not so much inflated as

"^"^ Nat.^size. '" '^ ii -^^ hartinr/i. Form of brain-case not

known. Teeth with pattern of enamel
folding as in 3Iicrotus 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.

3Icasuremcnts. — 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 Keyserliiig and Blasius.

ISih Arvicola ratticeps Keyserling and Blasius, Mem. prisentes a I'Acacl.
Imp. dcs Sci. Nat. de St. Petersbourg, iv, livr. 3, p. 333, 1841
(date on title-page of completed volume, 1845). (Welikii-Ustjug,
north-central Russia.)

1841. Arvicola arcnicola de S61ys-Longehamps, Bull, de I'Acad. Eoyale
des Sci. des Arts et Be'lles-Lettres de Bruselles, 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 ratticeps Blasius, Siiugethiere Deutschlands, p. 365.

1887. Microt2is ratticeps Lataste, Ann. Mus. Civ. Stor. Nat. Gcnova,
2nd ser., iv, p. 265, January, 1887.

1899. Arv[icola] (Microtus) ratticeps var. sfimmingi 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 ratticeps, M. ratticeps stimmingi, and M. arcnicola Troues-
sart, Fauna Mamm. d'Europe, p. 179.

Tjipe localitij. — Welikii-Ustjug, on the Dwina River, north-
central 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. — iiize 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 condyloliasal 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.

E.vternal characters. — Essentially as in Microtus agrestis, 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 agrestis, 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-buft'
and frecjuently 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. G4) 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 {sec Winge, Danmarks Fauna,
Pattedyr, p. 78, 1908).

709

blackish brown on nape, elsewhere a mere darkening of noi'mal
colour.

Skull. — The skull attains a greater length than that of
Microtus aijrestis, 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 M. (u/restis, 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 square ; upper surface rather strongly arched laterally,
not flattened as in 31. agrestis ; lateral ridges moderately
developed ; interparietal 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. Interorbital 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 3Iicroius
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 Chionomi/s. Mandible as in 31. agrestis,
but more slender.

Teeth. — Incisors essentially as in 3Iicrotits agrestis, but front
face of upper teeth less obliquely rounded ofl" at outer side.
Size of molars relatively to that of skull about as in Microtus

Fig. 146.
Microtus ratticeps. Nat. size.

710

Microtus ratticeps.
Enamel pattern. :

agrestis or jjerhaps a trifle less. Enamel pattern agreeing with
that of 31. arvalis in the absence of all trace of a postero-internal
loop in Ml'-, as well as in the characters of ?/i^, m., and jUj, common
to nearly all the European members of the sub-genus; there is
everywhere, however, less contrast in size and depth between
the inner and outer angles and triangles. Third ujjjaer 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 Eurojjean 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 ; antei'ior loop so short that it
bears no I'e-entrant angle on outer side, while
that of inner side is reduced to a shallow,
inconspicuous conca^■ity, 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 vi\
While dirtering conspicuously from that of the other European
species of true Microtus 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
reproduced.

Measurements. — Average and extremes of nine adults from
Lappmark, Sweden : hind foot (dry), 19'9 (19'4-21). Adultfrom
Csallukoz-Somorja, Pressburg, Hungary: hind foot, 20 "8. Adult
female from Texel Island, Holland : head and ])ody, 130 ; tail, 62 ;
hind foot, 20 ; ear, 13. For cranial measui-ements .see Table opposite.

Specimens examined. — Twenty-eight, from the following localities : —

Norway. — Gausdal, 4 ; Hjerkin, Kristiansamt, 1 ; Dovre Fjeld, 1
(U.S.N.M.).

Sweden. — Medelpad, W. Norrland, 1 ; Karesuando, Norrbotten, 3 ;
Lappmark, 5 (B.M. aud U.S.N.M. ; near Stockholm, 1 (U.S.N.M.).

Finland.— ^Nluonionniska, Uleaborg, 5 (B.M. and U.S.N.M.).

Germany. — Colpin, Brandenburg, 1 ; near city of Brandenburg, 1 (Berlin
Agric. ; cotype of stimmingi Nehring).

Austria-Hungary. — GsallokOz-Somorja, Pressburg, Hungary, 1.

711

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712 RODENTIA

Holland. — Lisse, near Leiden, 4 (Leiden ; cotj'pes of arcnicola de
S61ys-Longchamps) ; Texel Island, 1.

3 6.

Gausdal, Norway.

Christiania Museum

(E).

93. 3. 1. 11, 13.

6.

Gausdal.

Miller Collection.

7. 7. 7. 153G.

?.

Hjerkin, Kristiansamt.

Miller Collection.

7. 7. 7. 4535.

1.

Medelpad, W. Norrland,
Sweden.

Prof. Sundevall (p).

49. 11. 1. 21.

1.

Torneu, Lappmarlc, Swe-
den.
Lapland.

Prof. Sundevall (p).

45. 10. 25. 10.

1.

Prof. Sundevall (p).

49. 11. 1. 9.

2.

Lapland.

Tomes Collection.

7. 1. 1. 141-142.

1.

Lapland.

Miller Collection.

7. 7. 7. 3250.

1.

Muonionniska, Uleaborg,
Finland.

Stockholm Museum

(E).

90. 8. 1. 7.

6.

Texel Island. Holland.

J. L. Bonhote (c & i

).

8. 10. 26. 3.

lal.

C 0 1 p i n, Brandenburg,
Germany.

C. Gadow (c & p).

81. 10. 7. 1.

2.

C s a 1 1 6 k o ii-S 0 m 0 r j a.

Budapest Museum (e).

94. 3. 1. G4-5

Pressburg, Hungary.

Sub-Genus CHIONOMYS Miller.

1857. Paludicola Blasius, Saugethiere Deutschlands, p. 334 (part). Not of

Wagler, 1830.
1867. Praticola Fatio, Las Campagnols du Bassin du L6man, p. 34 (part).

Not of Swainson, 1837.
1896. Microtus MiUer, North American Fauna, No. 12, p. 62, July 23, 1896

(part).
1908. CJiionomys Miller, Ann. and Mag. Nat. Hist., Sth ser., i, p. 97,

January, 1908.

Type species. — Arvicola niralis IMartins.

GeogyapMcal distrihution. — Mediterranean 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 Pitijmijs ; 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 leas
well-defined.

lie marks. — The sub genus Chiononujs 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 lowdands 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.

MICROTOS 713

KEY TO THE EUROPEAN ]\IEMBERS OP 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. Ichrunii, p. 718.

Back clear light grey ; auditory bulloe small and
Uattened (Neighbourhood of Nimes, Gard,

France) ^f. I. lehrunii, p. 719.

Back with slight brownish tinge; auditory bullae
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) M. ulinus, 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) 3f. n. nivalis, p. 716.

Anterior loop of first lower molar with postero-
external salient angle usually broad and
rounded, sometimes obsolete (Pyrenees) M. n. aquitanius, p. 717.

MICROTUS NIVALIS Martins.
(Synonymy under subspecies.)

Geograpltical distribution. — Pyrenees, Alps (except extreme
south-western portion : Basses- Alpes and Var), Apennines and
Tirol.

Diagnosis. — Size rather large (head and body about 130 rum.
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 fos.sa
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 with 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. (Kjrestis. Ear rather 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 M. 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 annulations, 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-gi'ey, 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 Ijrown above though never sharply
bicolor.

Shdl. — The skull is larger than that of Microtus agrestis
agrestis, its general form broader and more depressed, the dorsal
surface much less marked by ridges for muscle attachment, and
the auditory bulhe more developed. Dorsal profile 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
(aViout 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 ^'iewed 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 conspicuous ; 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 trajisverse diameter). Depth of occiput
when viewed froui behind barely more than half height, the
* Somewhat exaggerated in fig. 148.

Fig. 148.
Microtus nivalis.

MICR0TU9 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 2i times in median length ; auditory bulhe 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 bi-eadth of brain-case much less than in Mlcrotus
agrestis ; 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 dejith 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 ; mesopterygoid
space nearly parallel sided, its anterior border rounded, its width
iu front scarcely equal to distance from margin of fossa to
alveolus of 711^ ; 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 M. 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 sj^ecies of true Ilicrotus, 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 rii}, m- and m,^ shows no peculiarities worthy of special
note ; in" without postero-internal loop. Third lower molar with
re-entrant angles on outer 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 with 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 31. agrestis when
second inner triangle has been cut off ; 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 oiiter 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.

Remarks. — 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 clilfs
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 ilartins, Ecvue Zoologique, p. 331 (Faulhoni, Bern,

Switzerland).

1843. Hyimdseus aljyinus Wagner, Schreber's Siiugthiere, Supp]., iii, p. 57G

(Andermatt, Uri, Switzerland).
1845. Arvicola nivicola Schinz, Syuops. Mamm., ii, p. 236 (Highest Swiss

Alps; probably near Andermatt).
1853. Hypucheus pctrophilus Wagner, Miinch. Gelehrt. Anzeigen, No. 38,

p. 307, March 28, 1853 (Oberstdorf, near Sonthofen, Allgau,

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 l\Iag. Nat. Hist., Sth ser.,

I, p. 99, January, 1908.
1910. Microtus (Chionomys) nivalis Troucssart, Fanne ]\Iamm. d'Europe,

p. 183.

Type localitjj. — Faulhorn, Bern, Switzerland.

Geofjraplilral distribution. — Alps (except south-eastern portion),
Tirol and northern 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. G15-618, 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
shai-jjly pointed, and rarely, if ever, except in immature
individuals, obsolete.

Measurements. — External measurements of adult male from
Chamonix, Hauto-8avoie, 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, 1 20 ; tail, 63 ; liind foot, 20. For cranial measurements
see Table, p. 720.

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

Prance: Chamonix, Haute-Savoie, 2 (U.S.N.M.).

SwiTZEBLAND : 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)
Siintis, Appenzell, 12 (U.S.N.M.) ; Meglis Alp, Appenzell, 2 (U.S.N.M.)
Hochmiittli, Murgsee region, St. Gallen, 2 (U.S.N.M.) ; Campfer, Grisons, 1
(Rothschild) ; no exact locality, 2.

Gebmany : Oberstdorf, near Sonthofen, Allgau, Bavaria, 2.

Italy : Valasco, Valle del Gesso, Cmieo, . 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. 14:3-144.

7 6, ?. Furka Pass, Uri. {E. H. O. Thomas (p). 2. 8. 4. 43-47.

ZolUkofer.) 4. 4. 5. 54-56.

1. St. Gothard, Uri. Baron E. de Selys- 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. (Mosclder.) 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.

1903. Microtns nivalis aquitanius Jliller, Ann. and Mag. Nat. Hist.,
8th ser., i, p. 99, January, 1908. Type in British Museum.

1910. Microtns [Chioywinys) nivalis a<2uitanius Trouessart, Fauiie Mamm.
d'Europe, p. 184.

Type locality. — Near I'Hospitalet, Ariege, France. Altitude
about 4800 feet.

Geixjrapliical distribution. — Pyrenees ; at pi'esent known fi-om
the French side only.

718 RODENTIA

Didgnrms. — Similar to 3Iicrotus 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 Microtis nivalis nivalis (a) and
M. nivalis aquilanius (Ji). 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, 1 7 • 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, Pyrenies-Orientales, 2 ; I'Hospitalet,
~ ■ — — . . - Pic-du-j\Iidi, Hautes-Pyren6es, 6.

G. S. Miller (c). 8. 8. 4. 240-241.

G. S. Miller (c). 8. 8. 4. 237-239.
(8. 8. 4. 238. Type of subspecies.)
O. Thomas (p). 8. 9. 1. 64-69.

Ariege, 5 ; Bareges, Hautcs-Pyr^nees, 1 ;

2 ?. Port6, Pyrenees - Orientales,
France.
L'Hospitalet, Ariege.

3(5.
3 6, 3

Pic-du-Midi, Hautes-Pyrenees.
{A. Robert.)

MICROTUS LEBRUNII Crespon.
(Synonymy under subspecies.)

Geographical distrihution. — Central and south-eastern France
from Puy-de-D6me to the Departments of Gard, Var, and Basses-
Alpes.

Diagnosis. — Like Microtits 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 differ
tangibly from that of M. nivalis.

Teeth. — As compared with those of 31. nivalis of the same
age, the teeth differ in their slightly smaller size, but I can find

MIOROTUS

719

no constant cliai-acter in the enamel pattern by which they may
be distinguished. In the specimens examined the terminal looji
of wi^ is invariably short and broad, not showing any tendency to
the elongation occasionally occurring in M. nivalis.

Bemarhs. — Microtus lehrunii is represented by two geographic
races, one of which is remarkable, in the sub-genus Chionomys, for
its occurrence at a distance from mountains and in a region of
unusually high summer temperature.

MiCKOTUS LEBRUNII LEBRUNII CrespOH.

1844. A\rvicold] lebrimii Crespon, Paune M^ridionale, i, p. 77.

1857. Arvicola nivalis b. Arvicola Iciicurus Blasius, Siiugethiere Dcutsch-

lands, p. 359 (part).
1908. Microtus Icbninii lebnmii IMiller, Ann. and Mag. Nat. Hist., 8th ser.,

I, p. 101, January, 1908.
1910. Microtus {Chionomys) Icbruni TrouGSsart, Faunc Mamm. d'Europe,

p. 184.

Tyjje locality. — Neighbourhood of Nimes, Gard, France.
Altitude, 180 m.

Geogra]}liical distrihution. — Known with certainty from the
type locality only, but probably extending through the Cevennes
and the mountains of Auvergne.

Diagnosis. — ^Back clear, very pale smoke-grey, without evident
wash of wood-brown ; skull with auditory buUte small and
flattened.

Colour. — Upper parts a \ery 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 bufiy 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 bullae 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 Nimes {B.M.,
Mottaz and Nimes) ; no exact locality, 2.

Remarls. — 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 JNIuseum collected in Auvergne (probably
near Clermont-Ferrand) by Lecoq, and presented in 1854, is

720

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MICROTUS

721

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722

ROD£NTIA

probably referable to Microtus lehrunii lebrunii, though in its
present faded and dust-stained condition exact identification is
impossiT>le. The skull is so impei'fect that it serves only to show
that the animal was not adult.

3 S. Nimes, Gard, France. O. Thomas (p). 8. 8. 10. 101-103.

(C Moltaz.) {TopotijiJcs.)

2. S. France. Tomes Collection. 7. 1. 1. 140-147.

Microtus lebrunii leucurus Gerbe.

1852. Arvicola Icuairus Gerbe, Revue Zoologique, 2nd ser., iv, p. 260,

June, 1852.
1857. Arvicola nivalis b. Arvicola leucurus Blasius, Siiugethiere Deutsch-

lands, p. 359 (part).
1908. Microtus lebrunii leiicurus Miller, Ann. and Mag. Nat. Hist., Sth ser.,

I, p. 102, January, 1908.
1910. Microtus (Chionomijs) lebruni leucurus Trouessart, Faune Mamm.

d'Europe, p. 184.

Type localltij. — Barcelonnette, Basses-Alpes, France. Altitude
about 1300 mm.

Geographical distribution. — South-western Alps (Departments
of Basses-Alpes and Var).

Diagnosis. — Ujjper parts pale smoke-grey, slightly but evi-
dently washed with wood-brown ; skull with auditory bullje large
and well inflated, essentially as in M. nivalis.

Colour. — The colour resembles that of M. lehrunii lehrunii,
except that the back is sufficiently sufiused with bufTy or pale
wood-brown to destroy the clear grey, almost hoary appearance
characteristic of the typical, lowland race. This suffusion is
usually more noticeable on rump than on sides, so much so that
in one specimen (No. 7. 1. 1. 14-5, probably a imratype) the
rump is a clear dull buff.

Skull. — The skull is distinguishable from that of M. lehrunii
lehrunii by its large, strongly inflated, normally developed auditory
bullae. As in M. nivalis the lower edge of the bulla extends
decidedly beyond line of cutting edge of molars.

Measurements. — 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.j\I.).

^, ?. Barcelonnette, Basses-Alpes, 0. 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.

AEVICOLA

723

MICROTUS ULPIUS Miller.

1908. Microhis ulpius Miller, Ann. and Mag. Nat. Hist., Stli ser., i, p. 100,

January, 1908. Type in British Museum.
1910. Microtus (Chionomys) ulpius Trouessart, Faune Mamm. d'Europe,

p. 184.

Tii2)e locality. — Hatszeg, Hunyad, Austria-Hungary. Altitude,
2000 'feet.

Geographical distribution. — 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 Microius 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.

Sladl 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 wjj usually similar to that of
M. n. aquitanius.

Measurements. — External measurements 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.

6 (J, 7 9. Hatszeg, Hunyad, Transyl- C. G. Danford (c). 3. 2. 2. 45-49.
vauia, Austria-Hungary. 3. 11. 8. 35-42.

(3. 2. 2. 48. Type of species.)

Genus ARVICOLA Lacepede.

1799. Arvicola Lacepede, Tableau des divisions, sous divisions, ordres et

genres des mammiferes, p. 10 (amphibius).
1836. Hcmiotomys de Selys-Longchamps, Essai monographique sur les

Campagnols des environs de Liege, p. 7 (sub-genus) part.
1857. Paludicola Blasius, Sdugethiere Deutschlands, p. 333 (sub-genus)

part. Not Paludicola Wagler, 1830.
1867. Ochetomys, Fitzinger, Sitzungsb. Math. Naturwiss. CI. K. Akad.
Wissensch., Wien, lvi, p. 47 (based primarily on the water-rats of

Europe).
1807. Praticola Fatio, Les Campagnols du Bassin du Leman, p. 36

(sub-genus) part. Not Praticola Swainson, 1837.
1888. Arvicola Lataste, Le Naturaliste, ii, p. 349 (sub-genus).

3 A 2

724

RODENTIA

1896. Arvicola Miller, North Amer. Fauna, no. 12, p. GG, July 23, 1896

(sub-genus).
1908. Arvicola Miller, Ann. and :Mag. Nat. Hist. 8th ser., i., p. 195,

February, 1908 (genus).

Ty2^e species. — Mus ampMbius Linnpeus.

Oeographical distribution. — Palsearctic region from Great
Britain eastward into Asia, and from the Mediterranean coast
north to northern Scotland and the shores of the Arctic Ocean.

Characters. — Like Microtus, but plantar tubercles only five
(a very rudimentary sixth rarely present) ; third upper molar as
in Chionomys ; external form modified for aquatic or subterranean
life.

Bemarks. — Though not sharply diflerentiated from Microtus,
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 aquatic in their mode of life,
others fossorial, while in two (^A. terrestris and A. scherman
scherman) the habits appear to vary locally or with season.

KEY TO THE EUROPEAN FORMS OP ARVICOLA.

Size large, hind foot more than .30mm.,condylobasal
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 m^
and nu forming evident protuberances on lower
surface of mandible in old individuals ; habits
strictly aquatic {avipliihius group).
Nasals at widest region conspicuously narrower

than rostrum (Great Britain) A. ampliibius,j). 725.

Condylobasal length of skull usually 42 to
44 mm. ; colour dark, but melanism infre-
quent (England) A. a. avijjhihius, -p. 130.

Condylobasal length of skull usually 40 to
42 mm. ; colourvery dark, melanism frequent

(Scotland) A. a. reta, p. 732.

Nasals at widest region nearly as wide as rostrum

(Spain and southern France) ^4. sapidus, p. 732.

General colour rather light, the sides and face
clear yellowish brown (Spain, except As-

turias ; lowlands of southern France) A. s. sapidus, p. 733.

General colour dark (as in amphibius), the sides
and face strongly " lined " with black

(Pj'reuecs and Asturias) A. s. tenebricus, 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
?», and w„ never forming protuberances on lower
surface of mandible ; habits aquatic or terres-
trial (terrestris group).

ARVICOLA 725

Interparietal tending to be somewhat ligulate in
outline ; skull representing the extreme of

fossorial adaptation A. scherman, p. 744.

Colour dark brown, the tail usually blackish
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 ?»( short and wide ; auditory

buUiTB usually somewhat flattened (Alps

and their immediate neighbourhood) A. s. cxitiix, p. 746.

Anterior loop of ■«(., long and narrow; auditory
bulltB evenly inflated (Pyrenees and their

immediate neighbourhood) A.s. inonlkola, 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. tcrrestris, 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. 7Husignani,]).74.i.

General colour dark brown, the back with
noticeable clouding of black.
Underparts slaty, washed with yellowish
brown (Italian Switzerland and north-
ern Italy) A. italiciis, p. 740.

Underparts slaty, washed with whitish

(Bosnia and Koumania) A. illyricus, p. 741.

ARVICOLA AMPHIBIUS Linnaeus.

(Synonymy under subspecies.)

Geoijraph'tod distribution. — Great Britain.

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 m^ and m., forming evident protuberances on
lower surface of mandible in old individuals; habits strictly
aquatic, never mole-like.

E.xtcrnal charnciers. — General 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 surface 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. jVEuzzle 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 about 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
iri-egularly 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, well 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 forwai'd it
extends about to shoulders or slightly beyond ; annulations
evident, though somewhat irregular, about 15 to the centimeter
at middle of tail ; hairs numei'ous, about 4 to 5 mm. in length,
nearly concealing the annulations and forming a slight pencil.
Mammaj : jj 2 — 2, «' 2 — 2 = 8.

Colour. — Upper parts ranging from broccoli-brown to mai-s-
brown or darker, in most specimens rather heavily overlaid with
black, though the latter is usually not in excess of the gi-ound-
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 odii-aceous-bufF or liglit law-umber,
though not sutliciently to produce any marked contrast with
surrounding parts ; chest and belly a varying mixture of
ochraceous-butf 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.

SJniJl. — The skull of Arvicola amphihius is large and massive,
becoming conspicuously ridged aiid angular in fully adult

FIG. 150.
Arvicola ami)hibius. Nat. size.

individuals. In general form and proportions it does not differ
notably from the skull of Microtus agrcstis, except that the brain-
case tends to be shorter and wider, the dorsal pi-ofile is mo*-e
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
visilile when skull is viewed from above. Brain-case somewhat
longer than wide, the well-developed postorbital processes
st[uaring its anterior margin and imparting the general outline
of a parallelogram ; lateral ridges strongly developed in adults
and marking oif a conspicuous flat median surface widest at

728 KODENTIA

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
23oint 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 moi'e so than in Microtus
aijrcstis, the entii'e 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 ; paroccipital processes large,
widely spreading, their extremities not applied to bulhe, 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 bullse
rather large (about 9 mm. in antero-posterior diameter), but not
highly inflated, their ventral border not extending to level of
cutting surface of molars. Interorlntal region deeply constricted
posteriorly, where in old individuals the width is less than half
that in lachrymal region ; ridges eai'ly 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 Microti! h aijrcstis, but .somewhat more
slender anteriorly ; nasals distinctly smaller relatively to rostrum
than in 31. agrcstis, 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 foivimina relatively shorter
than in Microtus (ujrcstis, their posterior border f.'illing short of
alveolar level by 1 mm. or more, their anterior extremity lying
decidedly nearer maxillo-jiremaxillary suture than to posterior
surface of incisor. Bony palate with no special peculiarities as
compared with that of Microtus agrcstis ; 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 ai/restis, 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 31. 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 711.-, 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
ti'ansverse 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 ; ni'^ 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 ; m^ 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 ;
nin, with four closed, sub-equal triangles (the antero-internal and

Arvicola amphibiiis.
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 ;
Wg 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.

Bemarls. — So far as is at present known Arvicola amphihius
is confined to Great Britain. Among the continental species it
finds its nearest relative in A. sajjidus of Spain and southern
France. Two local races appear to be represented among the
specimens examined, though their status is not well defined.

Arvicola amphibius amphihius Linnauis.

1758. [Mus] amphibius Linnaeus, Syst. Nat., i, 10th ed., p. 61 (England :
based on the Mus major aquaticus of Ray, Quadr., p. 217).

1816. 7 Arvicola aquatica Leaoh, Syst. Catal. Spec. Indig. Mamm. and

Birds, Brit. Mus., p. 7 (nomen nudum : " Water Campagnol ").

1817. Lemmus aquaticus F. Cuvier, Diet, des Sci. Nat., vi, p. 306, part

(substitute for ampUihius).
1812. Arvicola americana Gray, Ann. and Mag. Nat. Hist., x, p. 226 (half-
grown individuals supposed to have been taken in South America).

Co-types in British Museum.
1815. ? [Arvicola amphibius'] sub-var. nigricans de S^lys-Longchamps, Atti

della sesta Riunione degli Scieuziati Italiani, Milano, 1844, p. 322

(nomen nudum).
1857. Arvicola amphibius a. Blasius, Saugethiere Deutschlands, p. 344

(part).
1910. Arvicola amphibius amphibius Miller, Proc. Biol. Soc. Washington,

sxiii, p. 19, March ^i, 1910.
1910. Arvicola amphibius Trouessart, Faune Mamm. d'Europe, p. is.

Type locality. — England.

Gcograjjliical distribution. — England and southern Scotland ;
exact northern limits of range not known.

Diagno-iis. — 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 infrenuent.

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, Earlsfield, 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 cxa mined. Sexcnty -two, from the following localities : —
ScoTL.\^'D : Blackwood, Lanarkshire, 3 (Edinburgh) ; Stockbriggs,
Lanarkshire, 1 ; Bargsly, Kirkcudbright, 1 ; Stirling, 1.

731

England : Alnwick, Northumberland, 1 ; Wylara, Northumberland, 1 ;
Hull, Yorkshire, 4; Cheadlo, Staffordshire, 1; Diss, Norfolk, 1; Ashton,
Oundle, Northamptonshire, 2 ; Watcrbeach, Cambridgeshire, 1 ; Grafton-
bury, Hereford, 7; Groat Grimsby, Lincolnshire, 2; Kingsbury, ^Middlesex,
1 ; Hampton, Middlesex, 1 ; Chertsey, Surrey, 1 ; Garrett Park Lake,
Earlsfield, Surrey, 6 ; Shalford, Surrey, 9 ; Surbiton, Surrey, 1 ; New
Forest, Hampshire, 9 ; Eversloy, Hampshire, 3 ; Aldershot, Hampshire, 3 ;
Clcvedou, Somerset, 2; Blandford, Dorsetshire, 2; Gullompton, Devon-
sbiro, 2; Northlow, Dovoushiro, 1; Horsham, Susses, 1; Minster, I. of
Thanot, Kent, 1 ; Yalding, Kent, 1 ; no exact locality (supposed to bo from
South America), 2 (co-types of americana Gray).

9. Stockbriggs, L a n a r k -
shiro, Scotland.

(Taylor.)
9. Stirling. (Broimi.)
6. Alnwick, Northumber-
land, England.
6. Wylam, Northumber-
land.
2 d, 2 9. Hull, Yorkshire. (Capt.
Hume.)
Oundlo, Northampton-
shire. (Cox.)
Graftonbury, Hereford-
shire.
Great Grimsby, Lincoln-
shire.
Kingsbury, Middlesex.

(J. E. Harting.)
Hampton, Middlesex.

(IF. Dodson.)
Chertsey, Surrey.

E. R. Alston (p). 79. 9, 25. 46.

2 i.

3<?.
i, 9.

9.

6. Earlsfield, Surrey.

3 c?, 6 9. Shalford, Surrey.

1 (albino) Surbiton, Surrey.

S, 9 juv., Aldershot, Hampshire.

iaX.

4 (?, 5 9. New Forest, Hampshire.

E. R. Alston (r).
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 Haigh
(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 (5, 1 9. Eversley, Hampshire. Miller Collection.

6. Clevedon, Somerset.
i. Blandford, Dorset.

6. Gullompton, Devonshire.

R. I. Pocock (c & p).

J. C. Mansel Pley-
dell (c & p).

G. C. Shortridge
(c & p).

Purchased (War-
wick). (Co-types
Gray).

79. 9. 25. 49.
11. 1. 3. 406.

11. 1. 3. 294.

11. 1. 2. 51-57.

6. 8. 18. 1-2.

11. 1. 3. 297-299.

11. 1.3.295-296.

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,
2935-2936.

II. 1. 3. 291.
11. 1. 3. 292.

11. 1. 3. 290.

42. 4. 12. 6-7.
of A. americana

732

Ahvicola amphibius reta Miller.

1832. Arvicola afer Macgillivray, Mem. Wernerian Soc. Nat. Hist, vi,
p. 429. Not Hypudmus terrestris fi atcr Billberg, 1827.

1910. Arvicola amphihius reta Miller, Proc. Biol. Soc. Washington, xxiii,
p. 19, March 23, 1910 (substitute for atcr).

1910. Arvicola amphihius reta Trouessart, Faune Mamm. d'Europe, p. x.

Type locality. — Aberdeen, Scotland.

GeoffrapMcal distribution. — Scotland, except southern portion ;
limits of distribution not known.

Diagnosis. — Size less than in A. anqtJiihiiis am^thibins (hind
foot usually 30 to 32 mm., condylobasal length of skull usually
less than 42 inm.) ; normal colour darker than in the typical
race, the black usually in excess on upper parts ; melanism
fre(|uent.

Measurements. — Adult male from Glonfeshire 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 ; Glenfcshire
Forest, Inverness, 1 ; Cortachy, Forfar, 3 (B.M. and Wilson) ; Windygates,
Fife, 5 (B.M. and Edinburgh).

i. Llanbryde, Elginshire. W. Taylor (c & r). 11. 1. 3. 407.

Scotland.

6. Monar Forest, Eoss- Maj.-Gen. C. E. Luard 92. 8. 31. 1.

shire. (c & p).

6. Kingussie, Inverness- G A. Cooper (c. & r). 5. 5. 12. 1.
shire.

? al. Crieff, Perthshire. W. R. Oglivic-Grant 88. 5. 30. 2.

(c & p).

<J, 9. Windygates, Fife- N. B. Kinnear (c & p). 6.11.18.5-6.

shire.

6, ?. 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 ampliihius 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.

Memarlcs. — ^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 amphihius 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

FIQ. 152.
Arvicola sapidus. Nat. size.

take place. In the Pyrenees and their immediate neighbourhood
a form of sapidus 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 INIag. Nat. Hist., 8th ser.,i, p. 195,

February, 1908. Typo in British Museum.
1910. Arvicola sapidus scqndus Miller, Proc. Biol. Soc. Washington, xxiii,

p. 20, March 23, 1910.
1910. Arvicola sapidus Trouessart, Fauna Mamm. d'Europe, p. s.

Type locality. — Santo Domingo de Silos, Burgos, Spain.

Geographical distribution. — Essentially the entire Iberian
Peninsula.

Diagnosis. — General colour not so dark as in A. ampliihius
amphibius, the sides and face a clear yellowish brown without
noticeable sprinkling of blackish hairs.

Colour. — Upper parts a yellowish brown vai'ying 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 measvu-ements of type (adult female) :
head and body, 187 ; tail, 123 ; hind foot, 34 ; ear, 18. Average

734 KODENTIA

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). Average 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 cxammccl. — One hundred and twenty-four, from the
following localities : —

Spain : L6rida, 1.3 ; Panticosa, Huesca, 32 ; Jaca, Huesca, 6 ; Arre-
chavaleta, Vitoria, 1; Pajdres, Leon, 6; La Coruna, 2 (not typical,
approaching tcncbricus) ; Silos, Burgos, 20 ; vicinity of IMadrid, 1
(U.S.N.M.) ; Villalba, JIadrid, 1 ; Barracas, CasteUon, 12 ; Silla, Valencia, 1 ;
Vcnta del Baul, Granada, 1 ; Seville, 7 ; near Tarifa, Cadiz, 1 ; Audalucia, 5.

Feance : Foret de Bouconne, Gers, 1 (not typical) ; Leguevin, Haute-
Garonne, 6 (not typical, approaching tencbricus) ; St. Gilles, Gard, 1 (not
typical, approaching tenehriciis) ; Valescure, Var, 5 (not typical) ; Var, no
exact locality, 2 (not typical).

RemarTis. — Throughout Spain the characters of Arvicola sajndus
scqjidus are very constant ; only in the extreme north-west does
there appear to be any tendency toward the darker A. s. tene-
hriciis. The French specimens from the neighbourhood of
Toulouse and from the marshes near the mouth of the Rhone
agree with true sapklHn in the genei'al light yellowish hue of the
back and sides, though showing a distinct approach to tenehriciis
in the noticeable sprinkling of black hairs on cheeks.

4 (?, 3 <5 juv. L6rida, Spain. [N. Gon- 0. Thomas (r). 8. 2. 9. 192-201.
2 ?, 9 juv. zalez.)

8(5,4 9. Panticosa, Huesca, O. Thomas (p). 8.2.9.172-183.

Spain. {N. Gonzalez.)

3 i, 9. Jaca, Huesca. {N. Gon- 0. Thomas (p). 8. 2. 9. 184-187.

zalez.)

6. Arrechavalcta, Vitoria. O. Thomas (p). 8. 2. 9. 203.

{N. Gonzalez.)

2 9, 2 9. Pajares, Leon. (N. Gon- O. Thomas (p). 8. 2. 9. 188-191.

zalez.)

2 <5, 5 9. Silos, Burgos. G. S. Miller (c). 8. 8. 4. 113-119.

(8. 8. 4. 115. TyxJe of species.)

9. Burgos. S. & N. Gonzalez 8. 7. 7. 25.

(c).

7 <5, 5 9. Barracas, Castellon. 0. Thomas (p). 8. 2. 9. lGO-171.
(N. Gonzalez.)

i. Silla, Valencia. (N. Gon- O. Thomas (p). 8. 2. 9. 202.

zalez.)

4 9, 2 9. Seville. (Dr. Ruiz.) Lord Lilford (p). 95. 3. 3. 23-28.

Juv. al. Seville. Dr. V. L. Scoane 94. 3. 19. 3.

(c & p).

6. Villalba, Madrid. O. Thomas (p). 8. 2. 9. 204.

{N. Gonzalez.)

1. Tarifa, Cadiz. Col. L. H. Irby 94. 4. 27. 1.

(c & p).

3 al, 2. Andalucia. Lord Lilford (p). 94. 9. 2G. 1-5.

S. PoretdeBouconne.Gers. O. Thomas (p). 8. 9. 1. 7G.

{A. Robert.)

ARVICOLA 735

6. Leguevin, Haute- 0. Thomas (p). 8.9.1.77.

Garonne. (A. Robert.)

9. St. Gilles, Gard, France. G. S. Miller (c). 8. 8. 4. 2G1.

a J, juv. Valescure, Var, France. G. S. Miller (c). 8. 8. 4. 257-260.

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 S61ys-Lougchanips.

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, March 23, 1910.

1910. Arvicola sapidus tenebricus Trouessart, Fauna Mamm. d'Europc,

p. X.

Type locality. — Vicinity of Biarritz, Basses-Pyrenees, France.

Geographical distribution. — Pyrenees and Atlantic coast I'egion
of south-western France, north to the Garonne ; northern limit
of range not known ; two specimens from La Coruna, 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. amphibius ampliihius), the sides and face
conspicuously sprinkled with black-tipped hairs.

Colour. — Upper pai'ts 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.

Measurements. — 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, Pyrenees-Orientales, 2 ; I'Hospitalet, Ariege, 2 ; Luchon,
Haute-Garonne, 2 ; Montrejeau, Haute-Garonne, 5 (U.S.N. Si.) ; Biarritz,
Basses-Pyr6u6es, 7 ; Cadillac, Giroude, G (U.S.N.M.) ; Barsac, Giroude, 1
(Lataste).

Spain : — Coruua, Galicia, 2.

2 6. Port(5, Pyrenees - Orien- G. S. Miller (c). 8. 8. 4. 254-255.
tales, Prance.
6. L'Hospitalet, Ariege. G. S. Miller (c). 8. 8. 4. 256.

6 juv. L'Hospitalet, Ariege. O. Thomas (p). 8. 9. 1. 71.

(A. Robert).
5 (5, 2 9. Biarritz, Basses-Pyren6es. J. P. Davison (p). 0. 1. 21. 5-6.

6. 6. 4. 10-14.
(6. 1. 21. 5. Type of subspecies.)
2. Coruua, Galicia, Spain. Dr. V. L. Seoane 93. 12. 15. 1.

(p). 95. 6. 25. 1.

736

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RODENTIA

ARVICOLA TERRESTRIS Linnaeus.

1758. [Mus] terrcstris Liniifeus, Syst. Nat. i, 10th cd., p. 61 (Sweden).

1771. [Bins'] paludosus Linnaeus, Mantissa Plantarum, pt. 2, p. 522
(Sweden).

1827. IHypudxus terrestris] j3 atcr Billberg, Synopsis Faunae Scandinaviaa,
p. 4 (Gottland, Sweden).

1827. [Hypudxus imludosiis] /3 littoralis Billberg, Synopsis Faunae Scandi-
uaviae, p. 5 (Smaland, Sweden).

1827. [Hyjjudii'us paludosus] y aqtiaticus Billberg, Synopsis Faunae Scandi-
navite, p. 5 (South Sweden).

1858. Arvicola amphibitis a. Blasius, Saugethiere Deutschlands, p. 344
(part).

1910. Arvicola terrestris Miller, Proc. Biol. Soc, Washington, xxiii, p. 20,

March 23, 1910.
1910. Arvicola terrestris Trouessart, Faune Mamm. d'Europc, p. s.

Type locality. — Upsala, Sweden.

GeogrnpMcal distribution. — Scandinavian Peninsula, eastward
into Finland ; limits of range not known.

Diagnosis. — Size less than in Arvicola arnjjliihius, but genei'al
proportions not jDeculiar (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. aviphlbins, 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-cjuadrate in outline ; teeth rather heavy, but roots of vi^
and m.^ never forming protuberances on lower surface of mandible ;
habits both aquatic and mole-like.

External characters. — Aside from the somewhat less robust
general form correlated with the animal's smaller size there
appear to be no tangible external characters to distinguisli
Arvicola terrestris from A. amphihius.

Colour. — Upper parts varying from broccoli-brown to mars-
brown, heavily overlaid with black, the latter usually thougli
not always in excess and producing a general effect not far from
seal-brown, especially along median dorsal region ; underparts a
distinctly I'usty 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 visually producing a decided contrast
with surrounding parts ; feet varying from hair-brown to a decii
blackish brown ; tail blackish throughout, the under surface often
sprinkled with greyish hairs.

SJiull. — The skull is not so large as that of Arvicola amphibius
or A. sa2jidus. In form it differs from the skulls of the strictly
aquatic members of the genus in a decided tendency to assume a

ARVICOLA

739

fossorial structure. This is shown by the more pronounced
forward projection of tlio upper incisors, and more particularly
by the shortening and widening oi the brain-case in general and
the slightly greater obliijuity of the occipital region. Flattened

Fig. 153.
Arcicola terrestris. Nat. size.

area between lateral ridges on brain-case tending to assume a
diamond-shaped 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 Arvicola aniphibius.

Measurements. — Adult female from Holme, Mandal, Norway :
head and body, 167 ; tail, 99 ; 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 Jemtland, 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 : —
Norway : — Vefsen, Nordland, 1 ; Elverum, Hedemarken, 1 (U.S.N.M.) ;

Christiania, 1 (U.S.N.M.); Askor, Christiania, 1 (U.S.N.M.); Holme,

Mandal, 16.

Sweden :— Jemtland, 7; Upsala, 2 (B.M. and U.S.N.M.); Upland, 2

3 B 2

740

EODENTIA

(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.

Rewarlcs. — Arvicola terresfris is readily distinguishable from
A. ampluhius 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, Norclland, Nor- E. G. B. Meade 5. 7. 1. 3.

way. Waldo (c & p).

6 i, 9, Holme, Mandal. R. J. Cuninghame 8. 8. 9. 19-29.

(c & p).

6. Upsala, Sweden. Lord Lilford (p). 0. 5. 15. 1.
(E. Kolthoff.)

2. Near Stockholm. ' Prof. Snndevall (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 aniphibius var. italica Savi, Nuovo Giorn. dc' Letterati,

Pisa, XXXVII, No. 102, p. 202 (p. 5 of separate), February 1839 (for

date see de S61ys-Longchamps, Micrommalogie, p. 93). Vicinity

of Pisa, Italy.
1839. Arvicola pertinax Savi, Nuovo Giorn. de' Letterati, Pisa, xxxvii.

No. 102, p. 203 (p. 6 of separate), February, 1839 (MS. synonym

of italica).
1845. ? [Arvicola amphibius var. minor De Selys-Longchamps, Atti della

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, Fauno Mamm. d'Europe, p. x.

Tij2)e locality. — Vicinity of Pisa, Italy.

Geoc/rapMcal distrihntlon. — Italian Switzerland and northern
Italy, south at least to the vicinity of Pisa. Details of distribu-
tion not known.

Dia<jnosis. — Similar to Arvicola terrestris, but teeth not so
heavy and colour not so dark, the underparts washed with
yellowish brown instead of I'usty, 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, bufTy, 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 nnd teetli. — The skull resembles that of Arvicola terrestns
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 bulL-e slightly larger than in the Scandinavian species,
but not peculiar in form. Teeth as in A. ierrestris, but slightly
larger.

Measurements. — Average and extremes of five adults from
Lugano, Ticino, Switzerland: head and body, 180 -G (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 6,2 9. Locarno, Ticino, Swit- O. Thomas (c & p). 2. 7. 1. 2-5.
zerland.

9, 2juv. al. Modena, Italy. Marquis G. Doria (p). 90.3.5.1-3.

1. Italy. Purchased (Brandt). 4G. 1. 9. 20.

ARVICOLA ILLYRICUS Barrett-Hamilton.

1899. Microtus musignani illyricus Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7ch ser., in, p. 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, Faune Mamm. d'Europe, 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 bufl'.

Skull 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.

Remarks. — 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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744 tlOlDENTIA

identical. A specimen from Brelia, Pvoumania, in some respects
resembles the tyj^e of ArvicoJa ilhjrlcus, Init the small size of the
hind foot (25 mm.) renders its identification uncertain.

6. Bosnia. Dr. Floericke (c). 94. 1. 5. 1.

{Type of species.)
S. Breba, Roumania. [W. Dodson.) Lord Lilford (p). 4. 4. 6. GG.

ARVICOLA MUSIGNANI de .Sclys-Longchamps.

1830. Arvicola musignani de Selys-Longchamps, Revue Zoologique, p. 8,
January, 1839 (Rome, Italy).

1839. Arvicola destructor Savi, Nuovo Giorn. de' Letterati, Pisa, xxxvii,
No. 102, p. 204 (p. 7 of separate), February, 1839. (Maremma
Grossetana, Tuscany, Italy). For date of publication see de Selys-
Longchamps, Mioromammalogie, p. 93.

1845. ? [Arvicola musignani'] var. fuligijiosus de Selys-Longchamps, Atti
della sesta Riunione degli Scienziati Italiani, Torino, 1844, p. 322
(nomen nudum).

1857. Arvicola amphihius b. Blasius, Siiugethiere Deutscblands, 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 locdllti/. — Vicinity of Rome, Italy.

Geofjrapliical distribution. — Central Italy ; at present known
from the west coast only.

Diagnosis. — Size and general characters as in ArvicoJa iialicits,
but colour pale and yellowish, like that of A. saj/idus sapidus.

Colour. — Entire animal a light yellowish wood-brown, faintly
grizzled with black on median dorsal region and clouded by the
slate-grey underfur on throat and belly ; cheeks faintly more
yellowish than sides of body. Feet light drab. Tail obscurety
bicolor, blackish brown above, yellowish brown below.

Skull 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 "C
and 29 ; ear, 15 and 15. For cranial measurements see Table,
p. 743.

Specimens examined. — Seven, from the following localities in Italy:
Arsoli, Rome, G; Velletri, Rome, 1 (Mottaz).

6. Arsoli, Rome, Italy. G. Barrett-Hamilton (p). 11. 1. 2. 37.

ARVICOLA SCHERMAN Shaw.

(Synonymy under subspecies.)

Geograpliiral distribution. — West-central continental I^urope,
from the Pyrenees and Alps to the Baltic ; eastern limits of
range not known.

AUVICOLA 745

Dlagtiosin. — Essentlall}^ as in Arvicola terrestrin, 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.

SJcull and teeth. — The skull does not diflfer essentially from
that of Arvicola terrcsfris, but its fossorial characters are more
accentuated, especially in the terrestrial forms. Teeth without
.special peculiarities.

Hemarhs. — 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
related, 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. ? Slpalax] minor Leske, Anfangsgrunde der Naturgosch., i, p. 168

(Germany).
1801. Mus schermayi Shaw, Gen. Zool., ii, pt. i, p. 75 (Strassburg,

Germany).
1801. M[us] amph[ihius] albjis Bechstein, Gemeinn. Naturgesch. Deutsch-

lauds, I, 2d ed., p. 985 (Thiiringen, Germany).
1801. M[us]am2^h[ibius] ca»i<s Bechstein, Gemeinn. Naturgesch. Deutsch-

lands, I, 2d ed., p. 985 (Thiiringen, Germany).
1804. Mus terrestris Hermann, Observ. Zool., p. 59 (not of Linnaeus, 1758).
1804. [Mils'] scliertnaus Hermann, Observ. Zool., p. 59. Alternative for

terrestris; proposed but rejected on the ground that the animal

was the same as that of Linnteus (Strassburg, Germany).
1822. Arvicola argentoratensis Desmarest, Mammalogie, pt. ii, p. 281

(Strassburg, Germany).
1829. Lemmus arvalis fi buffonii Fischer, Synops. INIamm., p. 293 (near

Berlin, Germany ?). Based on Brants's account (Het Geschlact

der Muizen, p. 372) of two specimens in the Berlin IMuseum

supposed to be identical with the dark variety of " Le Campagnol "

described by Buffon, Hist. Nat. des Anim. \ii, p. 372.
1839. Arvicola terrestris de S61ys-Longchamps, Etudes de Micromamm.,

p. 97, pis. I and ii, fig. 6.
1858. Arvicola amphibius, c. A[rvicola'] terrestris auct., Blasius, Sauge-

thiere Deutschlands, p. 355 (part).
1910. Arvicola scherman scherman Miller, Proc. Biol. Soc. Washington,

XXIII, p. 21, March 23, 1910.
1910. Arvicola scherman Trouessart, Faune Mamm. d'Europe, p. x.

Tijijc locality. — Strassburg, Germany.

Geographical distribution. — Continental Eui'ope 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 • EODENTIA

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
terrestris. 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 fi'om
hair-brown to blackish. Tail blackish throughout or sprinkled
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.).

Feance : La Fert(5-Milon, Aisne, 1 (Lataste).

Germany: Stolpin, Pomerania, 1; Brunswick, 3 (U.S.N.JM. and
Merriam) ; Ingelheim, Rheinhesseu, 7 ; Heidelberg, Baden, 1 (U.S.N.M) ;
Lotzen, Saxony, 1 (U.S.N.JL) ; Ummerstadt, Thiiringen, 1 ; Strass, near
Burgheini, 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-HuNGABY : Hainspach, Bohemia, 4 (U.S.N.M.); Zuberer, 4.

1 al. Stolpin, Pomerania. Dr.H. Gadow (c & p). 82. 7. 31. 1.

4 (5, 3 ?. Ingelheim, Rheinhessen. C. Hilgert (c). 8. 11. 2. 52-58.

9. Strassburg, Alsace. O. Thomas (p). 8. 8. 10. 100.

(C. Mottaz.)
6, juv. Niesky, Silesia. (W.Bacr.) Dr. E. Hamilton (p). 97. 12. 4. 31-32
4. Zuberet"', Hungary. Budapest Museum (e). 94. 3. 1. G1-C4.

Arvicola scherjI(\.n e.xitus Miller.

1839. Arvicola terrestris Savi, Nuov. Giorn. de' Letterati, Pisa, xxxvir,

p. 300 (p. 3 of separate). Not Mus terrestris Linnaeus. (Geneva,

Switzerland.)
1845. ? [Arvicola terrestris'] var. nigcr De S^lys-Longchamps, Atti della

sesta Riunione degli Scienziati Italiani, Milano, 1844, p. 321

(Lausanne, Switzerland). Nomen nudum.
1845. ? [Arvicola terrestris] var. castaneiis De S61ys-Longchamps, Atti

della sesta Riunione degli Scienziati Italiani, Milano, 1844, p. 321

(Lausanne, Switzerland). Nomen nudum.
1857. Arvicola amphibius c. A[rvicola'] terrestris Auct., Blasius, Silugethiere

Deutschlands, p. .355 (part).

AUVICOLA 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 schcrman exitus Trouossart, Faune Mamm. d'Europe,p. x.

1911. Arvicola scherman cxilis Lydekker, Zool. Record, xlvii (1910),

Mamm., p. 54 (Accidental renaming of exitus).

Tupo localiii/. — St. Gallen, Switzerland.

Geographical (listrihution. — ^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-
lohasal length of fully adult skulls, 3.3 to 35 mm. ; colour of
upper parts a light yellowish brown, usually without much black
clouding ; tail buffy throughout or evidently bicolor (never
luiiformly blackish) ; auditory bullre 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 ampJnhius 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 bulk 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. ampliihius.
An exceedingly rudimentary sixth tubercle sometimes present.
Tail with annulations narrower than in A. ampliihius, about 20
to the centimeter at middle.

Colour. — Upper parts varying from a light broccoli-brown to
ochraceous-buflT, 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-
buft', in others more nearly a greyish cream-buS"; underparts a
paler, less slaty grey than in M. sclierman scherman, the throat
nearly the grey No. 9 of Ridgway ; chest and belly washed witli
cream-buff; feet ecru-drab, sometimes with a buffy tinge ; tail

(48

RODENTIA

whitish throughout or sprinkled witli blackish above, sometimes
rather distinctly bicolor.

SA-uU 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 shallowness of the rostrum as

Fig. \',i.
Arvicola sdierman exitits. Xat. 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 fi'om 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, 1 3 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 crania
measurements see Table, p. 750.

Specimens examined. — Eighty-two, from the following localities ; —
Feajstce : Gerbaniont, near Vagney, Vosges, 6 (Latasts) ; Cranves-
Sales, Haute-Savoie, 8 ; Lucinges, Haute-Savoie, 4 ; Jlontaubau, Haute-
Savoie, 2 ; Scientriers, Haute-Savoie, 1 (Mottaz) ; Ktupes, Doubs. 1
(IMottaz ; not typical).

ARVICOLA 749

Switzerland : Geneva, 9 (B.M. and I\Iottaz) ; Nyon, Vaud, 1 (]\rottaz) ;
Chesieres, Vaud, 2 (IMottaz) ; St. Corguos, Vaud, 1 (Mottaz) ; Lausanne,
Vaud, 2 (U.S.N.M.); Les Plans, Vaud, 10 (U.S.N.M.) ; Meiringen, Berne,
6 (U.S.N.M.) ; Vitznau, Lucerne, 2 ; St. Gallen, 13 (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.T\L) ; Roggvvoil, Thurgau, 3 (U.S.N.M.) ; Pribourg, 1 ; no exact
locality, 3.

AusTBiA-HuNGARY : Mittelbcrg, Vorarlberg, 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 c5. Cranves-Sales, Haute-Savoie,
Prance.

3 (J, 3 ?. Cranves-Sales, Haute-Savoic.

1200 m. (A. Robert.)
2. Jlontauban, Haute-Savoie.
1. Geneva, Switzerland.

6, 9. Vitznau, Lucerne.
6. Ziiberwangen, St. Gallen.
{E. H. Zollikofer.)

4 (5, 4 ?. St. Gallen. (E. H. Zullikofcr.)

1. Pribourg.

1. Switzerland.

2. Switzerland. (Moschlor.)

i. Mittelberg, Vorarlberg, Aus-
tria-Hungary. {Dr. Major.)

A. Robert (c & p). 5. 4. 9. 6-7.

0. Thomas (p). 5. 11. 18. 18-23.

A. Robert (c & p). 97. 1. 9. 5-6.

Baron de S61ys- 55. 12. 24. 358.

Lougchamps (p).

O. Thomas (c & p). 5. 8. 3. 19-20.

O. Thomas (p). 2. 8. 4. 48.

0. Thomas (p). 10. 8. 16. 1-8.

(10. 8. 16. 8. Type of subspecies.)

L. O. Galliard (p). 75. 9. 20. 1.

Baron de Selys- 45. 7. 5. 5.

Lougchamps (p).

E. R. Alston (p). 79. 9. 25. 47-48.

O. Thomas (p). 8. 11. 30. 17.

Arvicola SCHERMAN MONTicoLA de Selys-Longchamps.

1838. Arvicola monticola de S61ys-Longchamps, Revue Zoologique, p. 249.

1839. Arvicola monticola de Selys-Longchamps, Etudes de Jlicromamm.,

p. 92, pis. I and ii, fig. 3.
185.S. Arvicola ampliibius,c. A[rvicola'\ terrcsiris auct., Blasius, Saugethiere

Deutschlands, p. 355 (part).
1910. Arvicola scherman nionticola Miller, Proc. Biol. Soc. Washington,

XXIII, p. 22, March 23, 1910.
1910. Arvicola scherman vwnticola Trouessart, Paune Mamm. d'Europe,

p. X.

Tijj^)e locaVity. — •St. Bertrand de Comminge, Hautes-Pyrenees,
France.

Geographical distribution. — Pyrenees and their immediate
neighbourhood (known at present from the French side only) ;
Puy-de-Dume 1

Diagnosis. — Like Arvicola schermayi exitiis, but auditory bullae
usually larger and more evenly inflated, and first lower ifiolar
with anterior loop longer and narrower than in the Alpine form.
Habits strictly terrestrial, mole-like.

760

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A. scherman scherman.

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France ; La Perte-Milon
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Lotzen, Saxony .

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France : Caterille, Haute-Garonne

Luchon

Pyrenees no exact locality

Biarritz, Basses-Pyren6es

France: ISIontauban, Haute-Savc
Cranves-Sales, Haute-Sa

Lucinges, Haute-Savoie
Switzerland ; Geneva .

Chesi^res, Vaud

Vitznau

St. Gallen ."

Ziiberwangcu, St. G

Mels

Degersheim ,,

10^ EODENTIA

Measurements. — Average 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 four 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-Garoune, 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.).

Remarks. — 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 bullje and in the outline
of the anterior loop of ni^.

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 (5, 5 9. Biarritz, Basses-Pyrenees, J. P. Davison (c & p). 6. 6. 4. 15-22.
France.
1. Pyrenees. Baron E. do Selys- 45. 7. 5. 4.

Longchamps (p).
1. Pyrenees. Purchased (E^ank). 45. 5. 23. 7.

1. Pyrenees. Stockholm iluseum (e). 46. 6. 2. 66.

Genus PITYMYS McMurtrie.

1830. Psammomys Le Conte, Ann. Lye. Nat. Hist., New York, ni, p. 132

(P. pinetorum Le Conte). Not of Cretzschmar, 1828.

1831. Pitijuiys Mcilurtrie, Cuvier's Animal Kingdom, American edition, i,

p. 434 [Fsammomys pinetorum Le Conte).

1831. Ammmnys Bonaparte, Saggio Distrib. Metod. Anim. Vert., p. 20,
footnote {Psammomys pinetorum Le Conte).

1836. Pinemys Lesson, Hist. Nat. des Mammif. et Ois. decouv. depuis
1788, Compl. Oeuvres de Buffon, v, p. 436 (Psammomys pinetorum
Le Conte).

1857. Microtus Blasius, Siiugethiere Deutschlands, p. 387 (sub-genus). Not
Microtus Schcank, 1798.

1867. Terricola Fatio, Les Campagnols du Bassin du Leman, p. 36 (sub-
genus to contain Arxncola subterraneus de S^lys-Longchamps, and
A. savii de Selys-Longchamps). Not of Fleming, 1828.

1877. Micrurus Major, Atti della Soc. Toscana di Sci. Nat., in, p. 126 (sub-
genus to contain Arvicola nebroclensis Mina-Palumbo).

1887. Pitymys Lataste, Ann. I\Ius. Civ. Stor. Nat. Geneva, 2d ser., iv,
p. 266 (subgenus).

PITYMYS 753

189G. Pitymys Miller, North American Fauna, No. 12, p. 58, July 23, 1896

(sub-gonus).
1907. Pitymys Mottaz, Bull. Soc. Zool. de Prance, xx, p. 27, September

1907 (genus).

Type si)ecies. — Psammoiiiys pinctorum Le Conte.

Geograpliical diHirihution. — Central and southern continental
Europe, eastward into Asia Minor ; eastern and south-eastern
United States ; southern Mexico.

Characters. — Like M'lcroius but sole with five tubercles, and
maiumaj only four ; external form slightly or considerably modified
for underground life ; skull in most sjaecies showing some trace
of fossorial modification, this character often conspicuous ; enamel
pattern variable, but first inner and first outer triangles of m^
broadly communicating.

RemarJcs. — 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 qh^.

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 bullte about 05

per cent of occipital breadth.

Third inner re-entrant angle of Wi^ shallow

(Roumania) P. dacius, p. 760.

Third inner re-entrant angle of w^ deep.

Back strongly s.uff used with bufiy ; feet whitish
in noticeable contrast with back (Basses-

Alpes) P. drnentius, p. 762.

Back without noticeable bufiy suffusion ; feet
dusky, not noticeably contrasting with

back P. subterraneus, p. 755.

Brain-case moderately flattened ; colour

dark (Belgium to Transylvania) P.s. subterraneus, -p. 758.

Brain-case much flattened ; colour less

dark (Puy-de-D6me region, France) P. s. cajmeiniia, p. 760

Dorsal profile of skull obviously convex from
nasals backward ; brain-case slightly de-
pressed, its depth including auditory bullas
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.) 1'. viult'qilcx, 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 m^ well developed, with three
definite enamel sides and a distinct central
osteodentine 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. planiccps, 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 m^ broadly communi-
cating P. gcrbii, p. 773.

Interparietal strap-shaped, its lateral extremi-
ties truncate, broadly in contact with
temporals ; first and second triangles of
m^ usually isolated.
General outline of brain-case sub-quadrate,

the upper surface rather flat (Italy) ... P. savii, p. 768.
General outline of brain-case not evidently
sub-quadrate, the upper surface rounded
off at sides.
Mesopterygoid space rather wide (Sicily) P. ncbrodcnsis, p. 770.
Mesopterygoid space narrow (South-
western France) P. pyrcnaicus, p. 770.

General colour above clear hair-brown
without noticeable yellowish cast

(Pyrenees) , P. p. 2iyrenaicus, p. 771.

General colour above brown with a
noticeable yellowish cast (Plains

of south-western Prance) P.p. hrunneus, p. 772.

Outer triangles of vv^ 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. dcprcssus, p. 779.

Brain-case not much flattened, the occipital
depth noticeably more than one-half
occipital breadth.
Upper incisors slightly projecting (Portugal) P. lusitanicus, p. 77G.
Upper incisors nearly vertical.

Skull narrow; outline of brain-caso dis-
tinctly elongate (North - v^esteru

Spain) P. mariir, p. 777.

Skull broad; outline of brain-case not
specially elongated (North-central
Spain) P. pclandunius, p. 778.

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 Prance) P. p7-ovincialis,'p. 7 85.

Condylobasal length of fully adult skulls

24 to 26-5 mm.

Size large, condylobasal length of largest

skulls about 2G"5 mm.; transverse

diameter of auditory bulla about 7 mm.

(Montenegro) P. thomasi, p. 786.

Size medium, condylobasal length of largest
skulls not more than 25 mm. ; trans-
verse diameter of auditory bulla about
G 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. duodccimcostatus,

OcciiJut not very obliquely truncate, so P' '
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 upxser tooth-row 6 to 6 ■ 4 mm.
Colour above a dark bister brown

(Dehesa de Valencia, Spain) P. i.pascims, p. 783.

Colour above alight yellowish brown.

General colour above hair-brown

with or without a buf!y cast

(Central Spain) P. i. centralis, p. 782.

General colour above a pale huffy
broccoli-brown or drab (South-
eastern Spain) P. i. ibericus, p. 782.

PITYMYS SUBTERRANEUS de Selys-Longchamps.
(Synonymy under subspecies.)

Geogra-pMcal distribution. — Central Europe from Belgium to
central Fi'ance, eastward through Switzerland to Transylvania.
Details of distribution imperfectly known.

Diac/nosis. — 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, depi-essed, the dorsal profile nearly
flat or at most only a little convex from nasals backward ; brain-
case much depressed, its depth including auditory bulla; about
65 per cent of occipital breadth ; third 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 suflusion of
buify ; feet dusky, not contrasting with colour of back.

External characters. — General external form essentially as in

3 c 2

756

RODENTIA

Microtus 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 Microtus arvali.s, 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 Microtus arvnlis. Muzzle not
peculiar. Front foot as in Microtus arvalis, except that the claws
ai'e 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. Mammaj :
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-
brown or wood-brown, but rarely if ever with any distinct buffy
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 interjjarietal, 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 bullae,
the general outline of skull as viewed from the side rather
narrowly cuneate. Brain-case long as compared with that jjortion
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 ; lambdoid
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 bulke about 65 jjer cent of occipital breadth ; occiput

* A character that ofteu proves convenient in distinguishing between
imperfect sliulls 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 bulhe, their tips descending below lip of foramen
magnum, the ridges extending vipward 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 buUas moderately large, about as in Microius
nrvalis, 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
I'elatively smaller than in Mkrotus
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 Microius
arvalis, their posterior extremity at level of front of m^, 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-deflned, 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 M. 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
ip. relatively shorter, strongly convex anterior
^i border. Molars so like those of Microius
^^ arvalis that aside from the pecuharities of
"^ the first mandibular tooth there is nothing to
^, distinguish witli certainty between the two
"^ patterns. Crown of third upper molar distinctly
S longer than that of second. In Pitymys sub-
^ terraneus there is, however, a tendency for the
re-entrant angles to be wider and the closed
triangles relatively smaller, while the pattern of
nn'"^' Enamel" '"^ 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 looji and second triangle. Second
upper molar strictly as in M. arvalis. First lower molar
like that of Microfus 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.

Bemarks. — Although bearing a strong superficial resemblance
to Microius arvalis, and more particularly to the Alpine M.
inccrtns, this species is distinguishable externally by its rather
smaller ears, and by the number of mammaj and plantar tubercles.
In colour it is usually a clearer, less yellowish brown than the
smaller forms of Microtns. Old and faded museum specimens are
often impossible to determine with certainty. The skull is
recognizable by its small size, nearly Hat dorsal profile, wide
interorbital region, and by the almost perpendicular upper
incisors. Two geographical forms are known.

Pitymys subterraneus subterraneus de Selys-Longchanips.

1836. Arvicola subterraneus de Selys-Longchamps, Essai Monographique

sur les Gampagnols des environs de Li6ge, p. 10 (Waremme, Li6ge,

Belgium) .
1845. II\_ypud:nis'] rufesccnte-fuscus Scbinz, Synopsis Mamm., ii, p. 240

(Urserenthal, Uri, Switzerland). See ]Mottaz, Bull. Soc. Zool. de

France, xx, p. 27, September, 1907.
1S45. Hliipudnnis'] rufofuscus Schinz, Synopsis Mamm., ii, p. 240 (synomym

of rufcscente-fuscus).
1857. Arvic-?la subterraneus Blasius, Siiugetbiere Deutscblauds, p. 338.

PITYMVS 759

1900. [Arvicola acjreslis] fiisca Fatio, Rovuc Suisse cle Zool., vin, p. 472
(Untervats, Grisons, Switzerland). Sec Mottaz, Bull. Soc. Zool.
de Geneve, i, p. 15'J, November 15, 1908.

1907. Pitijnujs subterraneiis Mottaz, M6m. Soo. Zool. de Prance, xx, p. 27,
September, 1907.

1910. Pitijmijs subterraneiis Trouessart, Faunc Mamm. d'Europe, p. 185.

Type locality. — Waremine, Liege, Belgium.

Geographical distrihufion. — 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, Vaud, 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 foet, 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).

Feance : Near Boulogne, Pas-de-Calais, 3; Dinan, Cotes-duNord, 1 ;
Etupes, Doubs, 2 (B.JM. 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 Goscheuen, Uri, 6
(Mottaz) ; Furka Pass, Uri, 1 (Mottaz) ; Murgsee Eegion, St. Gallon, 3
(U.S.N.M. and Mottaz) ; St. Moritz, Grisons, 2 (Geneva) ; Poschiavo,
Grisons, 1 (Mottaz) ; Untervats, above Chur, Grisons, 1 (St. Gallen ; type
of fusca Fatio).

AusTBiA-HuNGARY : Hatszeg, Hunyad, Hungary, 8.

4 & 2 al. Liege, Belgium. Baron E. de S61ys- 37. 1. 3. 170-171,

Longchamps (c & p). 174-175, 177^178.
4. Li(5ge. {de Sehjs- G. R. Waterhouso (p). 44. 10. 4. 15-18.
Longchamps).
2 6,19. Boulogne, Pas -de - 0. Thomas (c & p). 98. 1. 9. 21-23.
Calais, France.
6. Etupes, Doubs. (C. O. Thomas (p). 8. 8. 10. 130.

Mottaz).
7 J, 1 ?. Hatszeg, Hunyad, C. G. Danford (c). 3. 11. 8. 44-51.

Transylvania.

i60

RODENTIA

PiTYMYS SUBTERRANEUS CAPUCINUS Miller.

1908. Pitymys subterrancns capucimis Miller, Ann. and Mag. Nat. Hist.,

8th ser., i, p. 202, February, 1908.
1910. Pitymys subterraneus capucinus Trouessart, Faune Mamni. d'Europe,

p. 185.

Tjipe locality. — Spruce forest near " Salon du Capucin,"
Mont-Dore, Puy-de-D6me, 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 Piidgway but with an e"vadent butfy cast ; feet a clear,
very pale smoke-grey.

Skull and teeth. — In all general features the skull agrees
with that 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 ofl" 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. IMiller (c). S. 8. 4. 266-267.

(8. 8. 4. 2C7. Type of subspecies.)

PITYMYS DACIUS Miller.

1903. Pitymys dacius Miller, Ann. and Mag. Nat. Hist., 8th ser., i, p. 202,

February, 1908.
1910. Pitymys dacius Trouessart, Faune Mamm. d'Europe, p. 188.

Ti/pe localiti/. — Gageni, Prahova, Pioumania (at foot of Car-
pathians, north-west of Bucharest), Rouraania.

Geographical distribution. — Known at present from the tyi^e
locality only.

Diagnosis. — Similar to PitijmijS subterraneus but skull larger,
the nasals more strongly bent downward anteriorly, the brain-
case more depressed posteriorly ; auditory bullfe larger ; posterior
upper molar with third inner re-entrant angle very shallow.

Colour. — The colour of the type is indistinguishable from that
of typical P. subterraneus.

Shdl. — In general the skull resembles that of Pitymys sub-
terraneus, but the brain-case is more depressed posteriorly and

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the nasals are more bent downward in front, the dorsal profile
being thus made faintly convex throughout. Though even more
flattened than in P. suhterraneus capucinus, the brain-case is as
long relatively to its breadth as in true suhterraneus. Auditory
bulliB larger than in P. suhterraneus though not peculiar in form.

Teeth. — The teeth resemble those of Pitymys suhterraneus,
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.

Specimen examined. — The type.

9. Gageni, Prahova, Roumania. Lord Lilford (p). 4. 4. 6. 65.
(W. Dodson.) {'^'upc of species.)

PITYMYS DRUENTIUS Miller.

1852. Arvicola (Miavtus) sclijsii Gcrbc, Rev. et ]\Iag. do Zool., 2d ser., iv,
p. 159, j\Iarch, 1852 (Near Barcelonuette, Basses-Alpes, France).
Not Arvicola sclysii Bonaparte, 1845.

1911. Pitymys driientius Miller, Proc. Biol. Soc, Washington, xxiv, p. .39,
February 24, 1911 (Substitute for selysii).

Type locnliti/. — Terres-plaines, near Barcelonuette, Basses-
Aljjes, France.

Geographical (llstrihution. — South-western Alps ; limits of
range not known.

Diagnosis. — In general similar to Pitynvjs suhterraneus, 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.

Slcull and teeth. — The skull and teeth resemble those of
Pitymys suhterraneus, 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 j«^ with a tendency to develop an evident

PITYMYS 763

convexity, the first stage in the formation of a postero-internal
loop. Such an incipient looji rarely if ever occurs in P. swbtcr-
rancus, though it is common and not infrequently well developed
in P. fat'ioi 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.

Specimens examined. — Eleven, from the following localities in France :
neighbourhood of Barcelounette, Basses-Alpes, 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. Barcelonnette, Basses-Alpes. 0. Thomas (p). 8. 8. 10. 126-129.

(C. Mottaz.)

PITYMYS FATIOI Mottaz.

1909. ntymys multiplex fatioi Mottaz, Bull. Soc. Zool. do Geneve, i,

p. 180, January 15, 1909. Type in Mottaz collection.

1910. Pitymys multiplex fatioi Trouessart, Pauue Mamm. d'Europe, p. 189.

Tiipe locality. — Zermatt, Valais, Switzerland.

Geographical distribution. — At present known only from the
neighl)ourhood 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 bullte 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 examined. — Twenty-six, all from the neighbourhood of
Zermatt (U.S.N.M., Mottaz and Geneva).

BemarJis. — 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

PITYMYS MULTIPLEX Fatio.

1905. [Arvicola^ multiplex Fatio, Arch. Sci. Phys. et Nat., Geneve, 4th ser.,

XIX, p. 193, February 15, 1905 (Lugano, Ticino, Switzerland).

1906. M\icrotns] leponticus Thomas, Ann. and Mag. Nat. Hist., 7th ser.,

XVII, p. 419, April, 1906 (Locarno, Ticino, Switzerland). Type in

British Museum.
1908. Pitymys multiplex Mottaz, BuU. Soc. Zool. de Geneve, i, p. 165

November 15, 1908.
1910. Pitymys multiplex Trouessart, Faune Mamm. d'Europe, p. 189.

Tijpc locality. — Lugano, Ticino, Switzerland.
. Geographical clistrihution. — Italian Switzerland and northern
Italy, south to Florence ; limits of range not known.

Diagnosis. — Similar to Pitymys druentius and P. fatioi but
decidedly larger (hind foot usually 16 to 17 mm., condylobasal
length of skull, 24 to 2.5 mm.) ; skull with dorsal profile notice-
ably convex ; auditory bullfe not specially inflated.

External characters. — Ears relatively smaller than in Pitymys
suhierraneus ; 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
suhierraneus. a.fi to need no detailed description. The dark wash
along back tends to be rather extensive, and the entire upper
surface is frequently suffused witli a light bister or raw-umbei'.
Feet usually with less dusky clouding than in P. snhterrancus.

Shull. — As compared with that of Pitymys subterrancus 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. subterrancus, 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 Microtits. Length of brain-case to back of
interparietal about equal to vvidth over roots of zygomata.
Occiput squarely truncate posteriorly, the condyles nearly or
entirely hidden when skull is viewed from above. Auditory
bullaj of the same relative size as in P. subferraneus, but with
smoother, more evenly inflated finish.

Teeth. — The teeth resemble in general those of Pitymys stib-
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 outer 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 oj^ening rather widely into terminal loop, though
sometimes completely isolated, the exact opposite to the condi-

765

Fig. 157.

PUyimjs miMvplex.
Enamel pattern. X 5.

: ear, 10. Adult

tion in P. suhterraneus. Second upper molar showing the
maximum of the tendency common to the three related
forms, druentiiis, fat'toi and multiplex, for the inner side of third
triangle to develop an imperfect postero-
intei'iial 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
Microtits agrestls, though in none is it com-
pletely isolated from the outer triangle.

Measurements. — Adult male from Gor-
dola, near Locarno, Ticino, Switzerland
(type of leponticus 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, Switzei'land : head and body, 1 00 ;
tail, 31 ; hind foot, 17. Adult male from Mirabello, Mon-
ferrato, Italy: head and body, 102; tail, 33; hind foot, IG'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 exanmied. — Thirty-uine, from the following localities : —
Switzerland : Near Locarno, Ticino, 5 (including type of Icponticus) ;
Bellinzona, Ticino, 1 ; Comano, Ticino, 4 (B.M. and Mottaz) ; near
Lugano, Ticino, 12 (B.M. and Mottaz) ; Pico Magadino, Ticino, 1
(Ghidini).

Italy : Porlezza, Corao, 3 ; Domodossola, Novara, 1 ; Ceresole d'Alba,
Turin, 1 (Turin) ; Mirabello, Monferrato, Alessandria, 1 (Turin) ; Vaccarezza,
G (Genoa) ; Caorsi, Antola, 2 ; Prato, Genoa, 1 (U.S.N.M.) ; Borzoli,
Genoa, 1 ; Florence, 1 (U.S.N.M.).

Beniarhs. — 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 i, 2 9. Locarno, Ticino, Switzer- O. Thomas (c & p). 5. S. 2. 11-14.
land. (5. 8. 2. 11. Type of P. Leponticus Thomas.)

?. Borzoli, Genoa, Italy. Marquis G. Doria (p). 8. 7. 18. 7.

766

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768

PITYMYS SAVII de Selys-Longchamps.

1838. Arvicola savii de S^lys-Longchamps, Revue Zoologique, p. 248,

October, 1838.
1845. A[7-vicola'] selysii Bonaparte, Atti della sesta Riunione degli Scien-

ziati Italiani, Milano, 1844, p. 350 (synonym of savii ; Pecchioli

cited as authority).
1857. Arvicola savii Blasius, Siiugethiere Deutschlands, 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 local! fj/. — Neighbourhood of Pisa, Italy.
GeograpJiical distribution. — 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 subler-
rancus, but feet more robust, ear relatively smaller, and palmar
and plantar tubercles more reduced ; skull usually more heavily
built, much less de2:>ressed, 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-hroAvn. Feet whitish gi'ey as in P. multiplex,
none of the specimens examined showing any trace of the
dusky suffusion characteristic of P. suhterraneus.

Shill. — 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 zygomata 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 tliat of P. multiplex,
less elongate. The rostrum is also

Fia. 158.
Pitymys saini. Nat. size.

and the brain- case
relatively shorter.

769

Pity my s savil.
Enamel pattern. X 5.

Tccih. — Except for the form of the third upper molar the
teeth show no pecuharities. This tootli is so shortened that tlie
length of its crown does not exceed tliat 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. suhterrancus, the
third triangle not normally showing any trace
of an incipient postero-internal loojj.

Measurements. — Adult male from Go/.zano,
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, 2.5 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. Augelo, Pavia, 3 ;
Serravalle, Piedmont, 1 ; Ceresole d'Alba, Turin, 5 (Turin) ; Frugarolo,
Alessandria, 21 (B.M., Genoa, and Mottaz) ; Mirabello, Monferrato, Ales-
sandria, 2 (Turin) ; Caorsi, Mount Antola, 3 ; Modena, 3 ; Pisa, 1 (Mottaz) ;
Florence, 29 (U.S.N.M. and Mottaz) ; Urbino, Marobe, 3 ; Benagna,
Perugia, 1 (Genoa) ; Ostia, Rome, 1 ; Chieuti, Foggia, 24 (B.M. and Genoa).

1.

Milan, Italy.

Baron E. de Selys-

Longchamps (p).

Marquis G. Doria (p).

Marquis G. Doria (p).

Marquis G. Doria (p).

<;, 2 9 al. St. Angelo, Pavia.
9 al. Serravalle, Piedmont

2 iJ, 2 9, 2 al. Frugarolo, Alessan-
dria.

6. Mirabello, Alessan- Dr. E. Festa (p)

dria.
(?, 2 9 al. Caorsi, Liguria.
6, 9, 9 juv al. Modena.
2 9, juv al. Urbino, Marche.
1. Ostia, Rome.

2 6, 9. Chieuti, Foggia.

Marquis G. Doria (p).
Marquis G. Doria (P).
Marquis 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.

770

PITYMYS NEBRODENSIS Miua-Palumljo.

18G8. Arvlcola nchrodensis Mina-Palumbo, Aun. Agric. Sicil., xri, p. Gl.
1910. Piti/mys savii nchrodensis Trouessart, Fauue Mamm. d'Europe,
p. 187.

Type locality. — Le Madonie, Sicily.

Geograph teal distribution. — Sicily.

Diagnosis. — Similar to Pitymys savii, hut skull witli deeper,
more elongate, less fattened brain-case, and more evidently
convex dorsal profile ; mesopterygoid space relatively wider than
■in P. savii.

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.).

.5,?. Palermo, Sicily. J. I. S. Whitaker (p). 98.10.6.16-17.

3 <5, 1 ?. Castelbuono. {A. Robert.) O. Thomas (p). \q g' -,' 26I27'

6 al. Madonie. Marquis G. Doria (p). 90. 3. 5. 14.

PITYMYS PYRENAICUS de Selys-Longchamps.

(Synonymy under subspecies.)

Geogrnpliical distribution. — Pyrenees (known from the French
side only) and region to the north as far as the Garonne. Limits
of range aiot known.

Diagnosis. — Related to Pitymys savii and P. nebrodensis ;
general form of skull somewhat intermediate between that of the
Italian and Sicilian s^jecies, but mesopterygoid fossa decidedly
narrower.

* 6. No. 98. 10. G. 16, Palermo, Sicily.

PITYMYS 771

Slnll. — The skull has much the same general outline as in
P. iichrod crisis, that is, the brain-case is shghtly longer than
broad and the dorsal profile is slightly convex, but the depth is
about as in P. savil and P. multiplex. It is readily distinguished
from that of P. snbterraneus 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 PltumyH 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 dc Selys-Longchamps.

1847. A[rvicola] pyrenaicus de S61ys-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 pyrenaicus Trouessart, Faune Mamm. d'Europe, p. 189.

Type locality. — Bagneres de Bigorre, Hautes- Pyrenees,
France.

Geographical distiihution. — 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.

Sjjecimcns examined. — Thirteen, from the following localities : —
France: L'Hospitalet, Ariege, 2; Ax-les-Thermes, Ariege, 3; Luchon,

Haute-Garoune, 1 ; Bareges, Hautes-PyrenSes, 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.
(.4. Robert.)

2 6. Bareges, Hautes-Pyr6n6es. 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.

6. No. 8. 8. 4. 271, Barfeges, Hautes-Pyr6n6es, France.

3 D

772

PrrvMVs Pyhunaicus bkunneus Miller.

l90S. Pili/mi/s pyrcnaicus bninncus IMiller, Ann. and Mag. Nat. Hist.,
8th ser., i, p. 203, February, 1908. Type in British Museum.

1910. Pitijmys pyrcnaicus bninncus Trouessart, Paune Mamm. d'Europe,
p. 189.

Tjipe locallfij. — Forest of Boucoiine, Gers, France.

Geographical distribution. — Lowlands of south-western France
from near base of Pyrenees to the Uaronne. Limits of distribu-
tion not known.

Diagnosis. — Like Pifi/mys pyrenai'cus pyrenaiciis, but colour of
upper parts with a decided cast of Ijutfy or pale wood-brown,
much as in P. savii.

Measunnncnts. — 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.

Specimens examined. — Fifteen, from the following localities in south-
western France : Forest of Bouconne, Gers, 14 ; Cadillac, Gironde, 1
(U.S.N.M.).

4 6, .3 ?. Foret de Bouconne, Gers, 0. Thomas (p). 6. 4. 1. 79-85.

France. {A.Robert.) (6. 4. 1. 82. TT/pe of subspecies.)

PITYMYS PLANICEPS Miller.

1908. Pitymys 2Jt(i,nice2JS Miller, Ann. and Mag. Nat. Hist., 8th ser., i,

p. 203, February, 1908. Type in Lataste Collection.
1910. Pitymys planiceps Trouessart, Faune Mamm. d'Europo, p. 190.

Type locality. — Bareges, Hautes-Pyrenees, France.

Geographical distribution. — Known only from the type
locality.

Diagnosis. — Size about as in Fiti/mys 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. pjyrcnaicus.
External characters not known.

Skull and teeth. — In general outline as viewed from above
the skull resembles that of Pitymys sacii 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. sub-
terraneus, but with the normal characters of the species carried
to the extreme. The dorsal profile is essentially straight from
posterior e.xtremity of nasals to back of interparietal, though
there is a faint concavity in interorbital region and a slight
•convexity at middle of brain-case. Interoil^ital region about as

PITYMYS 7 7. "5

in P. scuui, tlie temporal ridges wide apart and barely per-
ceptible. Auditory bulhe rather larger than usual in P. nubter-
raveus and more smoothly inflated. As compared with that of
P. pyrcnairus the skull is immediately recognizable by its flat
dorsal pi-otile and short, broad, excessively flattened brain-case.
Teeth as in Piii/)itus pyrenairiis.

Measurements. — External measurements not known. For
cranial measurements see Table, p. 775.

Specimen examined. — Tho type.

PITYMYS GERBII Gerl)e.

187'J. Arvicola [Microtus) gcrbii Gerbe, Le Naturaliste, i, p. 61, July 1,

1879.
1880. Arvicola (Microtiis) gerbei Gerbe, Bull. Soc. Zool. de France, Paris,

V, p. 49, pi. IV, figs. 1-9.
1910. Pitymys gerbei Trouessarfc, Faune Mamm. d'Europe, p. 186.

Type locality. — Dreneuf, Loire-Inferieure, France.

GiiKjra/phical distrihution. — 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.

External characters and colour. — Apparently about as in
P. pyrenaicus, so far as can be determined from a somewhat
faded mounted specimen.

SJmll. — The skull differs from that of Pitymys pyrenaicus in
greater relative depth and breadth throughout, shorter, heavier
rostrum, shorter incisive foramina, wider interorbital region,
and in the very peculiar form of the interparietal, as already
described ; na-sals conspicuously angular-emarginate posteriorly ;
auditory bullae rather large.

Teeth. — The teeth do not differ from those of Pitymys
pjyrenaicus 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 ihericus 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.

Specimen examined. — One from Dreneuf (paratype) in Paris Museum.

Peniarls. — This species is well characterized by the great
general breadth of skull, unusually wide interorbital region, and
vei-y peculiar interparietal. The last chai'acter might be sup-
posed to. be an accidental variation or abormality were it not

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776 KODENTIA

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 ihericus and its allies. The depth
of skull relatively to its length is intermediate between that of
P. pyrcnaicus and P. ibericus.

PITYMYS LUSITANICUS Gerbe.

1879. Arvicola (Microtus) lusitanicus Gerbe, Rev. et Mag. de Zool., 3rd ser.,

VII, p. 44.
1905. Microtus (Pitymys) lusitanicus Major, Anu. and Mag. Nat. Hist.,

7th ser., xv, p. 612, May, 1905.
1908. P[itymys~\ lusitanicus Miller, Ann. and Mag. Nat. Hist., 8th ser., i,

'p. 204, February, 1908.
1910. Pitymys lusitanicus Trouessart, Faune Mamm. d'Europc, p. 193.

Type locality.— 'Portiigal.

Geographical (listrihution. — Portugal ; limits of range un-
known.

Diagnosis. — 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, aljout 74 per cent of occipital breadth ; tliird 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 suhterrancus,
except that the underparts are a less clear grey. Upper
parts a dark hair-brown tinged with bister or sepia, and very
faintly grizzled by black tips and liglit reflections. Underparts
a dull ijuffy grey, in some specimens becoming almost a dull
ochraceous-buff' on cheeks and (in sides along border of dark area ;
feet bufty grey without dusky wash ; tail obscurely bicolor, bufly
white below, dusky mixed with buflfy 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 zygomatic 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. nebrodensis or
P. subierraneus, so that the condyles are usually visible when
skull is viewed from above. Auditory buUse 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 Pitjimi/s
.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 tirst 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 terminal 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 t;he 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. siihterraneus, the third
triangle without trace of postero-internal loop. Ujjper 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 exmnined. — Eighteen, all from Cintra, Portugal.
16 7 9, 2 al. Cintra, Portugal. 0. Thomas (c & p). 98. 2. 2. 37-52.

PITYMYS MARliE Major.

1905. Microtus (Pitjjmys) marix Major, Ann. and Mag. Nat. Hist., 7th ser.,
XV, p. 515, May, 1905. Type in British Museum.

1908. P[itymris] viariie Miller, Ann. and Mag. Nat. Hist., 8th scr., i,p. 201,
February, 1908.

1910. Pityynys -niarix Trouessart, Faune Mamm. d'Europe, p. 192.

Type locality. — Villalba, Lugo, Galicia, Spain.
Geographical distribution. — Known at present from the type
locality only.

* Essentially similar to those of P. ibericus, fig. 102 (p. 781).

778 EODENTIA

Diagnosis. — Similar to PiUjmys 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 probably similar to that of Pity my s
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 suhterraneus, 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 prohle is much less convex, and general outline
viewed from the side is less strongly cuneate. Auditory bullaj
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.— E,\gh\,, all from north-western Spain.

2 (5, 6 al. Villalba, 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 pdandonius Miller, Ann. and ^Mag. Nat. Hist., 8th ser., i,

p. 204, February, 1908. Type in British Museum.
1910. Fitymys pclandunius Trouessart, Faune Mamm. d'Europe, p. 192.

Type locality. — Silos, Province of Burgos, Spain. Altitude
about 3000 feet.

GeograpMcal distribution. — Not known at present elsewhere
than in the neighbourhood of the type locality.*

Diagnosis. — Similar to Pitymys murise 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.

Skull and teeth. — The skull differs from that of both Pitymys
lusitanicus and P. mcirise in its broader more quadrate brain-case.
Posteriorly the brain-case is rather low, as in P. marise, though
much less reduced in depth than that of P. depressus. Teeth as
in P. mari'cf^, the upper incisors nearly vertical.

Measurements. — 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

<le la Reina, Burgos: 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 Eeiua, Burgos, 1.

2 ?. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 294-295.

(8. 8. 4. 294. TijiJe of species.)

PITYMYS DEPRESSUS Miller.

1908. Pitymys deprcsstis Miller, Ann. and Mag. Nat. Hist., 8th ser., i,

p. 204, February, 1908. Type in British Museum.
1910. Pitymys dcpressus Trouessart, Fauue Mamm. d'Europe, p. 193.

Type locality. — Rascafria, south side of Hierra de Gundarrama,
Province of Madrid, Spain.

Geographical distribution. — Central 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 bullfe.

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. pelandonius 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 emarginate, 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 conspicuously beyond level of sides of brain-case. Length
of brain-case to posterior edge of interparietal less than breadth
over roots of zygomata, the general outline sub-circular ; occipital

4^^^n^

FIO. 160.

Pitymys depressus.

Nat. size.

780

RODENTIA

region obli(i[uely truncate posteriorly, 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 bullas very small and low, their greatest
diameter contained about four times in condylobasal length of
skull (about 3 to Sh times in lusitanicHs, 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.

Measurements. — 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.

Specimens 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. Eascafria, Madrid. M. de la Escalera (c). 6. 11. 4. 10-15.

(6. 11. 4. 15. Type of species.)
9. Villalba, Madrid. 0. Thomas (p). 8. 2. 9. 212.

{N. Gonzalez).

PITYMYS IBERICUS Gerlie.

(Synonymy under subspecies.)

Geographical distribution. — Probably the entire Iberian Penin-
sula, though at present known only from central and southern
Spain.

Diagnosis. — Larger than Pitymys Jusitanicits 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 inci.sors
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 difl'ering rather con-
spicuously from that of Pitymus snhterraneus and Microius arvalis
in its much more evident adaptation to underground life, the
head large, with incisors noticeably protruding from mouth,
the neck short and thick ; ears nuicli reduced, almost hidden in
the fur ; feet sti-ong 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 rolati%'ely smsdler area than in P. suhtcrraneus ;
tail short, scarcely more than one-cjuarter as lung 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 PHjimya Insitcmicus, 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 I'egion. 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
bullas 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 ihcricus 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. ici.
Pitymys ibericus. Nat. size.

Pitymys ibericus.
Enamel pattern. X

782

RODENTIA

PlTYMYS IBERICUS IBERICUS Gei'be.

1854. Arvicola ihericiis Gerbe, Revue Zoologique, 1854, p. 400. Type iu

Paris Museum.
1879. Arvicola (Microtus) ibericus Gerbe, Rev. et Mag. de ZooL, 3rd ser.,

VII, p. 42.
1905. Microtus ibericus Major, Ann. and Mag. Nat. Hist., 7th ser xv

p. 515, May, 1905.
1908. Pitymys ibericus ibericus Miller, Ann. and Mag. Nat. Hist., 8th ser.

I, p. 205, February, 1908.
1910. Pitymys ibericus Trouessart, Faune ilamm. d'Europe, p. 190.

Type locality. — Province of Murcia, 8paiu.

GeograpMcal 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 slightly
tinged with' bister along middle of back and suffused with lauffy
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. Feet clear greyish white. Tail whitish with a
varying amount of dark brown along median dorsal region, this
brown occasionally almost ab.sent.

Measurements. — Type (adult): head and body, 114 (skin);
tail, 26 ; hind foot, 18. Adult female from Venta del Paul,
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, 9. 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., 8th 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.

Geofjrapliical distribution. — Central Spain from the province
of Bui'gos south to Valencia and Seville ; exact limits of range
not known.

PITYMYS 783

Di(t(jnoms. — Not so large as P. iber'wus iberlcus (hind foot,
1() to 1 7 • 2 instead of 17-6 to 18-4); colour hair-brown usually
tinged with bufty, never with the pallid 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 Doilana, Huelva, 2.

8 (J, 2 ?. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 278-287.

(8. 8. 4. 278. Type of subspecies.)

1 al. Burgos. N. Gonzalez (c). 8. 7. 7. 45.
6. Catarroja, Valencia. 0. Thomas (p). 8. 2. 9. 208.

(N. Gonzalez.)
6 <5, 4 9, 2 juv. Seville. {A. Ruiz.) Lord Lilford (p). 95. 3. 3. 29-40.

2 al. Jerez de la Frontera, Abel Chapman (c 0. 5. 10. 2-3.

Cadiz. & p).

2 al. Jerez de la Frontera, A. Trevor Battye 7. 9. 12. 1-2.

Cadiz. (c & p).

1. Jerez de la Frontera, Abel Chapman (c 8. 3. 26. 6.

Cadiz. & p).

2. Coto Donana, Huelva. Abel Chapman (c 8. 3. 8. 10-11.

&P).

PiTYMYS IBERICUS PAscuus Miller.

1908. Pitymys ibericus fuscus Miller, Ann. and Mag. Nat. Hist., 8th ser.,
I, p. 206, February, 1908. 'Not Arvicola agrcstis fusca (= Pitymys
subterraneus) Fatio, 1900. Type in British Museum.

1911. Pitymys ibericus pascuus MiUer, Proc. Biol. Soc, Washington, xxiv,
p. 39, February 24, 1911 (Substitute ior fuscus).

Type locality. — Dehesa de Valencia, Province of Valencia,
Spain.

Geographical distribution.- — Apparently confined to the Dehesa
de Valencia.

Diagnosis. — Size as in Pitymys ibericus centralis, but colour
dark, resembling 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 oKscurely bicolor, whitish except median dorsal
region, which is tinged to a varying degree with dark brown.

784 RODENTIA

Skull and teeth. — As in P. ihericus 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 regulus Miller, Ann. and Mag. Nat. Hist., 8th ser.,

I, p. 206, February, 1908. Type in British Museum.

1909. [Piiymys] ilbericus] regnlaris Lydekker, Zoological Record, xlv

(1908), Mamm., p. 75, December, 1909. (Accidental renaming of
regulus.)

1910. Pitymys ihericus regulus Trouessart, Faune jMamm. d'Europe, p. 191.

Tijije 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 form.s
of Pitymys ihericus (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. ihericus 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 cxa?nined. — Eight, from the following localities in Spain :
Granada, 7 ; Malaga, 1.

2 6,i9. Granada, Spain. G. S. Miller (c). 8. 8. 4. 288-293.

(8. 8. 4. 290. Type of subspecies.)
1 al. INIalaga. J. G. Haggard (p). 8. 4. 28. 1.

PITYMYS DUODECIMCOSTATUS de Selys-Longchamps.

1839. Arvicola duodecimcostatus de Selys-Longchamps, Revue Zoologique,

p. 8, January, 1839.
1905. Microins duodecijncostatus iNIajor, Ann. and Mag. Nat. Hist., 7th ser.,

XV, p. 508.
1908. Pitymys duodecimcostatus Miller, Anu. 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 g6n6rosit6 do 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 distrihidion. — Mediterranean coast region of
France ; northward to western Switzerland 1 Limits of range
unknown.

Diagnosis. — Similar to Pitymys ihericiis, 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. ihericus centralis.

SJciill and teetJi. — Except for the peculiarities already noted
the skull so closely resembles that of Pitymys ihericus as to need
no detailed description. Teeth as in P. ihericus.

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 distrihution. — 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 dnodecinicostatus, except
for its much smaller size ; auditory bulke very small and Hat.

Colour. — Ujaper 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 grepsh.

Skull and teeth. — The skull of Pitymys p)rovincialis 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

* Ktudes de Micromanimalogio, pp. 110, 111.

3 E

786 EODENTIA

above sub-orbiciilar, the sides strongly rounded ofl". Occiput not
so obliquely truncate as in P. duodecimcostatiis, and interparietal
less narrowly ligulate ; in both characters it more closely resembles
P. ihericus. Rostrum very slender, though jserhaps not dispro-
portionately so. Auditory bulla? smaller and more flattened than
in any other member of the group except P. depressus of central
Spain, their contrast with the large bullte of P. diiodecimcostatus
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.

Mcasvrcnients. — External measurements of type (adult female) :
head and body, 9G ; tail, 22; hind foot, 14-6. For cranial
measurements see Table, p. 790.

Specimens 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 (5, 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 Selys-Long- 45. 7. 5. 9.

champs (p).

2. Var Purchased (Parzudaki). 52. 5. 27. 51-52.

PITYMYS THOMASI Barrett-Hamilton.

1903. Microtus [Pitymys) ihomasi Barrett-Hamilton, Ann. and Mag. Nat.

Hist., 7th ser., xi, p. 306, March, 1903- Type in British Museum.
1905. Microtus {Pitymys) tlwmasi Major, Ann. and Mag. Nat. Hist.,

7th ser., xv, p. '515, May, 1905.
1910. Pitymys tlwmasi Trouessart, Faune Mamm. d'Europe, p. 188.

Type locality. — Vranici, Montenegro.

GeograpMral distribidioji. — 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 ai'ea bi'ownish, the rest nearly white.

SJiull. — 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 bull;e appear to be relatively larger.

Teeth. — As in P. ihericus ihericus.

MeasHrements. — Type (adult female): head and body, 118;
tail, 22; hind foot (dry), 17-4; ear, 10. Adult male: head

787

and body, 107; tail, 23; hind foot, 17; ear, 10. For cranial
measurements see Table, p. 790.

Specimens examitied. — Four, from the following localities in llonte-
negro : Beri, 1 ; Daljans, 1 ; Vranici, 2.

BemarJcs. — Among the members of the group to which it
belongs this species is easily recognizable by its dark colour and
large size.

6. Beri, Montenegro. (L. Filhrer.
9. Dcaljans. {L. Filhrer.)
6, ?. Vranici. (L. Fiihrer.)

0. Thomas (p). 6. 8. 4. 23.
0. Thomas (p). 5. 8. 4. 25.
0. Thomas (p). 5. 8. 4. 24, 26.
(5. 8. 4. 26. Type of species.)

PITYMYS ATTICUS Miller.

1910. Pitymys 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 distribution. — Known only from the type locality.

Diagnosis. — Size much smaller than in the only other known
form occurring in the Balkan Peninsula, P. thomasi, and about
equal to that of P. duodecimcostatus. Differs from the latter in
less projecting upper incisors and less obliquely truncate occiput.

Colour. — tipper parts bufFy 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
bufiy white of underparts is rather abrupt. Basal portion of
fur everywhere slate-colour, tliis appearing at surface throughout
underparts. Feet bufly white in strong contrast with back.
Tail obscurely bicolor, pale cream-bufF below, huffy 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 rostrum and nasals, relatively wider inter-
orbital region, less strongly projecting incisors and less obliquely
truncate occiput. Auditory bullte flatter and less inflated than
in P. duodecimcostatus, though of about the same diameter (not
reduced as in P. provincialis).

Te^eth. — The teeth are essentially like those of Pitymys duode-
cimcostatus and P. thomasi, 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 liave compared it.

Measurements. — Type (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.Mottas.) Hon. N. C. Rothschild (p). 8.10.2.51.

( Type of species.)

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791

Suu-Famii.v MURINv^.

1857. Miirinx Baird, Mamm. N. Amor., p. 434 (part).

Gcograpltical 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.

Heniarks. — The sub-family Murinae contains about fifty
described genera, five of which occur in Europe.

KEY TO THE EUROPEAN GENERA OF MURINE.

Mesopterygoid fossa reduced to a triangular space
scarcely extending in front of hamulars ; coronoid
process of mandible obsolete ; back densely and

conspicuously spinous (Crete) Acomys, 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 ou

inner side.

Upper incisor with outer side of cutting edge entire;

first upper molar 5-rooted, its crown not so

long as that of m- and nv^ combined ; size

large (House-rats) Epimijs, 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 m^ combined ; size small (House-mice) Mas, 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

notioeably greater than depth at front of m'

(Field-mice) Apodemus, p. 791.

Tail prehensile, naked at tip above ; ear with con-
spicuous lobe closing meatus ; rostral XJortion
of skull so shortened that length of diastema is
less than depth at front of «j' (Harvest-mice)... Micromys, p. 840.

Genus APODEMUS Kaup.

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

I, p. 150 (agrarius).
1857. Mies Blasius, Siiugethiere Deutschlands, p. 309 (part).
1905. Micromys Thomas, Ann. and Mag. Nat. Hist., 7th ser., xv, p. 492,

May, 1905 (part).
1908. Apodemus Thomas, Ann. and Mag. Nat. Hist., 8th ser., i, p. 447,

May, 1908 (part).

Type species. — Mus agrarius Pallas.

Geographical distribution. — Northern temperate portion of

'^^'^ RorjENTIA

Old World from Ireland to Japan, south to northern India and
the Mediterranean region of Africa.

Chararicrs. — ykull without special modifications of form, the
rostral portion well developed (diastema noticeably exceeding
depth of skull at anterior root of »*'), the mesopterygoid fossa
scjuarish or broadly double- or single-rounded anteriorly ; first
and second upper molars with thi'ee cusps on inner- side ; ear
without valve for closing meatus ; tail not prehensile ; fur
without spines.

BomarJiS. — Among Palsearctic Muridse the genus Apodcmus 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 Thnmnomys.
About a dozen si)ecies are known, seven of which occur in
Eurojje.

KEY TO THE EUROPEAN FORMS OF APODEMUS.

First lamina of m' with crescentic form much dis-
torted; m- with no antero-external tubercle;
interorbital region with conspicuously beaded
edges; back with blacu median line (Eastern

Europe, west to Denmark) A. agrarius, p. 834.

First lamina of wt' with crescentic form not dis-
torted ; m- with small but distinct antero-
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 alwve but!y grey (Balkan Peninsula)... A. cpimclas, 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 imn. (25-29 mm.) A. flavkoUiK. p. 828.
Underparts usually creamy white, the pectoral
spot usually not forming complete collar
and not spreading bacliward over chest
(Continental Europe north of Mediter-
ranean region) A.f.flancoliis,-p. 829.

Underparts usually greyish white, the pectoral
spot usually forming complete collar and
spreading backward over chest (England) .1..^. wlattml, p. 831.

apodemus 793

Skull romaining light and smooth, cvon 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 with
noticeably worn teeth most frequently
23-24 mm. (21-26 mm.).*
General form of skull slender, especially in
rostral portion ; mandible with coronoid
process reduced; colour dusky above,
dull white below, the pectoral spot obso-
lete ; size maximum for this section, the
hind foot and oondylobasal length of
skull sometimes exceeding 20 mm. (Fair

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.

Oondylobasal length of skull 24 to 25 mm.

(Hebrides, Scotland) A. hebridcnsis, p. 824.

Oondylobasal length of skull 25-6 to

27 mm. (St. Kilda Island, Scotland) .4. hirtensis, p. 825.
Underparts usually whitish, occasionally
washed with yellowish brown ; ear

.iiormal A. sylvaticus, p. 797.

Size small, hind foot most frequently

22 mm., coudylobasal 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., oondylobasal 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. callipides, p. 809.

General colour above pale yellowish

or greyish brown without russet

(Mediterranean region in general) A. s. dichrurus, p. 810.

* These dimensions exceeded in the local form inhabiting Fair Isle,
Scotland.

»

794

APODEMUS EPIMELAS Nehrill

1882, Mus mystaciiius Winge, Vidcnsk. Jleddel. fra den naturh. Foreu. i
Kjpbenhavn, 4th ser., iii (1881), p. 21 (Dekelia, near Athens,
Greece). Not Miis mystacinits Dan ford and Alston.

1902. Mus epimelas Nehring, Sitz.-Ber. Gesellsch. Naturforsch. Freunde,
Berlin, p. 95, February, 1902.

1910. Mus (Epimijs) epimelas Trouessart, Faune Mamm. d'Europe, p. 144.

Type localifi). — Agoriani, Parnassus, Greece.

Geographical dislrihition. — 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 syli-aticus except for their
greater size, but molars relatively larger, the alveolar width of
m^ equal to about one-half width of palate, m^ and m'^ with
minute though evident fourth tubercle at postero-external
margin of crown, and outer ledge of m^ and in., somewhat better
developed.

External characters. — As in Apodemus 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
sui'face 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, ajjart from its conspicuously
greater size, the skull agrees with that of Apiodemus 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. flavicollis. Anterior
border of incisive foramina extending relatively further for-

APODEMUS

795

ward than in A. syhatims, the distance from foramen to alveolus
ot nicisor barely more than half combined width of the
two toraniina instead of neai'ly equal to it.

Fig. 163.
Apodemus epimelas. Nat. size.

Teeth—Thongh m general resembling those of Apodemus
sylmticus, the teeth of A. epimelas differ in several important
details. The crowns of all the cheek-teeth are perceptibly

riG. 164.

Apodemus epimelas. Cheek-teeth. X 10.

longer proportionately to their breadth than in the smaller
animal, but at the same time the distance between the upper
tootn-rows is so reduced that the alveolar breadth of m^ is fully

796

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APODKMUS

r97

to one-half width of palate in same region. Pattern of
enamel folding essentially as in Ajmdemus 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 m^ better developed than in
A. sylvaticus, those of the second tooth nearly as large as those of
first, a condition rarelj^ 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.

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

Montenegro : No exact locality, 6.

Albania : Northern Albania, S.

Greece : Corfu, 3 ; Cephalonia, G ; near Athens, 12.

BevLarks. — While Apodemus eijimelas is very unlike the other
European members of the genus it closely resembles A. mystacinus
Danford and Alston from Asia Minor, diflering 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 (?, 3 ?. Montenegro. (L.Filhrer.) O. Thomas (p). 5. 8. 4. 10-15.

4 (i, 4 9. North Albania. 0. Thomas (p). 5. 8. 4. 27-34.

{L. Filhrer.)
2 6. Corfu, Greece. (C.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, ?. Tatoi, Athens. (C. Mottaz.) Hon. N. C. Rothschild 8. 10. 2. 38-39.

APODEMUS SYLVATICUS Linnieus.

(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 vertebra?, about 8 0 to 110 ; hind foot, 20 to 25,
most frequently 22 or 23 ; condylobasal length oi' 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-bi'own 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 jiresent 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 tliird 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 rharacters. — External form slender, much as in the
house mouse, but feet, eyes and ears relatively 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 sliowing 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 outei' 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
across), its surface papillose in region among tubercles, rugose at
heel, smooth between last tubercle and wrinkled posterior
pdrtion ; 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.

APODEM0S

799

somewhat quadrate in section, the annulations distinct, about
eighteen to the centimeter at middle ; hairs rather sparse, not
conceaUng the annulations at middle but forming a small though
evident pencil at tip, their length equal to width of 2^ rings.
Mammas : p 1 — 1, i 2 — 2 = 6.

Colour. — For detailed description of colour see accounts of
subspecies.

SlmJl. — The skull is moderately large, smoothly rounded, and
without any striking characteristics. Dorsal surface straight
and gradually I'ising from tip of nasals to middle of frontals,
then moderately 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 ;«• ; palate nearly flat, without
special features, the pcjrtion formed by jialatine bones extending
forward to middle of ui^ ; mesopterygoid space narrow anteriorly,

I'lO.. 163.
Apodemiis si/lvaHcits. Nat. size.

800 EODENTIA

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 bulltB ; 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 rather slender, its
inner surface with noticeable longitudinal concavity ; dental
foramen at level of alveoli ; projection marking root of incisor
slight but evident, directly under base of coronoid process.

Teeth. — Relatively to size of skull the teeth are rather small :
alveolar breadth of m^ 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.

Apudennix Kiilraticas

Fig. im.
I'lieek-teetli, (a) slightly worn ; (h) much worn.

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 ditierentiated 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 with tubei'cles ari-anged acc<jrding t<j a ground-
plan of three longitudinal rows of tubercles, those of the median

APODEMUS 801

row much lar,ii;er than those of the inner and outer row.s, the
tubei'cles of the different rows partly joined by cross-ridges, and so
disposed as to form three transverse laminaj having somewhat the
form of forwardly-bowed crescents. In each tooth this complete
pattern is only approximated ; in m^ it is imperfect posteriorly,
in ni' both anteriorly and posteriorly, while in m^ it is scarcely
more than recognizable. For convenience of description the
tubei'cles 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 vi^ 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 of <1 and the very
broad, shallow angle between its inner border and the outer
border of <2 ; a small but evident projecting supplemental loop
usually present in angle between /3 and fl, a similar but less
developed loop occasionally present in that between t2 and il ;
second crescent essentially similar to first, and partly or com-
pletely joined to it by a narrow ridge extending between t'd 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 t9 and tl so that the three cusps of
this lamina lie in the same transverse line, and the re-entrant
angles separating them from /6 and /4 respectively are much
narrower than those behind iirst lamina ; t9 is broadly connected
with /6 anteriorly, so that the two might almost be interpreted
as parts of a deeply divided /6 ; t7 is similarly 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
Apodenius epimelas. Second upper molar sub-circular in outline,
its first lamina incomplete by the absence of <2 ; <3 very small,
terete ; second and third laminie essentially as in m^ ; postero-
external loop excessively minute, though usually present in unworn
teeth. Third upper molar similar to nr 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 <5. Lower molars consisting of a
series of paired, equal, nearly distinct tubercles, each tooth
terminating }iosteriorly in a single median tubercle usually much

3 F

802 EODENTIA

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 m^ the
most constant supijlemental 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 readily 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 Apodenms
sylvaticus is likely to be confused is A. JlavicolUs. Detailed
comparisons of the two animals will be found undei- 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
sylvaticMs 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 Apodemus sylvaticus
much less numerous than Mus spicilegus, and practically coniined to
ravines and river banks, while the smaller animal was abundant every-
where.

APODEMUS

803

ail area of iiitergradatiou between true nyJraticus ami the larger
Mediterranean form. Specimens from this region are indeter-
minate in characters, though 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 ajjpears 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 sufficiently distinct to admit of characterization. On
the other hand, the Hebrides, St. Kilda, and Fair Isle, oft' the
coast of Scotland, are each inhabited by a peciiliar, well-
difierentiated 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
sylvaficus. It seems probable, therefore, that they have been
recently introduced.

Apodemus sylvaticus sylvaticus Linnaeus.

1758. [AIus] sylvaticus Linnseus, Syst. Nat., i, 10th ed., p. 62 (Upsala,
Sweden).

1796. Mus sylvaticus parvus Bechstein, Getreue Abbild. Naturhist. Gegen-
stande, i, p. 100 (Thiiringen, Germany).

1796. Mus sylvaticus candidus Bechstein, Getreue Abbild. Naturhist.
Gegenstande, i, p. 100 (Thiiringen, Germany).

1796. Mus sylvaticus varius Bechstein, Getreue Abbild. Naturhist. Gegen-
stande, I, p. 100 (Thiiringen, Germany).

1796. Mus sylvaticus nigcr Bechstein, Getreue Abbild. Naturhist. Gegen-
stande, I, p. 100 (Thiiringen, Germany).

1801. Mus syl\yaticus'] albus Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2d ed., p. 965 (Thiiringen, Germany).

1801. Mus syllyaticus'] leucoccphalus Bechstein, Gemeinn. Naturgesch.
Deutschlands, i, 2d ed., p. 966 (Thiiringen, Germany).

1S21. Mus islandicus Thienemann, Reise im Norden Europas, ii, p. 153
(Iceland).

1834. Mus sylvaticus Melchior, Den Danske Stats og Norges Pattedvr,
p. 102.

1839. M21S 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 cdticxis Barrett-Hamilton, Proc. Zool. Soc, London,
p. 401 (Caragh Lake, Co. Kerry, Ireland).

1910. Mus sylvaticus intermedius and M. sylvaticus cclticus Trouessart,
Faune Mamm. d'Europe, pp. 149, 150, 151.

1910. [Apodemus'] sylvaticus Miller, Ann. and Mag. Nat. Hist., 8th ser.,
VI, p. 460, November, 1910.

'--' Type locality. — Upsala, Sweden.

Geograjjhical distrihufion. — Central Europe from Ireland

3 p 2

804 RODENTIA

eastward, and from central Sweden and Norway south to
northern Italy and the southern France ; Iceland (probably
introduced).

Dlu(jnosis. — vSize small ; head and body, about 95 ; tail
vertebras, 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 bufly on head, neck,
shoulders and sides ; everywhere there is a noticeable sprinkling
of blackish hairs, these usually ^^roducing 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 line
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 Mus muscidus group, though they may always be
distinguished by their relatively larger eyes and ears, relatively
longer hind foot, and by the absence of sujjplementary 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

Al'ODEMUS

805

adults from Bergen, Norway: head and body, 96-1 (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
el 5-1 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 (9.3-99); 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.).

Irkland : Clandeboye, 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, Tixsperary, 2 ; Ballagh-
moon, Carlow, 6; Cappagh, Wexford, 1; Duncannon, Wexford, 3;
Kilmauock, Wexford, 2 ; Caragh Lake, Kerry, 6 (including tj'pe of cclticus
Barrett-Hamilton) ; Castlegregory, Kerry, 4 ; Glencar, Leitrim, 1.

Scotland : Islay, 1 ; Skye, 1 ; Duurossness, Shetland, 3 ; South Sutor,
Cromartv, 7 ; Black Isle, Cromarty, 4 ; Craigellachie, Elgin, 1 ; Lossie-
mouth, Elgin, 2; Lhanbride, Elgin, 2 ; Dunphail, 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, StafEordshire, 3 ; Swincliffe, Shrop-
shire, 1 ; Swithland, Leicestershire, 12 ; Belvoir, Leicestershire, 2 ; Anglesey,
1 ; Aberia, ^Merionethshire, 2 ; Truoyn-y-peurleyn, Merionethshire, 2 ;
Penrhyndeudraeth, Merionethshire, 1; Laugharne, Carmarthenshire, 4;
Merlins Hill, Pembroke, 2 ; Southerndown near Bridgend, Glamorgan-
shire, 10 ; Bishopstone, Herefordshire, 3 ; Graftonbury, Herefordshire,
14; Ledbury, Herefordshire, 1 ; Darsley, 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 ; Colnc, Cambridgeshire, 1 ; Histon, Cambridgeshire, 1 ; Fulbourn,
Cambridgeshire, 3; Levesham, Suffolk, 2 (Wilson); Lowestoft, Suffolk, 4;
Haileybury, Hertfordshire, 1 ; Tring, Hertfordshire, 1 ; Hillingdon,
Middlesex, 1; Chiswick, Middlesex, 1; Regent's Park, London, 2; Ban-
stead, Surrey, 18 ; Wandsworth Common, Surrey, 4 ; Betchworth,
Surrey, 1; Yalding, Kent, 4; Crowborough, Sussex, 3; Dobbin Island,
Pagham Harbour, Sussex, 1 ; Alum Bay, Isle of W'ight, 4 ; Bembridge,

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 ; Combmartin, 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, Li^ge, 4; Waremme, Li6ge, 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 ;
Foret de Bouconne, Gers, 14 ; Montr6jeau, 11 (U.S.N.M.) ; Biarritz,
Basses-Pyrdn6es, 7 ; St. Paul, near Barcelonnette, Basses- Alpes, 27 ;
Chamonix, Haute-Savoie, 6 ((J.S.N.M.); Cranves-Sales, Haute-Savoie, 8 ;
Luciuges, Haute-Savoie, 6 ; Montauban, Haute-Savoie, 3 ; Etupes,
Doubs, 4 ; Manonville, Meurthe-et-Moselle, 2.

Germany : Brunswick, 34 (U.S.N.M.) ; Hausbruch, Hanover, 2 ;
Bodethal, Harz Mountains, 4 (U.S.N.M.) ; Bahrenberg, Harz Mountains, 4
(U.S.N.M.) ; Wolfshau, Riesengebirge, Silesia, 22 (U.S.N.M.) ; Eulengrund,
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-HuNCiARY : 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.

Switzerland : 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. GaUon, 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) ; Faido, Ticino, 3 (B.M. and U.S.N.M.) ; Lugano,
Ticino, 6 (U.S.N. INI.) ; Capolago, Ticino, 1 ; Locarno, Ticino, 7.

Italy : Padola, Cadore, 1 (Turin).

2 J, 1 9. Clandeboye, Down, Ire- W. R. Ogilvie-Grant 11. 1. 3. 318-320.

land. (c & p).

1 al. Meath. Dr. G. E. Dobson (p). SO. 12. 14. 5.

3 (?, 3?. Clare Island, Mayo. G.Barrett-Hamilton 11.1.2.132-137.

{R. H. Bunting.) (p).

2 <?, 2 ?. Inishmore, Aran Islands. G. Barrett-Hamilton 11. 1. 2. 138-141.

{R. H. Bunting.) (p).

3 6. Scarif!, Galway. G. Barrett-Hamilton 11. 1. 2. 68-70.

(R. F. Hibbcrt.) (p).

i. Clare. (R. F. Hibbert.) G.Barrett-Hamilton 11.1.2.71.

6. Nenagh, Tipperary. Capt. W. F. Smith 11. 1. 3. 321.

(c & P).
S.. Cashel, Tipi)orary. W. E. do Winton 11. 1. 3. 322.

(c & p).
2>^,2?. Ballaghmoon, Carlow. G. Barrett-Hamilton 11.1.2.64-67.
(J. G. Si/mex.) (p).

APODEMUS

807

6. Cappagh, Wat'.erforf!.

{W.J. Ussher.]
2 6. Kilmanock, Wexford.

(Council.
?. Kerry.

2 6, ?. Caragh Lake, Kerry.

1 al. Islay, Scotland.

6, 9. Tiree Island.

(P. Ajiderson.)

?. Isle of Skye.

2 6, 9. South Sutor, Cromarty.

6, 9. Black Isle, Cromarty.

2 S. Lossiemouth, Elgin.
9 al. Craigellachie, Elgin.
2 S,2 9. Dunphail, Elgin.

S, 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 JIan.
2 (5, 2 9. Swithlaud, Leicester-
shire, England.

6, 9 al. Belvoir, Leicestershire.

1 al. Swincliffe, Shropshire.

2 6. Aberia, Merionethshire,

Wales.
2 6,2 9. Laugharne, Carmarthen-
shire.

4 S. Bridgend, Glamorgan-

shire.
6, 4 9. Graftonbury, Hereford-
shire, England.

3 al. Bishopstone, Hereford-

shire.
i, Bloxham, Oxfordshire.
2 i,2 9. Wormsley, Oxfordshire.

3 6, 9. Yalding, Kent.

9. Chiswick, Middlesex.
9 6,2 9. New Forest, Hampshire.

3 6. Basingstoke, Hampshire.
3. Clifton, Gloucestershire.

2 6, 9. Clifton
9. Clifton

3 6, 9. Lowestoft, Suffolk.

6. Dobbin Island, Sussex,
England.

[J. E. Hartinq.)
4 6, 2 9. Scilly Islands, Cornwall,
England.
9. Brent Knoll, Somerset.
3 <5, 9. Opheim, Bergen, Norway.

G. Barrett-Hamilton
G. Barrett-Hamilton

Col. J. W. Yerbury

(c & p).

{Tiipe of cclticus
Col. J. W. Yerbury

(c & p).
H. Eussell (c & p).
Lord Elphinstone (p).

J. S. Elliott (c & p).
W. E. 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).

11. 1. 2. 72.

11. 1. 2. 62-63.

0. 3. 11. 1.

Barr.-Ham.)
11. 1. 3. 323-325.

9. 9. 11. 4.
8. 3. 16. 1-2.

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.

E. R. Alston (c & p). 79. 9. 25. 60.

C. H. B. Grant (c & p). 11. 1. 3. 346-348.

C. H. B. Grant (c & p). 11. 1 . 3. 349-351.

F. A. Butler (c & p). 11. 1. 3. 361-364.

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. Ridlev

(c & p).
O. V. Aplin (c & p).
W. R. Ogilvie-Grant

(c & p).
W. R. Ogilvie-Grant

(c & p).
Miss Sharpe (c & p).
Sliller Collection.

Miller Collection.
Miller Collection.
R. I. Pocock (c & p).
R. Powell (p).
O. Thomas (c & p).
E. R. Alston (p).

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. 357-360.
79. 9. 25. 41.

F. J. Cox (c & p). 11. 1. 3. 336-341.

A. B. Pcrcival (i^). 3. 12. 17. 1.

Miller Collection. 7.7.7.4464-4467.

808

RODENTIA

1. Aker, Christiania, ISTor- Christiania Museum 93. 3. 1. 10.

way. (e).

?. Upsala, Sweden. Lord Lilford (p). 11. 1. 1. 163.

{Eolthoff.)
2 9. HillerOd, Zealand. Den- 0. Thomas (c & p). 98. G. 7. 11-12.

mark.
6, ?. Oosterbeek, Guelderland, 0. Thomas (c & p). 98. 2. 1. 16-17.
Holland.

5, 2 ?. Noordwijk, Leiden, Hoi- 0. Thomas (c & p). 98. 2. 1. 18-20.

land.
i, 8 ?. Esueux, Li6ge, Belgium. Lord Lilford (p). 95. 1. 1. 12-20.

{H. GrUnvold.)
i, ?. Guines, Pas - de - Calais, 0. Thomas (c & p). 94. 6. 6. 14-15.

2 <5, 2 9. Boulogne', Pas-de-Calais. O. 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(5. Alderney, Channel Is- G.Barrett-Hamilton 11.1.2.73-74.

lands. [W. Eagle Clarke.) (p).
2(5. 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, Brittanv. G. Barrett-Hamilton 11. 1. 2. 115-122.

9. Huelgoat, Brittany. O. Thomas (c & p). 92. 9. 5. 2.

6 <5, 3 9. Solferino, Landes. O. Thomas (p). 6. 4. 1. 66-74.

(A. Robert.)
E) 6,2 9. Foret deBouconne, Gers, O. Thomas (p). 6. 4. 1. 59-65.

250 m. (A. Robert.)
2 (5, 2 9. Cranves - Sales, Haute- A. Robert (c & v). 5. 4. 4. 2-5.

Savoie, 900 m.
3(5. Cranves - Sales, Haute- 0. Thomas (p). 5.11.18.11-13.

Savoie. (-.4. Robert.)
8 (5, 8 9. Barcelonnette, Basses- O. Thomas (p). 8. 8. 10. 74-89.

Alpes. (C. Mottaz.)
i. 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-Pyrenees. J. F. Davison (c & v). G. 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. Bacr.)
2. i\Iagdeburg, Saxony. Dr. W. Wolterstorff 92. 12. 1. 8-9.

(c & p).
c?. Magdeburg, Saxony. Lord Lilford (p). 11. 1. 1. 158.

{Wolterstorlf.)
2 9. INlarxheim, Bavaria. ' Lord Lilford (p). 11. 1. 1. 161-162.

( Wolterstorff:)
6, 9. Scrass. Burgheim, Ba- Lord Lilford (p). 11. 1. 1. 159 160.

varia. (Korbitz.)
2 <5, 9. Schwarzburg. Lord Lilford (p). 95. 4. 18. 12-14.

(IF. Schllltcr.)
6. Kalbe, Germany. Lord Lilford (p). 11. 1. 1. 157.

( Wolterstorff.)
2 9. Ingelheim, Rheinhessen. C. Hilgert (c). 8.11.2.44-45

4 <5, 2 9. Strassburg, Alsace. O. Thomas (p). 8. 8. 10. 139-144.

(C. Mottaz.)

2. C s a 1 1 6 k (■) z - Somorja, Budapest ]Museum(E). 94.3.1.46-17.

Prossburg, Hungary.

APODKIIUS

809

6, 2 ?. Hatszeg, Hunyad, Tran- C. G. Danford (c). 3. 2. 2. 25, 30, 32.

sylvania.

1 al. Transylvauia. C. G. Danford (p). 80. 10. 28. 1.

4 al. Eszek, Slavonia. Budapest ]\Iuseum(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 6, ?. Wolfhalden, Switzer- Miller Collection. 7. 7. 7. 10-14.
land. (Zollikofer.)

9. Capolago, Ticino. O. Thomas (c & p). 2. 7. 1. 6.

3 <5, 3 9. Locarno, Ticino. 0. Thomas (c & p). 2. 7. 1. 7-12.

6. Faido, Ticino. O. Thomas (c & p). 2. 7. 1. 19.

3 6. Lucerne, Switzerlaad. Lord Lilford (p). 11. 1. 1. 154-156.
(Woltersto)f.)

Apodkmus sylvaticus callipides Cabrera.

1907. Micromys sijlvaticiis callipides Cahrera, Bol. Real Soc. Espau. Hist.

Nat., Madrid, vii, p. 227, November, 1907 (Villarrutis, La Corufia,

Spain). Type in Madrid Museum.
1910. Mas sylvaticus callipides Trouessart, Faune Mamm. d'Europe, p. 153.

Type locality. — Villarrutis, La CoruiJa, Spain.

Geoc/raphical distrihution. — Mountains of the Pyrenees-
Asturias chain ; southward into Portugal.

Diagnosis. — Size distinctly larger than in Apodcinns sylvaticus
sylvaticus, 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.

Measureinents. — 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,
lUo ; 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.

Specimejis examined. — One hundred and twelve, from the following
localities : —

France : Port6, Pyren^es-Orientales, 9 ; I'Hospitalet, Ariege, 8 ; Ax-
les-Thermes, Ariege, 31 ; Lucbon, Haute-Garonne, 11 ; Caterille, Haute-
Garonne, 5; Bareges, Hautes-Pyrenees, 19; Pic-du-Midi, Hautes-Pyr6n6es, 1.
Spain : Pajares, Leon, 10; La Corufia, 2 ; Villalba, Lugo, 5.
Portugal : Cintra, 7 ; Estoril,- 2 ; Oporto, 1.
4 6. Port6, Pyrenees-Orientales, G. S. Miller (c). 8. 8. 4. 200-203.

France.
c5, 9. Porte, Pyr6n6es-Orientales; 0. Thomas (p). 8.9.1.57-57.
16-1700 m. [A. PMbcrt.)

SIO

RODENTIA

5 6, 9. L'Hospitalet, Ariege. G. S. Miller (c). 8. 8. 4. 204-209.

6, ?. L'Hospitalet, Ariege. O. Thomas (p). 8. 9. 1. 54-55.

(.4. Robert.)

4(5,4 9. Ax-les-Thermes, Ariege. , G. S. Miller (c). 8.8.4.213-220.

5 <5, 3 ?. Luchon, Haute-Garonne. 0. Thomas (p). 6. 4. 1. 40-47.

(.4. Robert.)

8 6, 2 9. Caterille, Haute-Garonne. 0. Thomas (p). 6. 4. 1. 54-58.

(.4. Robert.)

3 J, 4 9. Bareges, Hautes- Pyrenees. G. S. MiUer (c). 8. 8. 4. 183-189.

6. Pic-du-Midi, Hautes-Pyre- 0. Thomas (p). 8. 9. 1. 58.

nees. (.4. 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. 10-18.

(c & p). 95. 4. 29. 2.

2(5,2 9. Cintra, Portugal. O.Thomas (c &p). 98. 2. 2. 22-25.

S, 9, 1. Cintra. {K. Jordan.) Hon. N. C. Roths- 10. 9. 28. 1-3.

child (p).

2 S. Estoril. O.Thomas (c (fc 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, Pr6cis des Decouvertes Somio-

logiques, p. 13 (Sicily).
1844. Mns pecchioli Pecchioli, Atti della quinta Unione degli Scienziati

Italian! , Torino, 184.3, p. 426 (Tuscany, probably vicinity of

Siena, Italy).
1900. Mus sylvaticus hayi Barrett-Hamilton, Proc. Zool. Soc, London,

p. 410 (part).
1910. Mus sylvaticus hayi and M. sylvaticus dichrurus Trouessart, Faune

Mamm. d'Europe, p. 154.

Type locality. — Sicily.

Geographical distribution. — Mediterranean region from the
Balkan Peninsula to central and southern Spain.

Diagnosis. — Size as in Ajwdemus 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 dull buffy wood-brown, slightly
darker along median posterior region, and slightly more greyish
anteriorly, the black hair tips, though numerous, nowhere
sufficiently abundant to produce 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 biiffy grey, intermediate between the
ecru-drab and cream-buff of Ridgway, but paler than either, the
"lining ' jiroduccd by tlie black hairs conspicuous. Yellow

APODEMUS

811

extreme : ground colour a clear buff slightly more yellowisli 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 extreme.

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.

Nl:

RODENTIA

Castiillo de la Reina, Burgos, 1; Panticosa, Huesca, 4; Jaca, Huesca, 1;
Jerez, Cadiz, 1; Inca, Majorca, Balearic Islands, 12; San Cristobal,
Minorca, Balearic Islands, 6.

Prance : Cerbere, Pyr^n^es-Orientales, 1 ; near Nimes, Gard, 21 ;
Valescure, Var, 11 ; Agay, Var, 1.

Italy: Vicinity of Turin, 19 (Turin); Moucalieri, Turin, 6 (B.M.,
U.S.N.M. and Turin) ; Genoa, 11 (U.S.N.M.) ; Viareggio, 4 ; Sorrento, 12
(B.M. and U.S.N.M.); Aspromonte, Lagouegro, 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.

CIbeece : Corfu, 9 ; Cephalonia, 13 ; Patras, 1 ; near Athens, 15 ;
Agorianni, 5 (U.S.N.M.).

4 (5 1 9. Guadix. Granada, Spain. G. S. Miller (c). 8. 8. 4. 75-79.

4(5 3 9. Venta del Baul, Granada. G. S. Miller (c). 8.8.4.80-86.

2i. Elche, Alicante. G. S. Miller (c). 8.8.4.87-88.

(5, 2 9. Alcoy, Alicante. O. Thomas (p). 8. 2. 9. 127-129.

(A^. Gonzalez.)

0. Thomas (p).

a.

4 i, 3 9.

7 <i,9.
9.
4(5.

9.

3 (5, 6 9.

4 (5,2 9.
6. juv.

4 (5, 4 9.

4 (5, 3 9.

9.

6, 9.

4.5.

2<5, 2 9.

2<5.

Villalba, Madrid.

(A''. Gonzalez.)
Bejar, Salamanca.

{N. Gonzalez.)
Silos, Burgos.
Castrillo, Burgos.
Panticosa, Huesca.

(N. Gonzalez.)
Jaca, Huesca.

{N. Gonzalez.)
Jerez, Cadiz.
Inca, ^Majorca, Balearic

Islands.
San Cristobal, Minorca.

Cerbere, Pyrenees Ori-

entales, Prance.
Nimes, Gard.

{C. Mottaz.)
Valescure, Var.
Agay, Var.

Moucalieri, Turin, Italy.
Viareggio.
Sorrento.

0. Thomas (i>).

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

O. 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 & v). 8. 3. 26. 5.
O. Thomas and K. I. 0. 7. 1. 9-17.

PoCOCk (C & P;.

O. Thomas and R. I. 0. 7. 1. 52-57.

Pocock (c & p).
O. Thomas and R. I. 0. 7. 1. 75-76.

Pocock (c & p).
O. Thomas (p). 8. 8. 10. 66-73.

Aspromonte, Calabria.
{A. liobert.)
6. Monte Sirino, Calabria.

{A. Robert.)
3 <5, 9. Castelbuono, Sicily.

{A. Robert.)
5 6. Ficuzza, Sicily.

(.4. Robert.)
5 6, 3 9. Madonna dell' Alto,
Petralia, Sicily.

(A. Robert.)
2 6, 2 9. Marsala, Sicily.

(.4. Robert.)

3 (5, 9. San Giuglielmo, Sicily.

{A. Robert.)

G. S. ]\Iiller (c).
G. S. IMiller (c).
Dr. E. Festa (p).
0. Thomas (c &. p).
R. J. Cuninghame

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

0. Thomas (p).

O. Thomas (p).

0. Thomas (p).

0. Thomas (p).

0. Thomas (p).
O. Thomas fp).

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.

/6. 8. 4. 42-43.
\8. 9. 1. 24-25.
1 6. 8. 4. 39-41.
18. 9. 1. 22-23.
8. 9. 1. 10-17.

fi. 8. 4. 35-38.
8. 9. 1. 18-21.

APODEMUS

813

6, 9. Asco, Corsica. O. Thomas (i-). 8. 11. 30. 7-8.
{Br. F. Mujur.)

2. La Foce di Vizzavona, Col. J. W. Yerbury 93. 9. 15. 4-5.

Corsica. (p).

3 al. Corsica. Col. J. W. Yerbury 98. 9. 15. 6-8.

2(5. Montenegro. (L.^i^/Z/f/w.) O. Thomas (p). 5.8.4.16-17.

6, ?. North Albania. O. Thomas (p). 5. 8. 4. 36, 37.

(L. Filhrer.)

6. Potamos, Corfu, Greece. J. I. S.Whitaker (p). 8.10.1.10.

6. Cephalonia. C. Mottaz (c). 8. 11. 3. 11.

8 <i, 4 ?. Cephalonia. (C. Mottaz.) J. I. S. Whitaker (p). 8. 10. 1. 11-22.

6. Itaci, Athens. Hon. N. C. Boths- 8. 10. 2. 26.

(C. Mottaz.) child (p).

4 ?. Tatoi, Athens. Hon. N. C. Roths- 8. 10. 2. 33-36.

(C. Mottaz.) child (p).
9. Kephissia, Athens. Hon. N. C. Roths- 8. 10. 2. 37.

(C. Mottaz.) child (p).

Apodemus sylvaticus creticus Miller.

1910. Apodemus sylvaticus creticus Miller, Ann. and Mag. Nat. Hist.,
8th ser., vi, p. 460, November, 1910.

Type locality. — Katharo, Crete.

GeograpJiical distribution. — ^Island of Crete.

Diagnosis. — Size small, e.ssentially as in true A. sylvaticus, 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. sylcaticus 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).

Remarks. — The Cretan form of Apodemus sylcaticus combines
the small size of the typical race with the dull yellowish colour
■of A. sylvaticus dichrurus.

2 (?, 2 9. Katharo, Crete. • Miss D. Bate (c). 5. 12. 2. 25-28.

(5. 12. 2. 27. T///JC of subspecies.)
6, 9. Kanea. Miss D. Bate (c). 5. 12. 2. 29-30.

814

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, much worn.

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6. 8. 4. 6
G. 8. 4. 7
6. 8. 4. 8
153345
153347

8. 9. 1. 11

153344
8. 9. 1. 13
8. 9. 1. 16
8. 9. 1. 17
8. 9. 1. 15
153339
153340
6. 8. 4. 39
6. 8. 4. 40

Locality,

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Monte Sirino
Sicily : Madonna dell

Ficuzza

APODEMUS

823

ad a'

a d

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*^^4 RODENTIA

APODEMUS HEBRIDENSIS de Wintoil.

1S'J5. Mas licbridcnsis de Winton, The Zoologist, 3rcl scr., xix, x'- 369,
October, 1905. Type in British Museum.

1895. Mus sylvaticus licbridensis de Wiuton, The Zoologist, 3rd ser., xix,
p. 426, November, 1905.

1900. Mus sylvaticus hebridensis Barrett-Hamilton, Proc. Zool. Soc.,
London, p. 403.

1910. Mus sylvaticus hebridensis Trouessart, Faune Mamm. d'Europe,
p. 151.

Type Jocalifi/.-^-Vig, Island of Lewis, Outer Hebrides,
Scotland.

Geographical ilistrihution. — Island of Lewis, and Barra Island,
Hebrides.

Diagnosis.— Hize about as in the large southern races of
Apodemus sylvaiicus ; 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 . si/h-aticus ; general
colour of upper parts essentially as in A. si/lraficus syhafictis, but
underparts so heavily washed with dull buffy brown that line of
demarcation along sides is ill-detined ; 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.

Measurements. — 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 e.ramined. — Thirteen, all from the Islands of Lewis and
Barra.

Memarlcs. — "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.

6. Island of Lewis, W. E. de Winton (c & p). 95. 10. 25. 1.

Hebrides. ( Tt/^jc of species.)

i 6, 5 9. Island of Lewis. W. E. de Wiuton (c & p). H. 10. 11. 13-21.

6. Barra Island. Col. E. W. Pinney (c ct p). 11. 1. 3. 314.

6 al. Barra Island. J. McRurie (c & p). 11. 1. 3. 415.

APODEMUS 825

APODEMUS HIRTENSIS Barrett-Hamiltuii.

18'jy. MiiK Jiirtcnsis Barrett-Hamilton, Proc. Zool. Soc, Loudon, p. 81.
Type in British Museum.

1900. Mus sylvaticus hirtenais Barrett-Hamilton, I'roc. Zool. Soc, Loudon,
p. 404.

1906. 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. 152.

Ti/pe locality. — Island of Saint Kilda, west of Outer Hebrides,
Scotland.

Geographical distrihiition. — Saint Kilda.

Diagnosis. — Similar to Apodemus hehridensis, but skull larger
(condylobasal length, 25 '6 to 26 '8) and colour darker.

Measurements. — 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 si3ecifically 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 Hebridean form, though of exactly the same
type. Notwithstanding its large size the skull retains all of the
characteristics of that of A. si/Ivaticus 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 (J, 3?. St. Kilda. W. Eagle Clarke 11.1.24.1-6.

(0 & 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 .4. sylvaticus
syh'dticns, but back and sides less bright and more clouded with

826 RODENTIA

black, and underparts entirely dull whitish, the pectoral spot
absent or barely indicated.

External characters. — In general external characters Aiwdemus
fridariensis agrees with A. si/Ivaticus and A. Jlavicollis, 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 Apodemus siihaticus sijlvnticus, 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. flavkolVn^, and without trace
of the brownish wash so frecjuently indicated in A. sylvaticus,
though a faint buffy tinge is sometimes present ; pectoral spot
usually absent, but 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 Apodonus fridariensis
rather exceeds that of the large Mediterranean forms of
A. sylvaticus, though by no means equalling the maximum of
A. Jlavicollis. 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.]\I., U.S.N.M.
and Edinburgh).

Bemarks. — Notwithstanding its large size this species, like
Apodemus hebridensis and A. hirtensis, appears to be an offshoot of
A. sylvaticus rather than of A. Jlavicollis. 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.)
S St. Fair Isle. N. B. Kinnear (c & p). 6. 11. 18. 1.

2 6,1 9. Fair Isle. N. B. Kinnear (c & p). 6. 11. 18. 2-4.

6, 2 ?. Fair Isle. Duchess of Bedford 10. 10. 11. 1-3.

(c & p).

APODEMUS

827

3

JO

O

Teeth much worn.

,, slightly worn.
,, much worn.

,, moderately worn.
,, much worn.

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•siqipaBM

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r^T^r^ rHiHiHi-lrHi-lTHTHiH rHrHiHrHi-lTHrHi-liHT-H

' •Bm9')seic[

CD CD O CM O O 00 O O CO <M O O 00 00 CO ^ 00 O CO 00 CO

t-t-00 t- CO 00 t- CO 00 t- 00 00 00 !>• t- t- t- t- 00 t- t- t-

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O CO O O "H O O CD CD CO O CM ^ oq 'iH -*

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JO iildgo;

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CO 00 Cn 00 00 CO 00 00 oo oo oo oo oo oo oo oo oo

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tHtHtH tHt-Ii— ItHt-I tH iHtHt-Ii— ItHt— IrHiHrH

•uonouisuoa
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Number. '

95. 10. 25. 1*

94. 7. 16. 1*
11.1.2.123
153507
1905. 120. 2. 7t
9t
16t
17t
1905. 120. 4t
1905. 120. It

6.11.18.2

6.11.18.4

196t

2S9t

153499

153500

153502

153503

103504

103506

locality.

.2

k1 '^.

nsis.

id of

sis.

Islan

isis.

A. hebride

Scotland : Uig, Islai

A. hirtens

Scotland : St. Kilda

A. fridariei

Scotland : Fair Isle

828

APODEMUS FLAVICOLLIS Melchior.

(Synonymy midcr subspecies.)

Gcixjnipluriil diHtrihiitioii. — 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 Apodcnnis sj/lvaticus sjil-
vaticHs, 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; condy-
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 upfjer 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. sylvaticus, frecjuently spreading laterally
to form a complete collar.

External characters. — As in Apodemns 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 ]:)arts, and the constant absence
of all trace of brownish wash below, the colour so exactly
resembles that of Apodemus sylraticns sylratlcus as to need no
detailed description.

Slcull 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
Large specimens of Apodemus flavi-
ikulls (if inunature A. epiinelas ; but

KiG. lev.

A podcmiis jlaKiciiUis.

that of the smaller animal.
colJis are ecjual in length to

Al'ODEMUS S'2d

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 i-esemble those of Apodemus s/jlvaticus.

BemarJiS. — The large general size, long hind foot, and massive,
angular skull serve to distinguish adults of this species from
individuals of Apodevius sylvaticus. 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 v/here A. sylvaticus
has been found, are both circumstances which suggest the
probability that Apodemus flancoll is 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. Mils flavicollis Melchior, Den Danske Sfcaats og Norges Pattedyr,
p. 99 (Sielland, Denmark).

1862. ? \Mns sylvaticus] vrt. major Radde, Eeisen im Siiden von Ost-
Sibirien, i, p. 180 (Bureja Mountains). Part, specimens mentioned
from Crimea which are probably this form.

1866. Miis cellarius J. F. Fischer, Zool. Gart., vii, p. 153, April, 1866
(near Luga, St. Petersburg, Russia).

1874. Mus sylvaticus 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 sylvaticus typicus Barrett-Hamilton, Proc. Zool. Soc, London
p. 404. Not Mus sylvaticus Linnaeus.

1900. Mus sylvaticus jirinceps Barrett-Hamilton, Proc. Zool. Soc, London,
p. 408 (Bustenari, in the Carpathians, N.W. of Bucharest,
Roumania. Altitude 480 m.). T3-pe in British Museum.

1910. Mus sylvaticus and M. sylvaticus princeps Trouessart, Fauna Mamm
d'Europe, pp. 149, 153.

Type localitij. — 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, LoUand, 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 prince'ps) : 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 from 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
looalites : —

Sweden : Jemtland, 2 (U.S.N.M.) ; Herjedal, 1 (U.S.N.M.) ; Medelpad,
Norrland, 4.

Denmark : Hillerod, Zealand, 3; Nysted, Lolland, 2 (U.S.N.M.).

France: Ax-les-Thernies, Ariege, 5 (B.M. and U.S.N.M.); Caterille,
Haute-Garonne, G ; Bareges, Hautes-Pyr6n6es, 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.M.) ; Bahrenberg, Harz Mountains, 3
(U.S.N.M.) ; Bodethal, Harz Mountains, 6 (U.S.N.M.) ; Mauseklippe, Harz
Mountains, -5 (U.S.N. IM.) ; 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.

EoDMANiA : Bustenari, Prahova, 15 ; Gageui, 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.); Degersheim, St. Gallen, 1 (U.S.N.M.); Wild-
kirchli, Appeuzell, 1 ; Albula Pass, Grisons, 4 (U.S.N.iNI.) ; Julier-Hospiz,
Gri,sons, 1 (U.S.N.M.); Vulpera-Tarasp, Grisons, 4 (Rothschild) ; Vitznau,

APODKMDS 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.

Geeece : Athens, 6 ; Corfu, 3.

6,9. Medelpad, Norrland, Lord Lilford (p). 8.10.19.28-29.

Sweden.
3 6. Hillerod, Zealand, Den- 0. Thomas (c & p). 98. 6. 7. 9-10. 13.
mark.
2 6, 9. Ax-les-Thermes, Ari^ge, V. Builles (c & p). 8. 3. 27. 9-11.
France.
6(5. Caterille, Haute-Garonne. O. Thomas (p). 6.4.1.48-53.

{A. Robert. )
9. Bareges, Hautes-Pyr6n6es. G. S. Miller (c). 8.8.4.221.

3<J, 9. Oranves- Sales, Haute- O. Thomas (p). 5,11.18.14-17.

Savoie. (A. Robert.)
1 al. Tharandt, Saxony. Berlin Museum (e.) 93. 1. 1. 19.

1. Kiesky, Silesia. (W.Bacr.) Lord Lilford (p). 99. 1. 9. 19.

2 i, 9. Mittelberg, Vorarlberg, O. Thomas (p). 8. 11. 30. 9-11.

Austria-Hungary.

{Dr. F. Major.)
5 (5, 2 9. Hatszeg, Hunyad, Tran- C. G. Danford (c). |3. 2. 2. 26-29. 31.

sylvania. (8. 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. O. Thomag (p). 5. 8. 4. 36.

(L. Filhrer.)

4 (J, 2 ?. Locarno, Ticino, Switzer- 0. Thomas (c & p). 2. 7. 1. 13-18.

land.
2 6, 9. Vitznau, Lucerne. O. Thomas (c & p). 5. 8. 3. 16-18.

3 (5, 3 9. Athens. (C. Mottaz.) Hon. N. C. Roths- 8. 10. 2. 27-32.

child (p).
6,2 9. Corfu. (C. Mottaz.) J. L S. Whitaker (p). 8.10.1.23-25.

Apodemus flavicollis wintoni Barrett-Hamilton.

1894:. Mils flavicoUis de Winton, The Zoologist, 3rd ser., xviii, p. 441,

December, 1894 (part).
1900. Mus sylvaticiis wintoni Barrett-Hamilton, Proc. Zool. Soc, London,

p. 406 (part). Type in British Museum.
1910. Mus sylvaticiis tvintoni Trouessart, Faune Mamm. d'Europe, p. 149.

Type locality. — Graftonbury, Herefordshire, England.

Geographical distribution. — 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. — Tyjae (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-() (17-19). Adult male from AVoolpit, Suffolk: head ancl

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see Table, p. 835.

Specimens exaynincd. — 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, Suifolk, 1 ; Balcombe, Sussex, 1.

Eemarlis. — The British race of Apodpnnis flavicolUn, though
not a well marked form, seems worthy of recognition on account
of the duller colour of the underparts and the more diffuse
nature of the pectoral spot.

6. Riding Mill - on - Tyne, Rev. H. H. Slater 11. 1. 3. 413.

Northumberland, (c & p).
England.

i. Graftonbury, Hereford- W. E. de Winton 0. 3. 12. 1.

shire. (c & p). (Tyjx'oi subspecies.)

4 <?, 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.

i. Bishopstone, Hereford- Rev. S. O. Ridley 81. 1. 1. 4.

shire. (c & p).

<5. Ledbury, Herefordshire. Col. J. W. Yerbury 11. 1. 3. 411.

(c & p).

5. Swithland, Leicestershire. F.A.Butler (c & p). 11.1.3.410.
9. Ouudle, Northampton- Lord Lilford (c & p). 11. 1. 1. 1G5.

shire.

6. Woolpit, Suffolk. W. D. Parker (c & p). 11. 1. 3. 412.

APODEMUS AGRARIUS Pallas.

1778. Mtis agrarivs Pallas, Nov. Sp. Quadr. Glir. Ord., p. 95 (Berlin,

Germany).
1801. ilf[MS] agf[arius] albirostiis Bechstein, Gemeinn. Naturgesch.

Deutschlands, i, 2d ed., p. 975 {Thviringen, Germany).
1801. Mills'] agr[arius] macxdatuH Bechstein, Gemeinn. Naturgesch.

Deutschlands, i, 2d ed., p. 975 (Thiiringen, Germany).
1816. Mus rubens Oken, Lehrb. d. Naturgesch., iii, pt. ii, p. 898 (Northern

Germany).
1857. Mus agrarius Blasius, Siiugethiere Deutschlands, p. 324.
1908. A[j)ode7]ms'] agrarius Thomas, Proc. Zool. See. London, 1908, p. 7,

June, 1908.
1910. Mas (Apodcmus) agrarius Trouessart, Faune Mamm. d 'Europe,

p. 155.

Type locality. — Berlin, Germany.

Geographical distribution. — Eastern and central Europe, west
along the Baltic coast to Denmark.

Diagnosis. — Size and external appearance about as in
Apodemns syhaticus, but colour of ujaper parts more reddish, and
back with a narrow, sharply defined, black median stripe ; skull
with interorbital region conspicuously beaded, even in half-grown
individuals ; mcjlars of about the same size as in A. syh-aticus,

APODEMUS

837

I

but tir with no iiiiiiule aiiturior cusp uu outer .sido, and i/ij and
m.^ without cinguhnn-like outei- ledge.

External characters. — The general external form is less slender
and delicate than in the members of the Apodemini syJvaticus
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 coarsely annulated. Palms as in
A. xyhmticus, except that the two intermediate tubercles are
relatively smaller. Feet with metatarsal portion relatively
shorter than in A. sylvaticus, 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. sylvaticus. Caudal
annulations very distinct, about fourteen to the centimeter at
middle ; caudal hairs essentially as in A. sylvaticus but less
abundant, the pencil ill-defined. Mammte :p2 — 2,i2 — '2 = ii.

Colour. — Upper parts a reddish brown between the russet
and wood-brown of Ridgway, brighter posteriorly, more dull
anteriorly, fading nearly to ochraceous-bufi" 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 ; underjjarts and inner surface of legs dull white
clouded by the slaty undercolour ; tail not bicolor, the general
effect hair-brown somewhat darker above than below ; feet vary-
ing from pale drab to dull whitish. The
young are essentially like the adults, differ-
ing only in a slight general dulness of
colour.

Slull. — In general appearance the skull
resembles that of Apodemus sylvaticus, 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 jiortion of the
supraoccipital is visible when skvdl is viewed
from above. Interparietal with antero-
posterior diameter as in A. sylvaticus, but
with latei-al extremities abruptly and
almost scjuarely truncate at region where
bone has width of about 1 • 5 mm., thus con-
siderably reducing transverse diameter ; pos-
terior border of interparietal lying along lambdoid sutvire, the outer
anterior angles of the bone therefore encroaching on parietals.*
Margins of interorbital region conspicuously beaded, the raised

* In Mas musculus, which has a similarly ligulate interparietal, the
lambdoid suture passes along the anterior border of the bono, so that the
parietcils arc not emargiuated (fig. 177, p. 866).

Fig. 16S.
ApodevnLs agrarius.

838

KODENTIA

edge narrow and abrupt in adults, lower in immature individuals,
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. sylvaticus. Incisive foramina
noticeably shorter and (relatively) wider than in the other
European members of the genus, the posterior margin extending
barely or not to level of alveolus of m\ the anterior margin
separated from alveolus of incisor by a distance fully equal to
combined width of foramina. Auditory bullie more inflated than
in ApodeiauH sylvaticus, so that hamular is nearly or quite in
contact with inflated portion instead of with a conti'acted beak-
like process as in the related sj^ecies. While in general
resembling that of A^wcleiuus sylvaticus the mandible is readily
distinguishable by the narrower less curved angular process and
the higher more curved coronoid, the extremity of which is
distinctly above level of condyle.

Teeth. — The teeth are noticeably smaller than in Ajjodoiiiis
sylvnticiis : alveolar breadth of m} scarcely more than one-thirtl
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 m^ slightly
longer than that of wt" and iic'
together-, its area decidedly more
than tliat 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 t'i I'educcd to a mere appendage
to the outer border of /2 ; t\ well
developed, distinct, very posterior in
position, this peculiarity together with
the small size ol t'i producing great
distortion of the anterior crescent ;
no supplemental loops in angles
between /3 and V2 or i2 and t\.
as in A. sylvaticus except for the
of the second crescent and the more abrupt rounding
of the posterior border I'esulting from the complete absence of
all trace of ^9. 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 /G
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 j/j^ and w(^ nearly obsolete.

Rest of
flattenin

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APODKMUS

839

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S40

RODENTIA

Measureiamts. — Average and extremes of eight adults iixmi
Berlin, Germany: head and body, 98 "l (97-112); tail, 78-.")
(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 ( 1 8 • 6-1 9 • 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 following 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.) ; IBieberstein, Saxony, 1 ; Saxony, no exact locality, 2
(U.S.N.M.) ; vicinity of Konigsberg, 32 (U.S.N.M.).

Austria-Hungary : Eszek, Slavonia, 1.

EouMANiA : Bustenari, 1 ; Grageni, 2.

Bulgaria: Rustschuk, 2 (Andersen).

Bemarlcs. — This animal is immediately recognizable among
European Muridie by the shai'ply defined black dorsal line, and
among the smaller members of the family by the noticeably
beaded margin to the interorbital j^ortion of the skull. A similar
colour pattern occurs in the structurally very dilferent Siclt<t(t
frizona, and as the ranges of the two animals partly coincide
confusion may easily result. Superficially Sicista trizona 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 (5, 5 ?. Berlin, Germany. Dr. L. Heck (c &: p). 7. 6. 12. 1-8.

4<5. Herrenkrug, Magdeburg, Dr. W. Wolterstorlf 92.12.1.10-17.
4 9 al. Saxony. (c & p).

9 Altenberg, Saxony. E. R. Alston (p). 79. 9. 25. 42.

(Muschler.)

1 al. Bieberstein, Saxony. Berlin Museum (e). 93. 1. 1. 18.

{Nitsche.)

1 al. Eszek, Slavonia, Hun- Budapest IMuseum 94. 7. 23. 1.
gary. (k).

6 Bustenari, Prahova, Rou- Lord Lilford (p). 4.4. G. 50.

mania. (W. Doclson.)

i, 9. Gageni, Prahova, Rou- Lord Lilford (r). 4. 4. G. 51-52.

mania. (1-7. Dodson.)

Purchased (Ver- 63. 11. 16. 3.

reaux).

Genus MICROMYS Dehne.

1841. Micronuji Dehno, IMicromys agilis, ein neucs Siiugthier 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).

Time .siwcies. — Mirromi/f; lu/iliff Dehne = Miit< ><oririnus Her-
mann.

Geixjrdjjhk-ul distrihiitiou.. — Central Europe and Asia from

MICROJIYS _ 841

(Jreat Britain eastward ; iu Europe north to Scotland and
Denmark, south to Italy and southern France.

Dingnosis. — Skull difl'ering from that of Apodcvms 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 Apodemus, but m^ 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.

MemarAs. — 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.)

Geoyraphical distribution. — Same as that of the genus
Micromys.

Didtjnosis. — General characters as in the genus ; size very
small (head and body, 55 to 75 mm. ; hind foot, 13 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 consjiicuous tuft of hairs neaidy 5 mm. in length.
Feet relatively broader than those of Apodemus sylvaticus, but
proportions of digits not peculiar ; palmar and plantar tubercles
essentially as in Apodemus 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 prehensile, 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 mimitits Pallas, Reise durch verschiedeno Provinzen des
liussischeu Reichs, i, p. 451 (Banks of the Volga, Russia).

842

KODENTIA

SJiiiIl. — The skull differs conspicuously from that of all other
European Murime in the striking contrast between the short
weak I'ostrum and the large, much elongated brain-case ; 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 i:)rofile 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
anteriorh^ than posteriorly, to interparietal, where a slight
bi-eak in the j^rolile is produced by the more abrupt convexity
of the overhanging posterior portion of brain-case ; ventral
profile without sjsecial features, essentially straight from incisor
to condyle, except for the slight break caused by the auditory
bullte. 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 j^art of brain-case, and
almost marked off by a constiiction at
front of interparietal ; in posterior view
the middle portion of brain-case rises
unusually high above interparietal, and
distance from lambdoid sutui'e to upper
margin of foramen magnum is unusunlly
reduced : paroccipital processes very short,
extending scarcely to level of lower lip
of foramen ; flooi- of brain-case with no
special features, the median I'idge and
lateral furrows well developed, the anterior
width of basioccipital less than median
length ; auditory bullsB rather large, evenly
inflated, without evident beak, the meatus relatively large,
without lip except along anterior border. Interorbital region
i-ather wide and short, smoothly I'ounded off at sifles though
tending to become slightly angular posteriorly in old individuals.
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 boi'der of zygomatic I'oot wlien ^■iewed from abo^"e, 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 Apodemug stjhmticus, 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 widtli of single foramen. Anterior portion of palate
* Not successfully represcutcd in fig. 170.

Fig. 170.

.MU-riimyx mhiulus.

Nat. size.

:microjiys

843

witlioiit special pL'tailiai'ities ; a sliglitly indicated transverse
ridge at level of posterior border of last molar ; mesopterygoid
space beginning about 1 mm. behind this ridge, much narrower
anteriorly than posteriorly, its margins straight to tip of hamular,
which is barely or not in contact with auditory bulla. Mandible
essentially like that of Apodevtiis niilnaticun, 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 Ai)ii(lc)iius
xylvaticns, 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 Apoclemus sijlraficus than as in
A. (ujrnr'ms, 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, <3 well developed
though closely crowded against t'2, t\ some-
what posterior in position, the three tuber-
cles less contrasted in size than in the species
of Ajyodciaus ; no intermediate folds in
narrow angles between bases of tubercles,
but a rather well defined ridge from ^1 to
inner base of /5 ; /6 very large, filling nearly
the entire space occupied by it and /9 in
Apodemus siilvaticus : td reduced to a mere „. "^' '.' ,

rudiment at posterior base of /6 ; postei'o (.;iieek-teetu. x lo.
external margin of crown with slight thick-
ening of the enamel as in AjMdciims sylvaticiis ; /S and tl showing
no special peculiarities. Second upper molar with first crescent
represented by a minvite though evident to and a small tl ; second
crescent with simple t6 about ecjual in size to <4 ; third crescent
with barely indicated td and small /7. Third upper molar much
as in Aimdcmus si/lvaticus, its crown showing traces of tubercles
1, 4, 5, 7 and 8. Maxillary molars as in Apodemus sijhnticus,
but outer ledge of m^ and m., slightly developed and without
evident cusps.

Bemarks. — Two well marked races are recognizable among
the harvest mice of western Europe. True Micruiinjs minutus is
aj^parently not found in this region.

KEY TO THE EUROPEAN FORMS OP MICBOMYS MINUTUS.

Head and shoulders reddish brown, essentially

coucolor with back (England and western

continental Europe) -V. minutus t:orkinus, p. 844.

Head and shoulders greyish, noticeably contrasted

with reddish brown back (Hungary and Rou-

niania) -i/. >n. pratensia, p. S4G.

844

RODENTIA

MiCROMYS MiNUTUS soRiciNUS Hermann.

1780. Mils soricinns Hermann iu Schreber, Siiugethicre, iv, p. 661,

pi. CLXXXiii B. (Strassburg, Germany).
1785. [Mus] triticcus Boddaert, Elenchus Anim., i, p. Ill (Hampshire,

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., p. 230 (Hampshire, England).
1794. ^^ Mus avenarius Wolf, Versuche die Feldraiiuse zu vertilgen, p. 16,

315," Hermann, Observ. Zool., p. 61, 1804 (Locality not men-
tioned).
1804. Mus pendulimis Hermann, Observ. Zool., p. 61 (Strassburg, Germany).
1804. Mus parmilus Hermann, Observ. Zool., p. 61 (Strassburg, Germany).
1816. ? Mus arvensis Leach, Syst. Catal. Spec. Indig. Mamm. and Birds,

Brit. Mus., p. 7 (Nomen nudum : "Harvest Rat ").
1822. 3fus campcstris Desmarest, JIammalogie, pt. 2, p. 543 (France).
1832. Mlus] mcridionalis 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, INIicromys agilis, ein neues Siiugthier der

Fauna von Dresden, p. 1 (Dresden, German}').

1841. Mus oryzivonis de Si^lys-Longchamps, Atti della seconda Riunione

degli Scienziati Italiani, Torino, 1840, p. 247 (Rice fields of
Lombardy, Italy).

1842. Mus pumilus F. Cuvier, Hist. Nat. des Mammif., Tabl. gen. et

Meth., p. 4. Described in fasc. xxxii, October, 1821 (Vicinity of
Paris, France).

1857. j\[us minutus Blasius, Siiugethicre Deutschlands, p. 326.

lyOC). Mus minutus agilis Barrett-Hamilton, Ann. and INlag. Nat. Hist.,

7th ser., v, p. 629, June, 1900 (Brunswick, Germany).
1900. Mus minutus campestris Barrett-Hamilton, Ann. and INIag. Nat.

Hist., 7th ser., v, p. 529, June, 1900 (Waremmc, 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.) mcridionalis Trouessart, Faune

Mamm. d'Europc, pp. 1.56, 157.

Tjipe locidUy. — Strassburg, Germany.

Geographical distribution. — Central Europe from Great Britain
to eastern Germany, and from Scotland (Aberdeenshire) and
Denmark south to Italy and southern France.

Diagnosis. — Colour of upjjer jiarts essentially uniform through-
out, the shoulders, neck and head never conspicuously greyish.

Colour. — Winter pelage (England and Switzerland) : upper
parts a nearly uniform reddish brown approaching the tawny of
Ridgway 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-buft" ; back very
inconspicuously sprinkled with black-tijiped hairs ; ear concolor

MICROMYS 845

with sides ; underparts 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-
buff, the skin light brown below, dark above. Summer pelage
(Germany) : upper jjarts noticeably darker and duller than in
winter, approaching the russet of Ridgway, but with a tinge of
rufous ; sides dull cinnamon ; underparts dull 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-detined 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) ; Rugies, near I'A-reux, Eure, 2
(U.S.N.M.).

Belgium : Waremme, Li6ge, 3 (U.S.N.M.).

Gebmany : Holstein, 1 ; Brunswick, 27 (U.S.N.M.) ; near KiJnigsberg, 9
(U.S.N.M.) ; Strassburg, 1.

Switzerland: St. Piden, 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(5,?

1.
3(5.

1

4 (5,2 9.

Haddiscoe, Norfolk, Eng-
land. (L. C. Fm-man.)
Congham, Norfolk.
Norfolk.

(Oxlcy Grabham.)
Cambridgeshire.
Colchester, Essex.

G. Barrett-Hamilton

Oxley Grabham (c & p)
G. Barrett-Hamilton

(P)-
J. Baker (c & p).
Dr. Henry Laver

(c & P).
E. R. Alston (p).

11. 1. 2. 124-128.

11. 1. 3. 414.
11. 1. 2. 129-131.

39. 9. 29. 31.
3. 4. 8. 1-6.

1.

Kingsbury, Middlesex.

79. 9. 25. 61.

(J. L. Uarting.)
1 al. Totton, Hampshire. E. B. Wharton (c & p). S3. 4. 30. 1.

846 EODENTIA

1. Somerset. [Hiigel.) E. R. Alston (p). 79.9.25.62.

1. Holstein, Germany. Purchased (Brandt). 47.4.5.2.

1. Strassburg, Alsace. C. Mottaz (c). 8. 8. 10. 90.

MiCROMYS MixuTUS PRATEXSis Ockskaj.

1831. Mus pratensis Ockskay, Nova Acta Phys.-Med. Acad. Caes. Leop.-
Carol. Nat. Cur., xv, pt. ii, j). 243, pi. lxviii (Western Hungary).

1882. Mus arundinaceus Petenyi, Termeszetrajzi Fuzetek, v, p. 142.

1900. Mus viinidus i^ratensis Barrett-Hamilton, Ann. and ilag. Nat. Hist.,
7th ser., v, p. 530, June, 1900.

1910. Mus {Apodmms) viinutus iwatensis Trouessart, Faune Mamm.
d'Europe, p. 157.

Tiipc locaHty. — Western Hungary.*

Geograpicnl (listrihutlon. — Hungary and Roumania ; limits of
range not known.

Diaqnosis. — Similar to Micromys mtnutiis soriciimx, but head
and anterior half of body decidedly greyish.

Colour. — Posterior half of back together with outer surface
of hind legs as in 31. soririnns. Head and anterior half of liodj-
a buffy grey, darker and more brownish along median dorsal
i-egion, 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 Csallukuz-Somorja, Pressburg, Hungary : hind
foot, 1:3-2, 13-6, 14 and 14. '

Specimens examined. — Nine, from the following localities : —
AuSTEiA-HuNGARY : Csallokoz-Somorja, Pressburg, Hungary, G ; Cepin,
near Eszek, Slavonia, 2 (in spirit).
Roumania: Gageni, Prahova, 1.

Reniarlcs. — The few specimens examined indicate that this is
a well-defined race.

G. Csallokuz-Somorja, Press- Budapest Museum (r.). 94. 3. 1. 53-58.
burg, Hungary.
2al. Cepin, Slavonia. Budapest Museum (k). 94. 7. 23. 11-12.

? Gageni, Prahova, Rou- Lord Lilford (p). 4. 4. G. 49.

mania. {W. Dodson.)

•Frequentius occurrit in Comicatus Nittrienses, valle Vagi."

847

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848

Genus EPIMYS Trouessart.

1857. Mus Blasius, Saugethiere Deutschlands, p. 309 (part).

1867. Rattus Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien, Math.-

Naturwiss. Classe, lvi, p. 6.3 (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 Linnaeus.
1910. Epimys Miller, Proc. Biol. Soc. Washington, xxiii, p. 58, April 19,

1910.

Ti/pe species. — Mus rattus Linnteus.

Geographical distribution. — Originally confined to the temperate
and tropical portions of the Old AVorld, 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 in^ 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
ol tl ; upper incisor compressed, set at such an angle that its
outer side is worn smoothly away by action of lower teeth.

Remarlcs. — The genus Epimys is the most widely distributed
and abundantly represented group in the sub-family Mnrinse.
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 OP EPIMYS.

Greatest breadth of brain-case across lateral ridges
about equal to length of parietal measured
along ridge ; first lamina of ■?»' and terminal
heel of w- with no well developed outer
tubercle ; external form rather heavy ; tail
decidedly shorter than head and body (Nor-
way rat) E. norvegiciis, p. 858.

Greatest breadth of brain-case across lateral ridges
decidedly more than length of parietal mea-
sured along ridge; first lamina of »i' with
distinct outer tubercle usuallj' 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. rattus, p. 849.
General colour slaty black, the underparts dark

slaty grey (Distribution general) E. ,: raitu.-;. p. 853.

General colour brown, the underparts buffy or

greyish (Southern) H. r. alexaixln'nux, p. 854.

I

S4y

EPIMYS RATTUS LilUia'US.

(Synonymy under subspecies.)

Ge<i[ini[iliii<il (}islrihii(iiin. — Orii^iiially confiiitjd to tlie iioftli
tenipurate portions of the (Jld AV'orld ; now essentially cosmo-
politan through artificial dispersal.

Di(i(/iiflsiti. — Skull with brain-case broad, its greatest breadth
across lateral ridges decidedly moi'e than length of parietal
measured along ridge ; anterior upper molar without trace of
(■ingulum at anterior border of crown ; first lamina of la^ with
distinct outer tubercle usually almost as large as inner tubercle ;
terminal heel of wi- with evident though incompletely separated
outer tubercle ; tirst and second lower molars usually with
evide?it 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
40 mm. in length ; coudylobasal length of adult skulls usually
oS to 43 mm.

External characters. — General form nither slender, the 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 when laid
forward, its substance thin and somewhat translucent, its
general outline broadly oval, the basal portion of anterior border
folded backward (inward) ; posterior border of meatus with
barely indicated ridge ; surface of ear finely papillose and very
inconsijicuously pubescent. Muzzle pad small but well defined,
with deep median groove continuous with cleft in upper lip ;
nostril crescentic, its inner and lower margins swollen, forming
an ill-defined, wart-like excrescence below. Feet moderately
large, with no special peculiarities 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
live, 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 digit,
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 postero-external 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 feet largest.
Tail slightly four-sided, uniformly and very thinly haired
throughout, the tip with no distinct pencil ; annulations well

o I

850.

RODENTIA

defined, regular, about nine to the centimeter at middle ; length
of hairs equal to about one and one-half width of rings. Fur
soft but interspersed with numerous coarse hairs which impart
to it a somewhat harsh quality ; with a lens these coarse hairs
aie seen to be very slender bristles, distinctly grooved on the
upper side. Mammse : jj 2 — 2, i 3 — 3 = 10.

Ciihnir. — The two main types of colour are (1) dark slaty,
becoming almost blackish on back, and (2) light brown above,
jiale buff or light grey below. All possible intermediate stages
exist connecting the two extremes.

SlnJl. — In general featui'es the skull shows no marked
peculiarities. Dorsal profile faintly convex throughout, but

Kir,. ITJ.
Jijiiinyx riiUiis. >'at. size

flattened over posterioi' lialf of nasals, between orbits and on
posterior half of brain-case ; occiput squarely and almost perpen-
dicularly truncate ; ventral profile nearly flat. Brain-case squarish
in outline when viewed from above, but slightly longer than
wide and somewhat narrowed posteriorly, the sides of its upper
surface 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
cf parietal measured along ridge. No trace of sagittal crest,
but a low ridge-like lambdal crest present in old individuals and

lU'iMVs 851

a kiiifc-]iko iiiodiau occiuital crest from interparietal to foramen
magiumi. 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 beliind, but central jiortion of brain-case not rising con-
spicuously above it ; paroccipital processes slender and rather
long, extending beyond level of lower lip of foramen. Floor of
bi-ain-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 incisoi's, its mai'gin 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.
Rosti'um much deeper than broad but without special peculiarities
of form ; nasals truncate or bluntl)^ 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 m.^, 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
bulke ; ectopterygoid well developed, tilted noticeably upward
from within outward (skull viewed from below). Mandible
robust, with 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 strongly
compressed, without special peculiarities, the cutting surface of
upper tooth vertical, flat, its edges entire. First upper molai'
5-rooted, the anterior I'oot largest, strongly projecting forwai'd
beyond base of crown, the median inner and median outer roots
smallest ; crown barely as long as that of m'^ and hi'* 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; tirst lamina moderately and evenlv crescentic,

3 1 -4

852

RODENTIA

Eliiiinj^

Fir;. 173.
(tllu.'<. C'licuk-tt

the outer tubercle nearly as large as U, and only a little anterior
to it in position, both small tubercles well deiiued from Vl, 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, tl absent. Second upper
molar with first lamina represented by large t\ ; second and third
laminte essentially as in m^, the third with well developed <9
nearly as large as that of preceding tooth and marked off from
/8 by a distinct re-entrant angle. Third upper molar with large,
distinct, terete /I, small ti and <5, the
latter joined by an obsolete ^6 to outer
extremity of i(8, the only remnant of
the third lamina ; as the crown wears
away the second and third lamina^
assume the form of a narrow loop, the
twu limbs of which are separated by a
deep re-entrant angle. Anterior lower
molar with crown scarcely as long as
tliose of the two succeeding teeth, its
area also less ; first lamina narrower
than second, its posterior border nearly
straight, its anterior border with deep
re-entrant angle on outer side of
middle ; second and third lamime 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 laminib, and a similar though smaller tubercle occasionally
present in that between lirst 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 lamina;.
Third lower molar with two laminse, 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.

lieniarks. — 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 Epuiiys ratliia have established themselves in
western Europe. True raltiia, 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, Eplmijs rattus alexandrinm, 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.

EriMYS ^^)-^

form lin,vc been n,i'tiiifiiill\' tliou^h unintentionally transplanted
into tiie area of the other, and to this circumstance, rather than
to individual variation or normal intergradation, should be
atti'ibuted the intermediate specimens now so abundantly found.

Epimys rattus rattus Linnanis.

1758. [Mii.i] rattus Linnseus, Sj-st. Nat., i, 10th ed., p. Gl (Sweden).

1833. Mils tectornni var. fuUginosus Bonaxsarte, Iconogr. Faun. Ital., I,

fasc. 3, pi. 22, fig. 1, name on plate only (Italy, no exact locality).
1842. Mus ftubaervleus Lesson, Nouv. Tabl. du Begne Anim., Mamm.,

p. 138 (Roohefort, Charente-Inferieure, France).
1857. Mus rattus Blasius, Saugefchiere Deutschlands, p. 317.
18G7. Ttattus doniesticus Fitzinger, Sifczungsber. kais. Akad. Wissensch.

Wien, jNIath.-Naturwiss. Classe, lvi, pt. i, p. 64 (Substitute for

rattus) .
1SG7. [Raltiis domcsticus'] fiiscus Fitzinger, Sitzungsber. kais. Akad.

NV'issensch. Wien, Math.-Naturwiss. Classe, lvi, pt. i, p. 64 '

(Germany).
1867. [Rattus doniesticus'] varius Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lvi, pt. i, p. 64

(Germany).
1867. [Rattus domcsticus~\ fulvaster Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lvi, pt. i, p. 64

(Austria and Germany).
18G7. [Rattus doniesticus] albus Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, ]\Iath.-Naturwiss. Classe, lvi, pt. i, p. 65

(Austria, Hungary, Germany).
1867. [Rattus doniesticus] 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. Soe.

Zool. de Geneve, i, p. 163, November 15, 1908.
1905. Mus rattus ater INIillais, Zoologist, 4th ser., ix, p. 205, June, 1905

(London, England). Type in British Museum.
1908. Epimys rattus Satunin, Mitth. 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.

Tjjpe localUy. — Upsala, Sweden.

Geographical distribution. — Europe north of the Mediterranean
region ; occurring in isolated colonies and every ^Yhere being
11 'placed by Epinufs norvegicus.

Diagnosis. — General colour slaty, the back darker, frequently
almost black.

Measurements. — Adult male and female fi'om 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,
I'ngland (type of ater Millais) : head and body, 235 ; * tail, 235 ; *

* From ^Millais.

854 RODEXTIA

hind foot, 36 ; ear, 25.* Adult female from Hatszeg, Hunyad,
Austria-Hungary : head and body, 190 ; tail, 225 ; hind foot, 38.
For cranial measurements see Table, p. 856.

Specimens examined. — Twenty, from the following localities : —
Scotland: South Ronaldshay, Orkney Islands, 5 (Edinburgh).
England : Great Yarmouth, Norfolk, 1 ; Tring, Hertfordshire, 1 ;
London, 6 (including type of atcr JMillais) ; Lundy Island, Devonshire, 2.
Prance : Sark, Channel Islands, 1.
Switzerland : No exact locality, 1.
Austria-Hungary: Hatszeg, Hunyad, 2.
Portugal : San Miguel, Azores, 1.

6 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).

^, 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 liter :\lillais.)

2 <?. Lundy Island, Devonshire. T. A. Coward (c & p). 7.12.19.1-2.

9. Switzerland. (Ncujer.) E. R. Alston (p). 79. 9. 25. 40.

6,9. Hatszeg, Hunyad, Tran- C. G. Danford (c). 3.2.2.22-24.
sylvania.

S. San Miguel, Azores. Hon. W. Rothschild 3. G. 5. 15.
(ir. B. O. Gi-ant.) (p).

Epimys rattus ALEXANDRixas Geoffroy.

1803. Mux ali'xandrinus Geoffroy, Catal. ilammif. du Mus. Nat. d'Hist.

Nat., Paris, p. 192 (Egypt).
1814. Musculua frugivorus Rafinesquc, Precis des Decouv. et Travaux

Somiologiques, p. 13 (Sicily).
1825. Mus ti'cturum Savi, Nuovo Giorn. de' Letterati, Pisa, x, p. 74 (Pisa,

Italy).
1827. Mijo.vus siculx Lesson, Man. de Mammalogie, p. 274 (Substitute for

Musculus frugivarus Rafinesque).
1841. Mus sylvestris Pictet, j\Km. Soc. Phys. et d'Hist. Nat., Geneve, ix,

p. 153 (Near Geneva, Switzerland). Alternative for Icucogaster.
1841. Mus leucogaster Pictet, ]M6m. Soc. Phys. et d'Hist. Nat., Geneve, ix,

p. 154 (iSTear Geneva, Switzerland).
1841. [J/«s] ncmuralis de Selys-Longchamps, Atti della seconda Riunione

degli Scienziati Italiani, Torino, 1840, p. 247 (Near Geneva,

Switzerland). Accidental substitute for sylvesti-is Pictet.
1845. Mus incieti Schinz, Synops. Mamm., ii, p. 142 (Substitute for Irucu-

gaster Pictet).
1857. Mus alexandrinus Blasius, Siiugethiere Dcutschlands, p. 31G.
1882. [Mus rattus] intermedins Niuni, Atti del reale Inst. Veneto, 5th ser.,

VII, p. 574 (Venice, Italy).
1910. J^lus (EphniiK) rattus intermedius and M. (F.) raltiis ale.viindrlnus

Trouessart, Faune Mamm. d'Europe, pp. 143, 144.

Tijiir /(ico/////.-- Alexandria, Egypt.

(ii'diirajiliicdl dis/rihntion. — Mediterranean region ; everywhere
abundant and not teniling to l)e replaced ))y Ejiiiiii/.i itnrcrtj'irus.

* From ^fillais.

855

DiiupiOKi's. — Upper jjarls li<^lit ln'tjwn; uiuJorpartfj strongly
contrasted jmle buff or light grey.

Measurements. — Two adult females from near Nimes, Gard,
France: head and body, 180 and 199 ; tail, 222 and 250 ; hind
foot, ;56 • 5 and ;58 • 5 ; ear, 24 and 27. Two females from Sorrento,
Italy: head and body, 177 and 178; tail, 243 and 217; hind
fo(jt, 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 (19G-
21G); 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. 856.

Sixcitnens examined. — One hundred and twenty- three, from the follow
iug localities : — ■

England: Sunderland, Durham (on ship), 1; Lundy Island, Devon-
shire, 6.

Spain : Pajires, Leon, 2 ; Arrechavaleta, Vitoria, 2 ; B6jar, 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.

Peance : Biarritz, Basses-Pyrenees, 7 ; near Nimes, Gard, 17 ; Agay,
Var, 3; Barcelonnette, Basses-Alpes, 1 (U.S.N.M.).

Switzerland : Lugano, Ticiuo, 1 (U.S.N.M.) ; Agra, Ticino, 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, Cephalonia, 14; Kephissia, Athens, 4;
Canea, Crete, 2.

i. Sunderland, Durham (on
board ship), England.
3 (5, 2 9. Lundy Island, Devonshire.
6, ?. Pajares, Leon, Spain.

{N. Gonzales.)
S, 9. Arrechavaleta, Vitoria.

(JV. Gonzalez.)
2 9. B(»jar, Salamanca.

(N. Gonzalez.)
2 (5. Silos, Burgos.
6. Castrillo, Burgos.
S, 9. Jerez, Cadiz.
9. -^Icoy, Alicante.

(N. Gonzalez.)
6. Barracas, Castellon.

(N. Gonzalez.)
9. Panticosa, Huesca.

{N. Gonzalez.)
6, 9. Sau Cristobal, Minorca,

Balearic Islands.
f, 9. Biarritz, Basses-Pyrenees,

France.
(5, 2 9. Agay, Var.
9. Giamutri Island, Tuscany.
1. Italy (Savi).
J, 3 ?. Palermo, Sicily.

W. B. Tegetmeier

T. A. Coward (c & p).
O. Thomas (p).

O. Thomas (p).

0. Thomas (p).

N. & S. Gonzalez (c).
N. & S. Gonzalez (c).
A. Chapman (p).
O. Thomas (p).

O. Thomas (p).

O. Thomas (p).

O. Thomas and R. I.

Pocoek (c & p).
J. F. Davison (c &. p).

G. S. Miller (c).
Genoa Museum (e).
Crisp Collection.
J. I. S. Whitaker
(c A P).

89. 1. 28. 1.

7. 12. 19. 3-C.
S. 2. 9. 98-99.

8. 2. 9. 9*-95.

8. 2. 9. 9G-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. 100.

0. 7. 1. 50-51.

G. 1. 21. 1-2.

8.8.4. 224-22G.
7. 2. 28. 3.
84. G. 3. G.
98. 10. 6. 8-11.

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858 RODENTIA

3 6. Montenegro. (L. Fllhrer.) O. Thomas (p). 5. S. 4. 7-9.

5 (5, 5 ?. Argostoli, Cephalonia, J. I. S. Whitaker (p). 8. 10. 1. 31-37.

Greece. (C. Mottaz.) 40-42.

3 (5, ? Kephissia, Athens. Hon. N. C. Roths- 8. 10. 2. 40-43.

(C. JIo«a,?.) child (p).

i, 9. Canea, Crete. A. Trevor Battve (p). S. 10. 24. 4-5.
(C. n. B. Grant.)

EPIMYS NORVEGICUS Erxleben.

1777. Mus norvegicns Erxleben, Syst. Regni Anim., i, p. 381 (Norway).

1778. AIus decumanus Pallas, Nov. Spec. Quadr. Glir. Ord., p. 91 (Europe).

1779. lf[Ms] surmolottits Severinus, Tentamen Zool. Huugaricae, p. 73

(Central Europe).
1800. iH[Ms] d[ecuvianus\ Jnjbridus Bechstein, Pennant's Allgem. Ueber-

sicht d. Vierfiiss. Thiere, ii, p. 713, described on p. 497. Walters-

hausen, Germany (melauistic specimen).
ISlG. Mus caspius Oken, Lehrb. d. Naturgesch., in, pt. ii, p. 895.

(Alternative for decumanus.)
1837. Mus hibcrnicus Thompson, Proc. Zool. Soc, Loudon, p. 52 (Rath-

friland, Co. Down, Ireland).
1857. Mus decumayvus Blasius, Saugethiere Deutschlands, p. 313.
1900. Mus norvegicus Rehn, Proc. Biol. Soc, Washington, xiii, p. 1G7,

October 31, 1900.
1908. Epimys norwrgicns Satunin, ilitth. Kauk. ilus., Tiflis, iv, Lief. 1-2,

p. 111.
1910. Mits {Epimys) norvegicus and 3/. (E.) norvegicus hibcrnicus Trouessart,

Faune jMamm. d'Europe, p. 142.

Tijpe hifalitij. — Norway.

Geographical dintrihution. — Original!}' 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 gi'eatest
breadth across lateral ridges about etiual to length of j^ai'ietal
measured along ridge ; anterior upper molar with evident
cingulum at anterior border of crown ; first lamina of ni'^ with-
out distinct outer tubercle ; terminal heel of 7/*'- 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 usually
45 to 54 mm.

E.vtcrnul diaracfers. — Size larger and form more I'obust than
in E. rattns, the tail never as long as head and body. Ear short,
extending barely or not to e\^e when laid forward, its substance
thick and opac[ue, its surface densely covered with tine short
haii-s, its form not jjeculiar. Feet more robust than in E. rattns,
but proportions of toes and number and position of tubercles the
same. In size the tubei-cles are, howe^•er, relatively smaller than

859

thosi! of E. rati Its, especially on the hind foot. Except for its
greater diameter and relative shortness the tail is essentially
like that of E. raffHs, 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 tlie less abundance of grooved bristles
and the excessive slenderness of such of these hairs as occur.

Mammie

: p

a 1-1; / 3-3=12.

Colour. — Upper parts a coarse, variable mixture of dull
ochraceous-bufi' and slaty grey, the back darker than the sides anc 1
heavily " lined " (though rarely clouded) with black. Occasionally
there is a decided tinge of russet on posterior half of back.
Underparts greyish white, usually with an inconspicuous wash
of cream-buft". 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.

SIlIiU. — Brain-case narrow and almost sub-cylindrical in aspect,
the lateral ridges evident but not greatly developed ; width of

KlG. 174.
Epimys nnrveijicus. Nat. size.

brain-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. rattufi ; width of basioccipital along
auterioi- border dccidediv more than half median lenirth.

S60

Rostrum deepened anteriorly so that dorsal profile is nearlj-
horizontal throughout. Upper border of plate forming outer
wall of infraorbital foramen nearly horizontal and very abruptly
rounded off anteriorly, the plate wider in proportion to its height
than in Epimys rattus. In other respects the skull closely
I'esembles that of E. rattus apart from its larger general size.
The auditory bulla; tend, however, to be less inflated and the
incisive foramina to be slightly shorter and wider, their posterior
border usually not extending quite to level of antei'ior molar
mot.

Teeth. — Incisors as in E. rattus. Molars with number of
roots and general size and proportions of crowns as in E. rattus,
liut upper teeth showing a greater tendency toward reduction
nf tlie 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 ^3, which is so reduced
as to be practically nothing more than
an outward prolongation of the outer
border of t'2, usually without trace
of antero-external re-entrant angle ;
second and third lamina essentially
as in Epimys rattus. Second upper
molar like that of E. rattus except
that /9 is greatly reduced in size and
completely merged in outer base of
/''^, tlie thii'd lamina thus forming a simple biconvex terminal
lieel. Third upper molar essentially as in Epimys rattus, t\
equally well developed, but other elements of tooth less distinct,
liower molars differing from those of Epimys rattus in the less
evident tendency to develop supplemental tubercles at outer
extremities of spaces between the lamime, such rudiments of the
small tubercles as may be present usually appearing as slight
thickenings at outer extremities of the laminR".

Measurements. — Adult male from Cortachy, Forfar, Scotland :
head and body, 22G ; tail, 179 ; hind foot, 41-^) ; ear, 20. Adult
male from Cambridge, England : head and body, 242 ; tail, 185 ;
hind foot, 42. Adult male from Tiverton, Devonshire, England :
liead and body, 238; tail, 212; hind foot, 42; ear, 21. Adult
male from Roydon, Norfolk, England : head and body, 2-50 ;
tail, 218 ; hind foot, 45. Adult male from Hampton, neai'
London, England : head and body, 255 ; tail, 187 ; hind foot, 41 ;
ear, 20 • 5. Adult male from Kilmanock, ^yaterford, Ireland :
head and body, 242 ; tail, 209 ; hind foot, 44 ; ear, 20. Adult
male and female from Upsala, Sweden : head and bod\', 250 and
252 : tail, 18.3 and 183; liind foot, 41 an.l 41. Adult male and

J-p!

861

femalo from Palenno, Sicily : liead and body, 242 and 222 ; tail,
200 and 18G ; hind foot, 45 and 43. For cranial measurements
see Table, ji. (S62.

Specimens cxami)icd.--One hundred and thirty-five, from the following
localities : —

Scotland : Stockbriggs. Lanarkshire, 2 ; Cortaohy, Forfar, 2 (Wilson).

England: Doncaster, 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, IMiddlesex, 3 ; Tiverton, Devonshire, 1 ; Northlew'
Devonshire, 1 ; Lundy Island, Devonshire, 2 ; Isle of ]\Ian, 1.

Ikeland: Castle Hamilton, Cavau, 1; Ballaghmoon,' 1 ; Kilmanock,
Wexford, S ; Inch, Wexford, 1 ; Stokestown, Wexford, 1 ; Clare Island, ]Mayoi
2 ; Inishmore, Arran Islands, 2 ; Co. Wicklow, 1 ; Castle Hamilton, 1.

NoBWAY : Holme, Mandal, 6.

Sweden : Upsala 8 (U.S.N.M.) ; Stockholm, 1 (U.S.N.M.).

Prance : Trinity, Jersey, 1 ; St. Martins, Guernsey, 1 ; Ktupes, Doubs,
1 ; Barcelonuctte, 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, Girondo, 2 (U.S.N.M.).

Spain: Pajares, Leon, 3; Bejar, Salamanca, G; Silla, Vallencia, 5;
Alcoy, Alicante, 4 ; Elche, Alicante, 1.

Germany: Brunswick, 1 (U.S.N.M.); Moritzburg, Saxonv 1
(U.S.N.M.); Uebigau, Saxony, 2 (U.S.N.M.); Jena, Thiiriugen,' 2;
Nuremberg, Bavaria, 1 (U.S.N.M.) ; near Konigsberg, 2 (U.S.N.M.f.

Switzerland : Neuchatel, 1 (U.S.N.M.) ; Roggweil, Thurgau, 4 ;
Bodcnsce, 1.

Italy: Palermo, Sicily, 26 (U.S.N.M.).

Greece : Corfu, 1 ; Argostoli, Cephalonia, 3.

2 6. Stockbriggs, Lanarkshire,

Scotland.
6, 9. Clare Island, Mayo, Ireland.

(A'. II. Bunting..)
6. ?. Inishmore, Aran Islands.

2 ?. East Norfolk, England.
(Dr. P. H. Emerson.)
i. Roydon, Norfolk.

6. Swithland, Leicestershire.

?. Epping Forest, Essex.
i. Tiverton, Devonshire.
i juv, ?. Lundy Island, Devonshire.
6. Kilmanock, Wexford.

( G. Barrett -Hamilton.)
6. 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.)
i. St. Martins, Guernsey,

Channel Islands.

(R. H. Bunting.)
9, Etupes, Doubs, France.

(C. Mottas.)

E. R. Alston (c k i>). 7'J. 'J. 25. 37. 87

G. Barrett-Hamilton
G. Barrett-Hamilton
Dr. E. Hamilton (p).

Rev. T. C. Hose

(c & p).
Hon. P. A. Butler

(c & V).
H. Doubleday (c&p).
T. P. Tracev (c & p).
T. A. Coward (c & p).
W. Eagle Clarke (p).

G. Barrett-Hamilton

G. Brooue (c & p).
C. H.B.Grant (c&p).
R. J. Cuuinghame

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

O. Thomas (p).

0. Thomas (p).

11. 1. 2. 142^

143.
11. 1. 2. 144-

145.

90. 3. 28. 1-2.

93. 11. 28. 1.

11. 1. 3. 41G.

4G. 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.

S. 9. 2. 6.
8. 9. 2. 22.

8. 8. 10. 99.

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MUS 80.:

6. St. GiUcs, Gard. G. S. Miller (c). 8. 8. 4. 223.

9. Bouconne, Gers. O. Thomas (p). 6. -1. 1. 39.

(^1. BobcH.)

2 6,2 9. Biarritz, Basses-Pyr6necs, J. F. Davison (c & v). ' ^ r a\\
100 ft. ' (6.6.4.1-2.

6. PajAres, Leon, Spain. O. Thomas {v). 8. 2. 9. 86.

{N. Gonzales.)
6, 9. Bejar, Salamanca. 0. Thomas (p). 8. 2. 9. 87-88.

(N. Gomalcz.)
2 6, 9. Silla, Valencia. O. Thomas (p). 8. 2. 9. 91-93

{N. Gonzalez.)
6, 9. Alcoy, Alicante. O. Thomas (p). 8. 2. 9. 89-90.

{N. Gonzalez.)
6, 9. Jena, Thliringen, Ger- Prof. K. von Bardc- 91. 4. 4. 1-2.
manv. lebcn (p).

2 (5, 2 9. Iloggwil, Thurgau, Swit- 0. Thomas (p). 2. 8. 4. 39-42

zerland.

{E. H. ZolliJwfer.)
6. Bodensee, Switzerland. O. Thomas (p.) 2. 8. 4. 38.

{E. H. ZoUilcofer.)
9. Corfu, Greece. (6\ ilfo/to^.) J. I. S. Whitakcr (p). 8.10.1.30.
2 9. Argostoli, Cephalonia. J. I. S. Whitakcr (p). 8. lU. 1. 38-39.

(C. Mottaz.)

Genus MUS Liiuueiis.

1758. Mus Linnffius, Syst. Nat., i, 10th ed., p. 59 {nm.vculiis, li}' tautonymy).
1814. Miiscidiis llafinesque, Preccs des Decouv. et travaux Somiologiques,

]}. 13 (Substitute for Mus).
1837. Leggada Gray, Charlesworth's Mag. Nat. Hist., i, p. 58G, November,

1837 (L. hooduga Gray and Mas platythrix Bennett).
1844. Dn/momijs Tschudi, Fauna Peruana, p. 178 (D. parvulus Tschudi =

]^lus vmseiilus Linnaeus. See Palmer, Index Gen. Mamm.,

p. 246, 1904).
1857. Mas Blasius, Siiugethiere Deutschlands, p. 309 (part).
1876. Nannomijs Peters, Monatsber, k. preuss. Akad. Wissensch. Berlin,

p. 480, August, 1876 {N. sefulosus Peters).
1881. Acromys Trouessart, Bull. Soc. d'Etudes Sci. d'Augers, x, p. 133

(Synonym of Drymomys wrongly attributed to Wagner. See

Palmer, Index Gen. Mamm., p. 246).
189G. Paendoconomys Rhoads, Proc. Acad. Nat. Sci., Philadelphia, p. 531,

December 8, 1896. Mns (Pseiidoconoiny.'i) proconodon Rhoads.
1900. Dryomys Philippi, An. Mus. Nac. de Chile, xiv, p. 20 (modificatiou

of Drymomys Tschudi).
1910. Mils Miller, Proc. Biol. Soc. Washington, xxiii, p. 59, April 19, 1910.

Ttjpe species. — Miis muscidii.s Linnpeus (by tautonymj').

Geot/raphical distribution. — Originally confined to the tropical
and temperate portions of the Old World ; now essentially
cosmopolitan through artiticial dispersal of tlie type species.

Characters. — In general like Epimys, but mechanical scheme
of molai's modified by the elongation of ci'own 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 ^~) and f6 ; m-' small and

864

EODENTIA

tending to disappear, in some species without trace of lirst
lamina ; upper incisor compressed, set at such an angle that a

I'pijer iUL-isiiis nf E/iimits (ri) ami Mi'n (b).
ilwh fiilargoil.

sub-apical notcli is normally cut in its outer side b\' action of
lower tootli.

Ilehtnrlis. - The genus Mils as now restricted contains the
Mus iiiiisi-iilioi group and the Asiatic and African species usually
referred to Lcg(jada. Though the species differ considerably
among themselves in details of structure the group is very
homogeneous as regards the yieculiar specialization of the
mechanics of the molars and the distortion of the first lamina
of ((/'. About twenty-live foi'uis are now known, seven of which
occLir in Europe.

KEY TO THE EUROPEAN FOIOIS OF MUS.

Hind foot broad and robust, its width at. base of
outer toes 4 to 6 mm.
Condylobasal length of skull in adults 21 -fj to
22 '2 mm.; hind foot 17-4 to IS -8 mm.

(St. Kilda Island, Scotland) M. DiuvaUs, p 871.

Condylobasal length of skull in adults 23 to

23-'i mm; hind foot I'J to 20 ram. (Faeroes)... M. furocnsis, 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 sliull fre-
quently more than 21 mm. ; hind foot usually

17 to I'J mm. (House mice) J/, muxalus, p. 865.

General colour dark and brownish, usually
without evidoiit yellowish tinge, the belly
dusky greyish (central and northern

Europe) M- '". /-u«r»te, p. 869.

General colour light, usually with a decided
yellowish tinge, the belly bulTy greyish
(Mediterranean region) M. m. (u-uriuus, p. 871.

Mus 865

Tail noticeably shorter than head and body
(exceptions very rare) ; colour of underpacts
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 entering houses)... M. spicilcgus, 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. spkiletjiw,'^. 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) il. s. liinpanivus, p. 879.

Colour of ujiper parts brownish grey, usually

with a decided russet tinge (Portugal) ... .V. a. hmitniiicus, 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 buffy 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
rudimentary 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

postei'ioi'ly, finely tuberculate antei'iorly ; plantar tubercles
Kraall, 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 tirst 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 sylvatiriis ; 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 e(iual 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 : p 3 — 3 ; «' 2 — 2 = 1 0.

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
di'ab or dusky, not contrasting with back ; tail dull brownish
thi'oughout, occasionally lighter below than above.

Skull. — As compared with that of small European Muridfe of
other genera, Micromjis excepted, the skull of Mus musculus is
immediately recognizable by its small size. Its general form is
cliaracterized by a peculiar flatness not
found in any of the others. Dorsal
profile 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 Apodemus sj/loaiicus. Brain-case
p.,g j.^7 short-oval in outline when viewed from

Mils iiiKsciiiiis. Nat. size. above, with a slight though evident ten-
dency toward squaring behind and in
front. Interparietal rather large, its area nearly e(iual to that
of one parietal, its genei'al form ligulate, the anterior border pro-
jecting forward at middle, the lateral extremities almost scjuarely

MDS 867

ti'iiucate ; lambdoid suture passing along anterior border of inter-
parietal so th.at the bone lies entirely within occipital ; '' sides
of brain-case abruptly rounded off or faintly angular, never
beaded. Occijjut broad and low when viewed from behind, but
central jiortion 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 Apochymiis
fsylvatiriis 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. Tnter-
orbital region distinctly wider than rostrum, slightly convex
laterally, the margins falling away scjuarely 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 anteorbit-fil 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 slightl}' beyond lachrymal level, and
usually a little exceeded by nasal branches of premaxillaries.
Incisive foramina very long (nearly equal to diastema), extending
from about 1 • 5 mm. behind incisors to level of middle of crown
of vi^ ; their greatest width is at I'egion of maxillo-premaxillary
suture ; jjosteriorly 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 m-^, 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 bulht*. Mandible rather short and deep as
compared with that of Apodentus si/lvatlcus, 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 Apodemus 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

ablv backward on outer side. Main axis of sliafts diverging
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 mai'gin 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 vr and m^ together, its area nearly 1 }j times as
great ; anterior border of ci'own 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 15 and /6 ; tS well developed, nearly
equal to ti or /6, but not so large as /I, its separation from t2
marked by an evident re-entrant extending almost to base of
crown ; tl separated from 12 by a broad, flattened or slightly
concave area which imparts to the antero-internal border of the
crown a characteristic oblicjuely-trun-
cate appearance ; second lamina more
i-egularly crescentic than first, but
^\'ith evident traces of the same dis-
tortion ; third lamina represented by
well developed /8 and <9 essentially
similar to to and /6, but with no
trace of tl 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-tei-ete tl about as large
as the outer tubercles of vi^ ; second
lamina with ti slightly larger than
id and connected posteriorly with
<8 as in the preceding tooth; third lamina about as in m',
but td reduced, decidedly smaller than /G, though marked
off from ^8 by an evident re-entrant angle. Third upper
molar relatively smaller than in any of the other European
Muridse, 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.

Mas mu^culus. t'heek-teetli. X 10.

(Antero-extenial re-entrant antrle

of nil usually much less evident.)

As in specimen represented in fig. 178.

Mus 869

true outer tubercle on first lamina ; secoiid and third laniiu;«' 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 tliose of anterior lamina, and
terminal heel greatly reduced. Third lower molar with crown
area eijual 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 vr.

ReviarJiS. — Although occasionally straying to considerable
distances from houses, Mus muftcuhts is characteristically an
urban species, and its spread from central Asia to Europe 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 Linni¥us.

1758. [Mils'] niusculiis LinriEeus, Sysfc. Nat., i, 10th ed., p. 62 (Sweden).

1801. i^f[Ms] m[usculus] albus Bechstein, Gemeimi. Naturgesch. Deutsch-
lands, I, 2d ed., p. 955 (Thiiiingen, Germany).

1801. JiT[us] m[iisculHs] flavus Bechstein, Gemeinn. Naturgesch. Deutsch-
lauds, I, 2d ed., p. 955 (Thiiringen, Germany).

1801. M{iis\ ni[iisculus] maculatiis Bechstein, Gemeinn. Naturgesch.
Deutschlands, I, 2d ed., p. 955 (Thiiringen, Germany).

1801. M[iis] ■m[nscuiiis'] niger Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2d ed., p. 955 (Thiiringen, Germany).

1827. [Mus musculus] 0 striatus Billberg, Synopsis Faunae Scandinavise,
p. 6 (Skane, Sweden).

1827. [Mus musculus] y albicans Billberg, Synopsis Fauuaj Scandinaviae,

p. 6 (Skilne, Sweden).
1827. [Mus musculus] S niveus Billberg, Synopsis Faunae Scandinaviae,

p. G (Molle, Norway).

1837. ? Mus brevirostrisWtLterhonse, Proc. Zool. Soc, London, p. 19 (Mal-

donado, Uruguay).
1844. 7Drymomys jMrvulus Tschudi, Fauna Peruana, p. 178 (Central Peru).
1857. Mus musculus Blasius, Siiugethiero Deutschlands, p. 320.

1867. [Mus 7HUsculus] lielvolus Fitzinger, Sitzungsber. kais. Akad. Wis-
sensch. Wien, Math.-Naturwiss. Classe, lvi, pt. i, p. 70
(Hungary).

1867. [Mus musculus] varius Fitzinger, Sitzungsber. kais. Akad. Wissensch.
Wien, Math.-Naturwiss. Classe, lvi, pt. i, p. 70 (Europe).

1807. [Mus musculus] cinereo-maculatus Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, IMath.-Naturwiss. Classe, i.vi, pt. i, p. 70

(Europe).
18C9. Mus poschiavinus Fatio, Fauue Vert. Suisse, i, p. 207 (Poschiavo,

Grisons, Switzerland).

870 RODENTIA

1872. Miis musculus var. flavcsccns Fischer, Zool. Garten, xiii, p. 223,

July, 1872 (Berlin, Germany).
1910. Mus musculus and M. musculus poschiavinus Trouessart, Paune

Mamm. d'Europe, p. 145.

Type locality. — Upsala, Sweden.

Geographical diiifrihution. — Europe north of the Mediterranean
region.

Diagnosis. — General colour dark and browni.sh, 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 (I7-6-19-4).
For cranial measurements see Table, p. 872.

Specimens examined. — One hundred and one, from the following
localities : —

Iceland : Near Reykjavik, 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) ;
Loudon, 3 (B.^f. and U.S.N.M.) ; Richmond, Surrey, b (U.S.N.M.) ; Wands-
worth Common, Surrey, 4; Martock, Somerset, 1 ; Newport, Isle of Wight,
1 ; Ramsay, Isle of ]\Ian, 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. I\L) ; Holme, JIandal, 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.JL).

Switzerland : Andermatt, Uri, 1 (U.S.N.JI.) ; 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, ?. Clare Island, Mayo, Ire- G. Barrett-Hamilton 11. 1. 2. 146-148.

land. (R. H. Bunting.) (p).

6, ?. Clifden, Galway. G. Barrett-Hamilton 11. 1. 2. 149-150.

{B. H. Bunting.) {v).

2 6. V. Oundle, Northampton- Lord Lilford (p). 11. 1. 1. 166-168.

shire, England.

6. Wandsworth, Surrey. J. Miller (c & p). 4. 2. 12. 1.

9. Martock, Somerset. C. B. Horsbrugh 11. 1. 3. 420.

(c & p).

6. Newport, Isle of Wight. R. J. Cuninghamc 11. 1. 3. 421.

(c & p).

2 6, ?. St. Ouen's Bay, Jersey, O. Thomas (p). 8. 9. 2. 7-9.

Channel Islands.

(E. H. Bunting.)

MUS

871

1 al. Alderney, Channel Is- W. Eagle Clarke (p). 98. 9. 20. 7.
lands.
2 c5, 3 ?. Holme, Mandal, Nor- R. J. Cuninghame 8. 8. 9. 14-18.
way. (c & p).

6. Unfcervatz, Grisons, 0. Thomas (p). 4.4.5.45.

Switzerland.

(E. H. ZolUkofer.)
3 <^, 9. St. Gallen. 0. Thomas (p). 4. 4. 5. 41-44.

(E. H. ZolUkofer.)

Mus MUSCULUS AZORicas Schinz.

1845. M[us] asorlcus Schinz, Synops. Mamm., ii, p. 161 (Azores).

1355. Mitsculus moUissimus Dehne, Allgem. deutsche Naturhist. Zeitung,

Dresden, neue Folge, i, p. 443. (Monte Pollino, Basilicata,

Italy).

Type locality. — Azore Islands.

Geographical distribution. — Mediterranean region ; Azores.

Diagnosis. — General colour less dusky and more yellowish
than in M. musculus musculm, 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.

Specimens 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; B6jar, Salamanca, 3;
Dehesa de Valencia, Valencia, 1 ; Elche, Alicante, 2 ; Venta del Baul,
Granada, 5 ; San Cristobal, Minorca, Balearic Islands, 1.

France : Biarritz, Basses-Pyr6n6es, 2 ; Ax-les-Thermes, Arifege, 1 ;
I'Hospitalet, Ari^ge, 1 ; Porte, Pyr6n6es-Orientales, 1 ; near Nimes,
Gard, 8 ; St. Gilles, Gard, 1 ; St. Pai^. Basses-Alpes, 6 ; La Poce di
Vizzavona, Corsica, 1.

Italy : Acceglio, Cuneo, 1 ; Calappiano, Empoli, 3 ; 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, Sicilv, 10
(U.S.N.M.) ; Lanusei, Sardinia, 1 (U.S.N.M.).

Greece: Corfu, 22; Patras, G; Athens, 4 (U.S.N.M.); Argostoli,
Cephalonia, 1.

5 6, ?. San Miguel, Azores. Hon. W. Rothschild 3. 6. 5. 17-22.
(W.B.Ogilvie-Grant.) (p).

S. Estoril, Portugal. 0. Thomas (c & p). 98. 2. 2. 36.

6, 9. Cintra. O. Thomas (c & p). 98. 2. 2. 28-29.

6, 9. Pajares, Leon. O. Thomas (p). 8. 2. 9. 133-134.
(N. Gonzalez.)

9. Silos, Burgos. G. S. Miller (c). 8. 8. 4. 109.

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EODENTIA

(5, 2?.

Bejar, Salamanca.

(N. Gonzalez.)

0. Thomas (p).

8. 2. 9. 136-138

i.

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.

Ct. S. Miller (c).

8. 8. 4. 110-112

?.

Venta del Baul, Granada.
(N. Gonzalez.)

0. Thomas (p).

8. 2. 9. 130.

i.

San Cristobal, Minorca,

0. Thomas and R. I.

0. 7. 1. 58.

Balearic Islands.

Pocock (c & p).

?.

St. Gilles, Gard, France.

G. S. Miller (c).

8. 8. 4. 222.

4 9.

Nimes, Gard. [C. Mottaz.

) 0. Thomas (p).

8. 8. 10. 93-96.

<5, 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.

(5.

Marsala, Sicily.

(.•1. Robert.)

0. Thomas (p).

6. 8. 4. 44.

3 i, 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 (J, 4 9.

Corfu, Greece.

(C. Mottaz.)

0. Thomas (p).

8. 10. 1. 43-48.

3 J, 2 9.

Corfu, Greece.

C. Mottaz (c).

8. 11. 3. 12-16.

2 i, 9.

Patras, Greece.

Hon. N. C. Roths-

8. 10. 2. 44-46.

(C. Mottaz.)

child (p).

<i.

Argostoli, Cephalonia.

C. Mottaz (c).

8. 11. 3. 17.

MUS MURALIS Barrett-Hamilton.

1899. Mas mnralis Barrett-Hamilton, Proc. Zool. Soc, London, p. 81.

Type in British Museum.
1910. Mas niusculus mnralis Trouessart, Faune Mamm. d'Europe, p. 146.

Tijj.ic locaJlty. — Island of St. Kilda, Scotland.
Geographical distribution. — Island of St. Kilda.
Diagnosis. — Similar to Mus mmculus, 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.

E.derital characters. — Except for the less slender
tail and more robust feet the external characters do
not differ appreciably from those of 3Ihs mH.'<cuhis.

Colour. — The colour is essentially as in Mus
miisculits, 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 narrower
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 immature speci-
mens than in atlults, is constantly diflerent from that charac-

Mus 875

teristic of the large series of European specimens of Mux inim-idus
examined. Teeth as in Mua nmsculus.

Measurements. — External measurements of type (adult
female): head and body, 90; tail, 85; hind foot, IT" 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).

9. St. Kilda, Scotland. G. Barrett-Hamilton (p). 8. 7. 16. 1.

(H. Evans.) (Type of species.)

6, 9 al. St. Kilda. J. E. Harting (ip). 94. 7. 16. 2-3.

1 al. St. Kilda. Kelvingrove Museum, 96. 0. 6. 1.

Glasgow (p).

2 <£, 3 9. St. Kilda. W. Eagle Clarke (c & p). 11. 1. 24. 7-11.

MUS FiGROENSIS Clarke.

1904. Mus musculus fxroensis Clarke, Proc. Roy. Phys. Soc. Edinburgh,

XVI, pt. II, p. 163. Type in Royal Scottish Museum.
1910. Mus miisculus fxroeyisis 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. niusculus.

E.rternal characters. — Except as already pointed out the
externa] characters agree with those of Mus musculus.

Colour. — The few specimens examined indicate that the colour
does not differ essentially from that of Mus imisculus.

Shdl 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
anteriorl}', and the pterygoids, though imperfect, evidently not
far from parallel. The skull from Myggenivs, however, as
pointed out by Dr. Winge 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 niusctilus, 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 ;

S76

EODENTIA

Observations.

Teeth moderately worn.

„ slightly worn.
,, moderately worn.

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8

M. muralis.

Scotland : St. Kilda Island

M. faeroensis.

Faeroe Islands : Nolsoe

Myggeuips

MDs 877

liiiid foot, 20; ear, 1:5. Adult feinalo from the tyjie locality:
hind ft)ot (in alcohol), I'J'cS. For cranial mcasurcnionts .see
Table opposite.

Sjiecimcns examined. — Six, three from Nolsoe (Edinburgh aud Copen-
hagen), and three from Myggenfes (Copenhagen;.

Bemarhs. — The Faeroe house mouse ditlers 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 species. The enlarged extremities may be,
as suggested by Dr. Winge, the result of changed habits due to
a climbing life among the cliti's 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 feeroensis is likely to be
confused is Mus vmralls 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 dai'ker than in Mus ')imralis, 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.)

Oeograpliical distribution. — Eastern and southern Europe,
west to Portugal, Hungary and Sweden.

Diagnosis.-— >iize less than in Mus )nusculus, the head and
body rarely attaining a length of 90 mm., the hind foot in adults
ranging about from 15 '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 j^ellower races the colour is strikingly
different from that of Mus musculus. 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. — 3Ius spicilcgus 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 centi'al Asia. Among the
European representatives of the species three races may be
distinguished, one of which ranges somewhat further north than
the others.

878

KODENTIA

Mus spiciLEGUS SPiciLEGUS Petenyi.

1882. Mus spicilcgus Petenyi, Termeszetrajzi Fiizetok, BudaxJest, v, p. 114.
1882. Mus acervator Petenyi, Termeszetrajzi Fiizetek, Budapest, v, p. 114

(alternative name).
1882. Mus acervif»;x Petenyi, Termeszetrajzi Fiizetek, Budapest, v, p. 114

(alternative name).
1882. Mus canicularius Petenyi, Termeszetrajzi Fiizetek, Budapest, v,

p. 114 (alternative name).
1882. Mus caniculator Petenyi, Termeszetrajzi Fiizetek, Budapest, v,

p. 114 (alternative name).
1896. Mus musculus flavesccns Barrett-Hamilton, Zoologist, 3d. ser., xx,

p. 179, May 1896 (not Mus musculus Mir. Jiavcscens Fischer, 1872).
1910. Mus sjiicilcgus Trouessart, Faune Mamm. d'Europe, p. 148.

Type JocalUy. — Hungary.

Geogrcqjhical disfribuiion. — 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. — Ujjper parts a light wood-brown, becoming some-
what bufify 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 colovu*
of underparts. Basal portion of hairs slate-grey. Underpai'ts
rather sharply defined bufiy white or i)ale cream-buff, the slate-
grey bases of the hairs producing a slight greyish cast. Feet
and under surface of tail es.seutially concolor with belly ; upper
surface of tail brownish without sharp line of demarcation.
Ears sprinkled with brownish hairs externally, with huffy
internally, neither surface with any decided colour.

Measurements. — Average of seven adults from Ujisala,
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 : liead and body, 75 ; tail, 60 ; hind foot, 16 ; ear, 12.
Average and extremes of five adults from Patrily, Pioumania :
head and body, 77-5 (72-83); tail, 6-21 (59-5-70) ; hind foot,
16-2 (16-16-5); ear, 12 (11-5-13 5). For cranial measure-
ments see Table, p. 880.

Specimens examined. — Fiftv-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.^NL).
Austria-Hungary : Csallokoz-Somorja, Pressburg, 5 ; Hatszeg,
Hunyad, 2 ; Pester Comitat, 2.

MDS 879

RoDMANiA : Gagoni, Pi-ahova, 1 ; Patrily, 7.
Montenegro : Various localities, 5.
Albania : Various localities, G.

5. Csallokoz-Somorja, Press- Budapast Museum 9i. 3. 1. 48-52.

burg, Hungary. (e).

2 ?. Hatszeg, Hunyad, Trau- C. G. Danford (c). 3. 2. 2. 33-34.

aylvauia.
6, 9. Pester Comitafc. Budapest Museum 94 7 26 1-2

i. Gageni, Prahova, Rou- Lord Lilford (p). 4. 4. 6. 41.

mania. [W. Dodson.)
7 6. Patrily, Roumania. Lord Lilford (p). 4. 4 G 42-48

{W. Dodson.)
2cJ, 3 9. Montenegro. (L. Filhrcr.) O. Thomas (p). 5.8.4.18-22.

3(^,3 9. North Albania. 0. Thomas (p). 5. 8. 4. 38-43

{L. Fiihrcr.)

Mus spiciLEGus Hi.spANicus Miller.

1909. Mus spicilegiis hispanicus Miller, Ann. and Mag. Nat. Hist 8th ser

III, p. 421, May, 1909.

1910. Mtis spicilegus hispanicus Trouessart, Paune Mamm. d' Europe

p. 148. .

Type locality. — Silos, Province of Burgos, Spain.

Geographical distribution. — Central and southern Spain.

Diagnosis. — General colour of upper parts butty or pale butty
grey.

Colour. — Back and sides ranging from buff" to a pale butty
grey lighter and less j-ellow than the cream-buff" of Ridgway, the
median dorsal region faintly "lined" with black, the sides
gradually becoming clear buff" or butty 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, 1-4-4 ; 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 (15-6-1G-8);
ear, 12-5 (12-13). For cranial measurements see Table, p. 880.

Specimens examined. — Eighty-five, from the followiug localities : —
Spain; Veuta del Baul, Granada, 7: Elche, Alicante, 24; Alcoy,
Alicante, 21 ; Silos, Burgos, 12 ; Castrillo de la Reina, Burgos, 3 ; Pajdres]

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882 RODENTIA

Leon, 1 : San Cristobal, Minorca, Balearic Islands, 7 ; Inca, Majorca,
Balearic Islands, 6.

France : Cerbere, Pyr^n^es-Orientales, 1 ; near Nimes, Gard, 2 ;
Valescure, Var, 1. The Gard and Var sxaecimens not typical.
4 6, 9. VentadelBaul, Granada, G. S. Miller (c). 8. 8. 4. 102-106.

Spain.
4 6 2 9. Elche, Alicante. G. S. Miller (c). 8. 8. 4. 89-94.

13,^,8 9. Alcoy, Alicante. O. Thomas (p). 8.2.9.139-159.

(A''. Gonzalez.)
4 6, 9. Silos, Burgos. G. S. Miller (c). 8. 8. 4. 97-101.

(8. 8. 4. 101. Ti/pe of subspecies.)
6 9 Castrillo, Burgos. G. S. Miller (c). ' 8. 8. 4. 95-96.

'(5. PajAres, Leon. 0. Thomas (p). 8. 2. 9. 132.

(N. Gonzalez.)
4 (5, 2 ?. Inca, Majorca, Balearic 0. Thomas and R. I. 0. 7. 1. 18-23.
Islands. Pocock (p).

6 6, 9. San Cristobal, Minorca, O. Thomas and R. I. 0. 7. 1. 69-65.
Balearic Islands. Pocock (p).

S. Cerbere, Pyr6nees-0rien- 0. Thomas and R. I. 0.7.1.77.

tales, France. Pocock (p).

£ St. Genies, Gard. O. Thomas (p). 8. 8. 10. 92.

[C. Mottaz.)

Mus spiciLEdUS LUSiTANicus Miller.

1896. Mus sp. Thomas, The Zoologist, 3d ser., xx, p. 137, April, 1896.

1909. Mus spicilecjus lusitanicus Miller, Ann. and Mag. Nat. Hist., 8th ser.,

Ill, p. 422, May, 1909.

1910. Mus ."spicilegus lusitanicus Trouessart, Faune jMamm. d'Europe,

p. 148.

Tiijie forallfj/. — Cintra, Portugal.

Geographical (li.^tiibHil(>n.--K.n(>\vn only from the vicinity of
the type locality.

.7>/(((//M/s/.v.-- 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 buffy, 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 sylcatkus.

Mfasiirements. — Type (adult male) : head and body, 77 ; tail,
60; hind foot, lG-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
(1:2-13). For cranial measurements see Table, p. 881.

Specimens examined. — Six, all from the neighbourhood of Cintra.

4 i, 2 9. Cintra, Portugal. U. Thomas (c & p). 98. 2. 2. 30-35.

(98. 2. 2. 30. Type of subspecies.)

883

Genus ACOMYS I. GeofFroy.

1838. Acomys I. Geoffrey, Ann. des Sci. Nat., Paris, 2d ser., x, Zool.,
p. 126, August, 1838.

1841. Acosminthus Gloger, Gemeinn. Hand- u. Hilfsbucb der Naturgesch.,

I, p. 95 (dimidiatus).

1842. AcantJwtmjs Lesson, Nouv. Tabl. Regne Anim., Mamm., p. 185

[hispidus).

Type species. — 3Ius 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.

Reviarls. — The genus Acomys contains about fifteen species,
one of which occurs within the limits of western Europe as here
detined.

ACOMYS MINOUS Bate.

1906. Acomys dimidiatus minous Bate, Proc. Zool. Soc, London, 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 al)out one and a half rings. These bristles are of
a peculiar silvery horn-colour, and are so arranged as to form
continuous longitudinal striie, which, in connection with the
conspicuous rings, impart to the tail a peculiar lizard-like appear-
ance. Among the bristles occur numerous longer terete hairs so

3 L 2

884

RODKNTIA

slender that they can scarcely be detected without the aid of 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 buff, 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 bufl' with
the extreme tips horn-colour, the result being a peculiar coarse
grizzle in which the bufF decidedly predominates. On hind leg
the buft" 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.

Slnill. -In general appearance the skull resembles a miniature
of that of Epinii/s rnttus, 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,
liostrum rather deeper and
narrower than in other European
Muriripp. Interparietal very large,
strongly convex posteriorly, straight
or slightly convex anteriorly, its
area about efjual to that of parietal.
Incisive foramina extending back
to about middle of first molar,
their length oidy about 1 mm.
less than that of diastema. Apart from its peculiar backward
extension in median line characteristic of the genus, the bony

Fir,. 180.
Aeoiiiyx tiii'novs. Nat. size.

885

palate is very short, its leiif^tli from incisive foramen to the trans-
Terse i-i(lge liomologous with the noi-nial posterioi- bm'der scarcely
exceedini^ breadth at middle of tirst molar. Haniulars 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 buUfe moder-
ately large but not sj^ecially 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 Ephnys rattus or
Mas musculiis). Plate forming outer wall of infraorbital foramen
broad, rather stjuarely rounded oft" above. Infraorbital foramen
rather widely oj-ien 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 I'oot of lower incisor slight.

Teeth. — Upper incisor strongly compressed, I'ather more so
than in Mus musculus, the cutting edge normal. Lower incisor
essentially like the iqiper tooth
in cross section, its anterior and
posterior diameters nearly equal.
Molars suggesting those of Mus
7nmculus 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 m'^
combined. First upper molar
with crown relatively wider
than in 3Ius musculus, but
with <1 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 irr ; t'i usually
represented by a minute antero-external tubercle ; second and
third laminae rather distinct, oblique, separated by a narrow
groove, the extremities of which produce a slight re-entrant
angle on each margin of crown, the tubercular elements entering
into formation of the laminw 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 ?H^

Fig. 181.

Acomns minou!'. Clieek-teeth,

somewhat worn, x 10.

886

SPALAX 887

Mns musculus, but anterior portion of m^ broader and witli
two sub-equal tubercles nearly as large as those oi second lamina,
and (H.J relativelj' larger, its crown area slightly more than half
that of )«2-

Measurements. — 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 : —
Crete : Canea, 3.

2 6, 9. Canea, Crete. Hiss D. Bate (c). 5. 12. 2. 31-33.

(5. 12. 2. 33. Type of species.)

Family SPALACID^.
1821. Spalaciih'. Gray, London 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, for»vard-sloping occipital shield,
the outer margin of which extends to level of outer border of
zygomatic root ; jugal short, extending forward only a little
bej'ond 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 ^j, rooted, hypsodont, the subterete crowns with one inner
and one or two outer re-entrant folds.

Remarl's. — Both externally and in the structure of the skull
the family Spalacidse 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 Spalax.

Genus SPALAX Gueldenstaedt.

1770. Spalax Gueldenstaedt, Nov. Comm. Acad. Sci. Imp. Pefcrop., xiv,

pt. I, p. 410 {rnicrophthalmus).
1783. Mijospalax Hermann, Tabula Affinitatum Animalium, p. 83 {lax-

vianni Hermann = Spala.r inicrophtltalnius Gueldenstaedt).
1799. Talpoidcs Lac6pede, Tableau des divisions, sous-divisions, ordres et

genres des Mammiferes, p. 10 {typhlus).
1804. Aspalax Desmarest, Nouv. Diet. d'Hist. Nat., xsiv, Tab. Jileth.

Mamm., p. 24 (Mus typhlus Pallas).
1815. Anotis Rafinesque, Analyse de la Nature, p. 68 (Substitute for

Talpoides Lac6pede).

8R8

rodp:ntia

ISIO. ? Ommatosteigus Nordmanu in Keyscrliiig aud Blasius, Wirbelthicre
Europas, j)p. vii, 31 (0. imllasii Nordinann). Nomen nudum.

IS'JH. Microspalax Nehring, Sitz.-Ber., Gcsellscli. Naturforscli. Freundc,
Bei-liii, for December 21, 1897, p. 168 (Provisional sub-generic
name for smaller species of Spalar). Not Microspalax Trouessart,
1885.

1908. Nannofipalax Palmer, Science, N.S., xvii, p. 873, Tklay 29, 1903
(Substitute for Microspalax Nehring).

Type f^pecics. — Spnlax iiiicrophthaliiniti Gueldenstaedt.

Geocp-aphicnl dintrlbution. — South-eastern Europe and adjoin-
ing portions of Asia ; south into Egypt. Limits of range
imjjerfectly known.

Characters. — Essential characters as in the family Spalaridie.
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 tlie teeth wear away,
undergoing considerable changes in form, the re-entrant folds
tending to become isolated as enamel islands (fig. 184, p. 893).

EeinarJcs. — The members of the genus Sjjalax 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 su})p(ised to occur in Greece.

KEY TO THE EUROPEAN SPECIES OF SPALAX.*

Skull not unusually deep, the distance from alveolus
of HI/' to sagittal crest about 39 per 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. dolbioge;r, p. 889.

Skull unusually deep, the distance from alveolus of
m^ to saoittal crest about 43 per cent of condy-
lobasal length ; width of auditory bulla about
8 ram. ; greatest width of infraorbital foramen
conspicuously more than half height.
Breadth of rostrum in adult male about 11 mm. ;
alveolar breadth of «!■' equal to least palatal

width (Hungary) .S'. hungaricus, p. 894.

Breadth of rostrum in adult male about 12 mm. ;
alveolar breadth of «i' less than least palatal
width (Greece ?) S. gmrus, p. 895.

* The important work by Dr. L. M6hely on this genus— Species generis
Spalar. A Fiildi Kutyak Fajai, Budapest, 1909— has unfortunately been
received too late to be utilized by the author.- 0. T. (See Appendix,
p. 1000.)

889

SPALAX DOLBROGEiE Milk'r.

I'JOl. iSjxilax liuncjariciis Matschie, Sitz.-Ecr. Gesellsch. Natuiforscli.

Freunde, Berlin, p. 237, November, I'JOi (Roumania).
1903. Spalax dolbro(je:f IMiller, Proc. Biol. Soc, Washiugtou, xvi, p. 161,

November 30, 1903 (Dobrndscha). Type in U.S. National

Museum.
1910. Spalax dolbrogen' Trouessart, Faune INIamm. d'Europe, p.' 204.

Type locality. — Malcoci, Dobrudscha, Roumania.

GeiigrapJiical distrihntifin. — Koumania and Bulgaria ; limits of
range not known.

DiiK/iums. — .Size medium (hind foot about 2r) nnii., condylo-
basal length of skull about 50 mm.) ; skull moderately deep, the
distance from alveolus of m'-^ 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 bulUe rather long and narrow ; breadth of h«' 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 pi'ofile 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 ovitline, the
simple, round nostrils situated a little below middle ; a narrow
groove curving outward and upward from each nosti'il to anterior
margin of pad ; lip 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 furrow on side next thumb, the anterior
margin raised and thickened, sometimes appearing as a ridge

890 RODENTIA

crossing palm oblicjuely 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 others, 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 ; uuder-
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 ; occijiital 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-likfe 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 scjuarish 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 891

nearly twice length excluding condyles, width along anterior
suture about two-thirds median length ; surface of basioccipital
irregularly depressed at side; auditory bulla? rather large, but
displaced upwai'd so that lower Ijorder is about on level with
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 dolbrogese. 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 zj^gomatic 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

RODENTIA

than lialf height. lli)strum ratlier long, nearly parallel-sided
wlien \ie\ved from abo\e, though slightly wider somewhat
beyond middle than at base, and tapering rather, noticeably in
front, its width everywhere greater than that of interorbital
constriction; nasals wide, abruptly rounded off anteriorly, where
they project so as to conceal incisors, tapering gradually to a
narrow base, where they are abruptly truncate at level of
posterior extremity of nasal branches of premaxillaries, and of
middle f>f 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 sj^ace in front of them, twice in
that behind). Palate very narrow, its least breadth scarcely
eijual to that of alveolus, 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
Ijehind that of alveolus of m^ ;
hamulars simple, straight, slightly
diverging, widely separated from
bulhe ; ectopterygoid narrow, the
pit deep and widely open at bottom.
MandiT)le slender, appearing dispro-
portionately small for the very large
incisor which projects conspicuously
behind as well as in front, and to
wliich the ramus is little more than
a sheath ; coronoid process high,
slightly recurved, its anterior border rising abruptly from level
of front of }ii.^ ; 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 outwai'd 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 equal to greatest antero-posterior diameter ; posterior

S/talax di'Wroiji'ie. Skull fiiiiu
behiiiil. Nat. size.

SPALAX '^O^

border narrowly rounded ; inner and outer boi'ders about equal ;
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 (lolbrogeie. Cheek-teeth showing enamel pattern at various stages of wear.
I'pper 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 longitvidinally 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 jiattern of m^ and in'^ consisting of two
backwardly curving re-entrant folds on outer side and a single

894

EODENTIA

foward-curving re-entrant fold on inner side, the anterior outer
fold of VI- shallower than the others and usually becoming
converted into an "island" at an early stage of wear; m^ 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 wi^ and m.^ with
a small supplemental basal 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.
Mrasuremenfs. — 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 Rustsehuk, 6 skulls (Andersen).
Roumania: Malcoci, Dobrudscha, 3 (B.M. and U.S.N.M.).
6, ?. Malcoci, Dobrudscha, Rou- G. Dalgleish (c). 5. 10, 25. 1-2.
mania.

SPALAX HUNGARICUS Nehring.

1898. Spalax tyyhlus hungarieus Nehring, Sitz.-Ber. Gesellsch. Natur-

forsch. Freunde, Berlin, for December 21, 1897, p. 173, fig. 8.
1898. Spalax hungarieus Nehring, Zool. Anzeiger, xxi, p. 479,

September 5, 1898.
1898. ? Spalax monticola Nehring, Sitz.-Ber. Gesellsch. Naturforsch.

Freunde, Berlin, p. G (Ulog-Obruja, Herzegovina).
1910. Spalax hungarieus and ? S. monticola Trouessart, Faune Mamm.

d'Europe, pp. 203, 204.

Tyye locality. — Hungary ; exact locality not stated.
Geoyra-pMcal distrihution. — Hungary ; probably extending
southward into north-western portion of Balkan Peninsula.

FIG. 185.

Aiitc'iior liase of zygoma in Spaltix tlalbnyne (a)

. and S. hungarieus (6). Nat. size.

SPALAX , 895

Diagnosis. — Like Spalax dolbrogese, but skull tending to be
somewhat deeper ; anteorbital foramen moderate, nearly as wide
as liigh ; anterior base of zygoma heavy ; auditory bullae rather
short and wide ; breadth of m^ 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. O. Thomas (p). 3. 1. 14. 1-2.

(L. Fiihrer.)

SPALAX GR^CUS Nehring.

1898. Spalax gracus Nehring, Zool. Anzeiger, xxi, p. 228, March 21, 1898.

Type in Munich Museum.
1910. S^mlax gr.rcus Trouessart, Faune Mamm. d'Europe, p. 204.

Type locality. — 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 Spalax hungaricus, but skull with
somewhat broader rostrum and palate.

E.ciernal characters and colour. — As in S. hungaricus.

Shill and teeth. — In all respects the skull and teeth resemble
those of Spalax hungaricus, as opposed to S. dolbrogese (great
depth tlirough frontal I'egion ; form of auditory buUse ; 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.

RemnrJcs. — Through the kindness of the authorities of the
Munich Museum I have had the oj^portunity to compare the
original specimens of Spalax grsecus with S. hungaricus and S.
dolbrogene. 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 sj^ecitic. 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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Family SCIUUID^.

18:21. Sciurichc. Gray, London Med. Kopos., xv, p. 301, April 1, 1821.

Geograpldcal distrihution. — Eastern uTid western hemispheres
except at the extreme north and south ; absent from Madagascar,
New Guinea, Austraha and the Pacific Islands.

Chantctcrs. — Form slender or robust according as the animal's
habits ai'e arboreal or terrestrial ; tail without horny scales on
lower surface, densely long-haired, usually flattened ; ribs twelve
or thirteen pairs ; skull varying in form but never highly
modified ; postorbital processes present ; infi'aorbital foi'ameu
small ; jugal long, its anterior extremity wedged between
lachrymal and zygomatic pi'ocess of maxillary ; palate rather
broad ; upper incisor with root in front of anterior cheek-teeth,
lower incisor with root -not extending conspicuously into
ascending jwrtion of ramus ; molars rooted, tubercular and
transversely ridged, premolars j or }-.

HevtarJcs. — The family Scmridse, containing the squirrels,
spermophiles, marmots and their allies, is one of the most widely
distributed and abundantly represented group of rodents. It
contains 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 SCIURID^^.

(For convenience of comparison the cranial and dental cbaracters of
Sciuroptcrus arc 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 Scluropterns, p. 911.]

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 isarallel.
Incisors compressed, the width of anterior face not

more than half that of lateral face ; small

upper premolar minute, 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 compressed, 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 Ji

898 KODENTIA

Brain-case not excessively flattened, 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 condj-lobasal
length of skull ; size largo, form badger-
like (Marmots) Marmola, p. 931.

Genus SCIURUS LiniK.ni.s.

1758. Sciurus Linuteus, Syst. Nat. i, 10th ed., p. 63 (vulgaris, by

tautonymy).
1867. Sclunis Blasius, Saugethiere Deutschlauds, p. 271.
1893. Aphrontis Schulze, Zeitschr. fiir Naturwissensch., Leipzig, lxvi,

p. 16.5 {vulgaris).

Ti/pe xpccics. — Sciurus vul/jari'.s Limia'u.s.

(ji'dtjrapliical (ii.strihutiov. — Foi'ested rcf^dons of tlif northcni
liemisphere, in the Old World from Ireland to Japan and north-
eastern Siberia ; southern limits of distribution at present
uncertain owing to lack of pi'ecise definition of the genus.

Characters. — Strictly arboreal Sciuridie, of medium size and
typically sciurine aspect, the tail bushy and conspicuously
Hattened, more than half as long as head and body ; skull witli
deep, sti'ongly convex Ijrain-case and weak rostrum ; postorbital
processes slender, directed backward ; incisors strongly com-
pressed ; anterior upper premolar vestigial, usually present as a
practically functionless terete spike ; crowns of molars low, those
of u])per teeth with two moderately developed cross ridges and a
broad, low inner cusp (not forming a U^^t^'^'P^d pattern with
wear), those of lower teeth distinctly basin-shaped.

Ii(')uarl:s. — In the present uncertainty regarding the classifica-
tion of the squirrels, it is impossible to frame a satisfactory
diagnosis of the genus Sciuruti, 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 ivh'cs.

SCIURUS VULGARIS Linn;eus.
(Synonymy under subspecies.)

Geographical diiitrihutio)i.- -'Wood&\ portions of Europe from
the extreme northern limits of ti-ee growth to the Mediterranean
coast, and from Ireland eastward into Asia.

Diagnosis. — Size medium (head and body i-anging in the
different races from 205 to 280 mm., tail from 160 to 242, hind
foot from 55 to G6, condylobasal length of skull from 45 to 55) ;

sciuEus >^99

tail (Icc'idtiilly inoro tliaii half as long as head and body ; car
conspicuously tufted in winter ; colour reddish, greyish or dark
brown above, whitish (never distinctly yellowish) below ; minute
anterior upj^er premolar present ; cross-ridges on U2:)per 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
Cijual to head and body. Head rounded posteriorly, narrowed in
front, the muzzle deep and not much produced (distance from eye
to nostril about equal to depth at I'egion half way between) ; ear
rather large, just covering eye when laid forward, the A\'idth at
middle about three-quarters height from crown ; anterior bortlei-
nearly straight below, somewhat convex above, its lower two-
thirds abrujjtl}^ folded backward, the inner root of this fold lying
uearly over meatus ; tip rather abruptly rounded ofl"; posterior
border convex except for a flattened or slightly convex region
just below tip, its lower border extending forward as an abruptly
angled j^i't'jection 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 lingers ; thumb reduced to a mere
tubercle with closely appressed nail ; two middle lingers sub-equal
and longest, second and lifth svib-ecjual and about half as long ;
palm with Ave tubercles, the two posterior lai'gest and best
deflned ; claws long, compressed, strongly curved. Hind foot
long, the Ave toes all well developed and bearing strong claws
shorter and less curved than those of fore foot ; inner toe
extending about to base of others, outer extending a little beyond
middle of foui'th ; fourth toe longest, third and second successively
a little shorter ; sole densely hairy in winter, jjartly bare in
summer ; four small but evident tubercles at bases of toes. Tail
strictly tlistichous throughout, the hairs at sides and above about
30 to 50 mm. in length. Mammae, p 1 — 1, a 2 — 2, i 1 — 1 = 8.

Colour. — Though the actual colour shows great diflerences
according to phase and season the pattern is constant and
charactei'istic. The head, back, sides and tail ai'e essentially
uniform, the tail usually a little darker than back, the sides of
body and outer surface of limbs somewhat lightei', the middle of
back fi'equently with more I'ed ; face with an ill-defined j^ale area
on cheeks and muzzle ; ear-tufts usually concolor with sur-
rounding parts but sometimes dai'ker or more red ; inner surface
of legs and upper surface of feet not so dark as outer surface of
legs ; tail with a lighter, grizzled or whiti.sh area along median
line below, this sometimes pai'tly or wholly concealed ; white of
iniderparts extending forward to chin or to back of interramial
space; posteriorly the white does not reach base of tail, and

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 pi-edominance 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 gener'al 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 moi'e greyish than the summer
pelage. In regions where both colour phases occur four normal
combinations of phase and pelage will be found in any consider-
able series of specimens. Apart from these combinations and
their intermediate stages individual variation in colour is not
unusually great.

Skull. — In general the skull is br(jad, smooth and rounded,
with large, deep l)rain-case and small rostrum, tlie basicranial

Vin. ise.
Sciunix vulijaria. Nat. size.

axis bent downward so that foramen magnum is entirely below
level of alveolar line. Dorsal pi'ofile fiat or faintly and irregularly

RCItJRUS

901

convex to i-egion between postorbital pi'ocesses, 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 i-idge 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
brain-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 bull?e 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 m'\ their lateral diameter to inner margin of meatus
barely 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 oi'bital border sometimes complete so as to trans-
form thfi notch into a foramen ; postorbital processes long, slender,
sti'ongly bent downwai'd, the orbit surrounded by bone through
about three-quarters of its circumference. Zygomata not widely
spi'eading either in front or behind, the arches essentially parallel
with each other through the greater part of their extent ; jugal
projecting posteriorly liehind zygomatic process of squamosal, its

902 RODENTtA

Upper l)order rising at middle to form a sliglit but evident angle
marking hinder margin of orbit, its anterior extremity wedged
between lachrymal and zygomatic process of maxillary ; anteorbital
foramen small and slit-like, a little in front of level of anterior
premolar, 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 sijuarely 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
01' 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 posteriori}', where the hamulars curve
rather noticeably outward ; ectopterygoid 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 anterior border moderately convex ; articular
process with shallow liut noticeable concavity on outer surface ;
angular process well defined liy the deep curve of its lowei-
border.

2Vr///." UpptM- iiirisor robust, but course of shaft not con-
spicuously marked on sitle of rcjstruin, and root producing no
visible 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
covei'ing less than one-third circumference of tooth, its surface
linely 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
ova,t(^, 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>?«'*. Molariform teeth resembling eacli other in
general size aiid form, each with two roots on outer side and a
single lai'gei- I'oot on inner side, the inner border of crown formed

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-
verse 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

Fig. IS".
Sciitnis vulgaris. Clieek-teeth. x 5.

border of crown as an ill-defined ridge extending nearly to
base of fourth outer cusp. The principal deviations from this
pattern are : in jj»t*, first outer cusp nearly as large as second,
fourth transverse ridge with an evident tubercle near its inner
extremity ; m'\ all the outer cusps together with their transverse
ridges obsolete except the second, the posterior tM'o-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 jJiHt with antero-external border so
rounded off as to give the crown a somewhat triangular outline.
Main portion of crown occupied by a smooth depression, on both
outer and inner edge of which are three low tubercles, the
anterior and posterior outer having the broadest and best
defined bases, while the anterior internal is the highest ; on each

904

EODENTIA

side the median tubercle is much smaller than either of the two
between which it lies.

Memarl's. — 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 EUEOPEAN FORMS OP SCIURL'S VULGARIS.

White of underparts restricted to median region ;

hind foot about 50 mm. (Southern Spain) S. i: hieticux, 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
buff, never strongly reddish or blackish (British

Islands) S. r. levcoiirvs, p. 907.

Tail varying from red to black, never drab or buffy.
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. v. nuiiirint!u.t, p. 914.

Tail usually with conspicuous whitish median
area on under side (this sometimes repre-
sented by a few white hairs only).

Muzzle and cheeks rusty (Central Spain) S. r. iiifiixiriiii>:, p. 91G.

Muzzle and cheeks wbitish or pale grey

(South-western Spain) S. r. segunf,-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'. v. rulimrii:, p. 905.

Whiter pelage pearl-grey, the back scarcely
or not tinged with red (Extreme north
of Scandinavian Peninsula, east into

Russia) ,S'. n. mews, p. 900.

Dark brov/n phase not infrequent, sometimes
dominant (Central and southern).
Upper parts in dark brown phase much
darker anteriorly than posteriorly

(Greece) .". ,s'. v. Ulceus',^. 913.

Upper parts in dark phase not noticeably
darker anteriorly than posteriorly.
Rostral portion of skull unusually short

and broad (Pyrenees) s. v. fdpiiius,^). 914.

Rostral portion of skull normal.

Winter pelage of red phase essentially
without suffusion of grey on sides

(Italy) S. r. itftlh-nf:, p. 912.

Winter pelage of red phase strongly
suffused with grey on sides.
Rod in both winter and summer,
clear bright rufous (East-central

Europe) S. r. fuscnter, p 910.

Red in uoth winter and summer,
rufous strongly tinged with chest-
nut (West-central Europe) ,S'. r. nno-ns, p. 909.

905

SciURUS VULGARIS vuLOARis Linna>us.

1758. [Sciurvs'] vulgaris Linnteus, Syst. Nat., i, 10th ed., p. G.3.

1792. Scliurus] vulglaris] rufm Kerr, Anim. Kingd., p. 255 (Rencaming of

vulgaris).
1827. Sciurns vulgaris fi. albo votatiis Billberg, Synopsis Faunae Scan-

dinavife, p. 2 (Southern Sweden).
1827. Sciurus vulgaris y. albns Billberg, Synopsis Paunse Scandinavife,

p. 2 (Skiine, Sweden).
1827. Sciurus vtdgaris S. niger Billberg, Synopsis Faunse ScaudinaviaJ,

p. 2 (Skane, Sweden).
1857. Sciurus vulgaris Blasius, Siiugethiere Deutschlands, p. 272 (part).
1899. Sci2irns vulgaris typicus Barrett-Hamilton, Proe. Zool. Soc,

London, p. 6.
1910. Sciurus vulgaris Trouessart, Faune IMamm. d'Europe, p. 117.

Type locality. — Upsala, Sweden.

GengrajMcal distribution. — 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.

Measurements. — 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 : —
NoEWAY : Tolgen, Hedemarken, 1 ; Foldalen, Hedemarken, 1 ; Gausdal,
Kristiansamt, 1 ; Sundal, Bergenhus, 2 ; Konnernd, Buskerud, 1 ; Spjotsod,
Telemarken, 3 (U.S.N.M.) ; Aas, Akershus, 2 ; Aker, Akershus, 12 (B.M.
and U.S.N.M.); Asker, Chrjstiania, 2 (B.M. and U.S.N.M.) ; Maersness,
Uhristiania, 1 ; Holmestrand, Jarlesborg, 1 ; Kristiansand, INIandal, 2 ;
Frederickshold, Smaalcuene, 2 ; Gudbrandsdal, 1 ; no exact locality, 4.
Sweden : Upland, 2, near Upsala, 15 (B.M. and U.S.N.M.).

9. Tolgen, Hedemarken, ChristianiaMuseum (e). 97. 3. 13. 12.

Norway.

?. Foldalen, Hedemarken. ChristianiaMuseum (e). 97. 3. 13. 13.

i. Gausdal, Kristiansamt. ChristianiaMuseum (e). 97. 3. 13. 14.

i, ?, 2. Sundal, Bergenhus. C. B. Horsburgh (c & p). 99. 9. 5. 1-2.

6. Konnerud, Buskerud. Christiania Museum (e). 97.3.13.21.

2 6. Aas, Akershus. Christiania Museum (e). 97.3.13.17-18.

6. Aker, Akershus. Christiania INIuseum (e). 93. 8. 11. 8.

7 <5, 2 9, Aker, Akershus. ChristianiaMuseum (e). 97. 3. 13. 1-11.
2 juv.

9. Christiania Fjord. Rev. H. H. Slater (c & p). 99. 11. 27. 5.

6. Holmestrand, Jarles- Christiania Museum (e). 97.3.13.22.

berg.

2. Kristiansand, Mandal. Christiania Museum (e). 97. 3. 13. 15-lG.

2. Frederikshold, Sraaalc- Christiania Museum (e). 97. 3. 13. 19-20.

nene.

6. Gudbrandpdal. G. Barrer.t - Hamilton 11. 1. 2. 151.
(J. L. Lonhote.) (r).

90G RODKKTIA

2<5, 2 9. Near Christiania. W. E.deWinton (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. Kolthoff.) Lord Lilford (?). 11. 1. 1. 50-51.

SciURUS VULGARIS VARii's Gmeliii.

1789. [Sciurnii vulgaris] j8 varius Graelin, Syst. Nat., i, 13th ed., p. 14G.

1792. Sc[i?M-M.s] vtilg[aris] varius Kerr, Anim. Kingd., p. 256.

1829. [Sciurns vulgaris] 0 cinereus Fischer, Synopsis ]\Iammalium, p. 353.

Not Sciurus cinereus Linnfeus.
1857. Sciurus vulgaris Blasius, Saugethiere Deutschlands, p. 272 (part).
1899. Sciurus vulgaris varius Barrett-Hamilton, Proc. Zool. Soc,

London, p. G.
190G. Sciurus vulgaris varius Trouessart, Bull. IMus. d'Hist. Nat. Paris,

XII, p. 3G0.
1910. Sciurus vulgaris varius Trouessart, Fauue ^Mamm. d'Europe, p. 117.

Type Ivcality. — Northern Europe.

GefHirapMcal distrlhition. — From the extreme north of Scan-
dinavia eastward across Finland and into Kiissia.

DicKjrKms. — Similar to Sciurus vulgaris vulgaris but with grey
of winter pelage much lighter, closely approaching the pearl-
grey of Ridgway, and median dorsal region scarcely if at all
washed with chestnut.

ft;/o«y.— Winter pelage : haii's of upper pai'ts slate grey at
base (10 mm.), the underfur tipped (4 mm.) with a pale grey
most nearly approaching the grey No. 10 of Ridgway, but less
blue and with a slight smok}- cast, the longer hairs (30 mm.)
greyish to level of tips of imderfur, 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
i-eddish. Crown and face not so pale as back, occasionally with
a distinct brownish or reddish wash. Cheeks like sides of body ;
muzzle huffy whitish. Ears grizzled grey like sides of body
internally, the tufts A'arying from orange-rufous to blackish.
Outer surface of legs like sides of body, usually tinged to a vaiy-
ing degree with dull orange-rufous or light russet, this generally
the dominant colour on upper side of feet ; soles light huffy grev.
Tail greyish or light reddish ru.sset above, clouded by the blackisli
sub-terminal bands (20 mm.) of hairs ; tips of hairs incon-
spicuously light rus.set. 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.

Mensiiremenis. — Adult male from Kii-kkonummi, Finland :
head and body, 205; tail, 100: hind foot, -"•.">; ear, 30. For
cranial measurements see Table, }>. '.• 1 S.

sciuRUs 907

Specimens examined. — Eight, from the following localities :^
Norway: Hainmerfest, Finmarken, 2; Alten, Finmarken, 1.
Sweden: Enontekis, Norrbotten, 1.
Finland : Muouionniska, 3 ; Kirkkonummi, 1.

2. Hammerfest, Finmarken, G. Barrett-Hamilton 11. 1. 2. 152-153.

Norway. (J. L. Bonhutc.) (p).
9. Alten, Finmarken. Christiania Museum 93. 3. 1. 9.

(P)-
6. Enontekis, Norrbotten, I). Meinertzhagrn 97. 10. 21. 5.
Sweden. (c & p).

4. Muonionniska, Finland. D. Meinertzhagen 97. 10. 21. 1-4.

(c & p).

5. Kirkkonummi, Finland. Lord Lilford (p). 98. 11. 4. 2.

(P. Mevilainen.)

SciURUS VULGARIS LEUCOURUs Kerr.

1792. Sc\iurus] mdg\aris\ leucoiirus Kerr, Anim. Kingd., p. 25G.

1857. Sciurus vulgaris Blasius, Siiugethiere Deutschlands, p. 272 (part).

189G. Sciurus vulgaris leucurus Thomas, The Zoologist, 3d ser., xx,

p. 402, November, 189G.
1899. Sciurus leucurus Barrett-Hamilton, Proc. Zool. Soc, London, p. 3.
190G. Scimms vtilgaris leucurus Trouessart, Bull. IMus. d'Hist. Nat., Paris,

XII, p. 3G2.
1910. Sciurus vulgaris leucur2is Trouessart, Faune ]\Iamm. d'Europe,

p. 118.

Ti/pe local itij. — England.

Geographical distributibn. — Great Britain and Ireland.

Diagnosis. — Dark phase absent ; tail drab, fading in summer
to cream-buff, never tinged with red ; summer pela,ge : general
colour above light hazel or dull rufous, clearest and brightest on
sides and legs ; winter pelage : general colour above a dull
broccoli-brown. Size small, the condylobasal lengtli of skull
rarely if ever exceeding 48 mm.

Colour. — Summer ijelag(! : upper parts a I'eddish brown varying
from a light hazel to a dull rufous, darker and less reddish along
median area from face to rump, brighter on sides, on outer
surface of limbs, and on feet ; underparts dull white, usually
marked off from brown of sides by a narrow line of tawnj--
ochraceous, this spreading to suffuse inner surface 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. AVinter pelage : upperparts an incon-
spicuously grizzled greyish brown approaching the broccoli-
brown of Ridgway ; the middle of back faintlj' darker, tiie
shoulders and sides of neck usuall}" tinged with ru.sset ; outer
surface of limbs and narrow line along sides ru.sset or hazel,
rarely with the decided reddish tinge seen in summer ; under-
parts dull white ; tail and ear-tufts drab, occasionalh' with a
dusky cast.

Mensureiiicnt.t. — Adult male and female from Dunphail, Elgin,
Scotland: head and bodv, -18 and 213; tail, 179 and 165;

908 EODENTIA

liind foot, 58 and 56. Average and extremes of ten adults from
Saflron 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.

S2Jc'civiens examined. — One hundred and seventy-four, from the follow-
ing localities : —

Ireland : Craigio, Kilmanock, 1 ; Ahaseragh, Galway, 1 ; Woodpark,
Clare, 1 ; Graigne, Kilkenny, 1.

Scotland: Black Isle, Cromarty,!; Dunphail, Elgin, 3; Ballends-
loch, Banffshire, 2; Loch Earn Head, Perthshire, 3; Pitlochry, Perth-
shire, 2: Stockbriggs, Lanarkshire, 3; Tillicoultry, Clackmannanshire, 3.

England : Guisborough, 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, Crloucester-
shire, 1 ; Newland, Coleford, Gloucestershire, 1 ; Clarberton Road, Pem-
brokeshire, 1 ; Glamorganshire, 1 ; ]\Ionmouthshire, 1 ; Clevedon,
Somersetshire, 1 ; Wormsley, Oxfordshire, 5 ; Thornhaugh, Northampton-
shire, 4; Luton, Bedfordshire, 6; Pelden, Hertfordshire, 3 ; Tring, Hert-
fordshire, 1; Saffron Walden, Essex, 40; Whitley, Surrey, 1 (U.S.N.:\I.) ;
Milford, Surrey, 2 (U.S.N.il.) ; Horsham, Surrey, 1 ; Knockholt, Kent, 5 ;
Upton, near Andover, Hampshire, 1 ; Blandford, Dorsetshire, .55.

BeiiiarJis. — B}^ its small size and the peculiar colour of its
tail the British squirrel is one of the most sharply differentiated
races of Sciiirus ml</ari>i. 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 to produce bleacliing. 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 convenience it is here treated as a race.

9. Ahaseragh, Galway, Ire- G. Barrett-Hamilton 11. 1. 2. 154.

land. {W.S.Witherbi/.) (p).

6. Woodpark, Clare. G. Barrett-Hamilton 11. 1. 2. 155.

(/?. F. Hibbert.) (p).

?. Graigne, Kilkenny. G. Barrett-Hamilton 11. 1. 2. 150.

(L. I. Finn.) (p).

V. Black Isle, Cromarty, W. 11. Ogilvie-Grant 11. 1. 3. 422.

Scotland. " (c & p).

V. Dunphail, Elgin. W. R. Ogilvie-Grant 11. 1. 3. 423.
(c & P).

3 S. Stockbriggs, Lanarkshire. E. R. Alston (c <t p). 79. 9. 25. 2G-28.

<?, 2 V. Tillicoultry, Clackman- R. G. Wardlaw liam- 11. 1. 3. 424-42G.

uanshirc. say (c & p).

1 . Glamorganshire, Wales. Rev. A. Morgan (c & v). 71. 7. 9. 1.

1. Monmouthsliiro. Rev. A. Morgan (c & p). 73.7.12.1.

6. Oswaldskirk, Yorkshire, A. Houston Boswell 11. 1. 3. 427.

Engiaud. (c & p).

9. Guisborough, Yorkshire. A. E. Pease (c & p). 11. 1. 3. 428.

sciuRus 909

J, 29. Thornhaugh, Northanip- Rev. H. H. Slater 11. 1. 3. -129-431.

toushire. (c & )>).

•2.6. Bury St. Edmunds, Suf- Dr. A.Giinther (cct p). 83.2.3.1-2.
folk.

6. Horsham, Surrey. E. R. Alston (i). 7'J. 9. 25. 29.

(J. E. Ifarting.)

55. Blandford, Dorset. J. C. Manscl Pleydell 97. 1. 6. 1-55.

(c & p).

SciURUS VULGARIS RUSSUS Miller.

1857. Sciurus vulr/aris Blasius, Siiugothiere Deutschlauds, p. 272 (part).
1899. Sciurus vulgaris rufus Barrett-Hamilton, Proc. Zool. See, London,
p. 5 (part). Not of Kerr, 1792.

1906. Sciurus vulgaris rufus Trouessart, Bull. Mus. d'Hist. Nat. Paris,

XII, p. 360 (part). Not of Korr,'l792.

1907. Sciurus vulgaris russus Miller, Ann. and Mag. Nat. Hist., 7th sor.,

XX, p. 427, November, 1907. Type in British IMuseum.
1910. Sciurus vulgaris russus Trouessart, Fauna Mamm. d'Europc, p. 119.

Type locality. — Dinan, Cotes-du-Nord, France.

Geographical flistrihution. — West-central continental Europe
from Brittany to Holland, east to central France and jaerhaps
further ; details of distribution not known.

Diagnosis. — Dark phase present but less common than red
phase. Hummer pelage : red phase : essentially as in S. vulgaris
vulgaris, 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 blackish.
AVinter pelage : light phase : body hazel, the sides a little
sullused with light smoke-grey, tail rufous, much darker than
that of Ridgway ; dark j^hase not seen.

Measurements. — 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.

Spcciinens examined. — ^Twenty-six, from the following localities : —
Holland : s'Graveland, 8 ; Oosterbeek, Guelderland, 2.
France : Seinc-Inf6rieure, 5 ; Dinan, C6tes-du-Nord, 4 ; Meurthe-et-
Moselle, 7 (perhaps referable to S. v. infuscatus).

2 c5, 4 9. s'Graveland, Hilversen, F. E. Blaauw (c & p). 98. 5. 17. 1-2.

Holland. 98. 6. 11. I-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. Prance. (p).

4 6. Dinan, Brittany. W. J. Bramley (c & p). 97. 11. 6. 1-4

(97. 11. 6. 2. TyjK of subspecies.)

5 9. Manonville, Meurthe- Lord Lilford (p). 11. 1. 1. 97-101.

et-Moselle. (Lomont.)

910

SciUliUS VULGARIS FUSCOATEli Altum.

1804. Scinriis vulgaris var. cincrca Hermann, Observ. ZooL, p. 65

(Germany '?) Not Sciurus cinereus Liunieus 1766.
1857. Sciurns vulgaris Blasius Siiugethiere Deutschlands, p. 272 (part).
1S7G. [Sciurus vulgaris] var. fuscoatra Altum, Forstzoologie, 2d ed., i,

p. 75 (Harz Mts. and Silesia).
187G. [Sciurus vulgaris'] var. nigresccns Altum, Forstzoologie, 2d ed., i,

p. 75 (Silesia).
1876. [Sciurus vulgaris] var. hrunnca Altum, Forstzoologie, 2d ed., i,

p. 75 (Alsace-Lorraine).
1876. [Sciurus vulgaris] graca Altum, Forstzoologie, 2d ed., i, p. 75

(synonym of hrunnca).
1876. [Sciurus vulgaris] alpma 'Altum, Forstzoologie, 2d ed., i, p. 75

(synonym of brunnea). Not Sciurus alpinus F. Cuvier, 1812.
IS'J'J. Sciurus vulgaris rufus Barrett-Hamilton, Proc. ZooL Soc. London,

p. 5. Not of Kerr, 1792.
1905. Sciurus vulgaris var. gotthardi Fatio, Arch. Sci. I'hys. et Nat.,

Geneve, 4th scr., xix, p. 512 (South slope of 'Mt. St. Gothard,

Switzerland).

1907. Sciuriis vulgaris rutilans IMiller, Ann. and ilag. Nat. Hist., 7th ser.,

XX, p. 426, November, 1907 (Rudolstadt, Thiiringen, Germany).
Type in British ]\Iuseum.

1908. Sciurus vulgaris fuscuatcr Miller, Ann. and ^lag. Nat. Hist., 8th ser.,

1, i>. 128, January, 1908.
1910. Sciurus vulgaris fnscoutcr Troucssart, Faune 3Iamm. d'Europe,
p. 118.

Tj/iic Idciilitij. — llarz Moiintain.s, (Ttu'iiiany.

GcLKjrdjiliicdl (lixtrihation. — East-central Europe fruui (lermany
through Austria-Hungary to Roumania; south through tlie Alps.

Diaiino><is. — Hioiilar to S. vnhjaria riismafi, but red phase in
l)oth winter and summer a clear bright rufous, tinged with grev
on sides and sometimes on back. Dark phase present but less
connnon than red phase, the relative abundance of the two
subject to much local variation. Summer pelage : light phase :
body clear rufous, closely agreeing with that of Eidgway, and
without chestnut sufiusion, tail the sanae clear rufous but more
intense ; dark phase as in winter but general effect more sooty.
AVinter pelage : light phase : body rufous tinged with light
smoke- grey along sides, tail clear rufous; dark phase: liody a
grizzled hair-brown suH'used with mummy-brown over back : tail
slaty black.

Measurements. — Average and extremes of live adults from
Ingelheim, liheinhessen, Germany: head and bo<lv, 229 •■! (220-
236); tail, 192 (189-198); hind foot, GO-4 (59-61); ear, 32-i:
(31-04). Young adult male from Rudolstadt, Thiiringen,
(nirmany (type of rntllaiis Millei') : head and body, 223 ; tail,
1 75 ; hind foot, 57 ; ear, 27. Adult female from Niesky, iSilesia,
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, 18-1 and 190; hind foot, 61 and

SCIUEUS nil

/■'■,, ^'^-^

theidnlts from P^J. ?f ' '^^^ • ^^'"^'"''^ ^^^^ *^-^tremes of

^'l3 -^o^r f -1 ^^f «"^"' l^outfJ^ni^ •• head and body, 220 -S
eir 30 oioo -'-^^;.^ (170-185); hind foot' 58-6 (57-60)-
tar, 30 J (29-02-0). iV cranial measurements see Table p 920^

loc^Mes:-^ ™-^.-One hundred and seventy, from the following

(Motta.); Barcelonnette, EaicSpet^s m\T Tnd' MoT/f ".'"n"' '
Basses-Alpes, 3 (Mottaz) ^ "'^ JMottaz) ; Digue,

BuSirixony:'riu??T"M)-''St.b; ^^---^^^l- Hanover, 2;
Wern>gerodo, Sa/ony. 1 ; -^fusSl, £^^2 ^ IlSbu.^ ,?. (J^^^f f •> =
1 ; Aiumeuslablen, 1; Eudolstadt Thiirincr/n ^ -^^^enuuig, j Magdeburg,
12; Ingclheim, 10 HartwSrBaden 1 Fs V in""^"'''''l'' T^^^'^S^".
1 (U.S.N.M.); Strass, near' Bu^L i^ bS ' ^-'^'m^S'^^^
.^lonheim, Bavaria, G ; Niesky, SileSa 1 ^™"'^' "^ • ^^arxheim, near

AusTKiA-HuNGARY : Csalfokoz-Somoria Pressbnro- 9 . rv c ■■, c. ■
mark, 1; Haida, Arva Bobpniis fi t't i u i t,^' ^' ^refail, Steier-

Hunyad,' Bohemia 1"' Hu^ar; no exS' .' rf^^^' '' '^^''''''^'
Zuberec Hungary, 2. -"""°^'^>' 'lo exact locality, 1 (U.S.N.M.) ;

RouMANiA : Bustenari, Prahova 5
MoSrF^^^^ri'S:i^z?-^^t2ie''.'f;?r^ 1 (U.S.N.M.);

Gnsons, 3 B.M. and U S N M) ■ Rahinrrs.? i /f^. -C^^') ' Untervatz,

Ticino, 1 (U.S.N.MO; Cul'eiia ¥idno r°^^^^

Lugano, Ticino, 2 : Lugano THnor; m VV'^/'^ ' ^^a^^'esco, near
(U.S.N.MO; no exact locality'' l' (^^°"*^) ^ ^umino, Ticino, 1

Co^M^^^^^: ^j^St; S'i^^^r^' ^°^^^^' ^ (Turin);
Sommariva del Bosio, Cuieo'2 (^ndnSt' tS^^ c^ Y'""' ' ^^"""^ '
'■ ''s^o^F^nce''^'^'" A. Robert (c&P). ' 0,. 1. ,. ,.

(5,2?.

Montauban, Haute- O. Thomas (r). 0 4 o ,,

bavoie. (.4-. Robert.) ^ ' • ^- -^- ■^•

Marburg, Hessen-Nassau, Lord Lilford (r) n 1 1 r-. .„

Germany. (Schneider) ^'' ^l- L L (6^/S.

(5. Blumcnthal Hanover. Lord Lilford (r) 11 1 1 q^-

(WoUeniiurff.) ^ '' ^- ^- '- ^'•

?. Wernigerode, Saxonj-. Lord Lilford (r). 11 1 1 ss

(Molterstorff.) ^ ' ^- ^- '- °^-

^. V. Neustadt, Saxony. Lord Lilford (r). 11 l 1 ss ^^j

(Schneider.) ^ ' ' ^- -^- o3-Sl.

<i, ? ?. Ilseuburg, Saxony. Lord Lilford (p). n 1 . ..^ rn

(Bartclh.) ^ ' -Li- L 1. jj^eo.

1. Gross Ammenslablen, Lord Lilford (r). 11 l 1 ni

Saxony. (Schuchardt.) ^ ' ' ^- ^- ^^■

/■ Magdeburg Saxony. Lord Lilford (p). 7971

5 .5, ?. Ummerstadt, Thiiriugen. Lord Lilford p u ]'\ V^ .,

- . {Schuchardt.) ^ '' ^^- ^- 1- '0-'5.

2d, 3 9. Rudolstadt Schwartz- Lord Lilford (p). 95 4 m ^ 11

burg, Thurmgen. ^95 4 iq 7 t, J f o ^^- '"^l-
a, 5 9. Ingelheim, RhJnhessen. C ffil 'ert (cf ' '^■'•«ff filler.)

3 6, 9. Ingelheim, Rheinhessen. Lord Si^d^p) J^ ^^^ f 32-37.

(Hilgert.) ^ '' ' -^- '^-0^.

■ • ^^^^^^^Ut^:) Lord Lilford (P). 11. 1. 1. 85-86.

912 ■ RODENTIA

9. Strass, Bavaria. Lord Lilford (p). 11. 1. 1. 89.

6, 2 9. Marxheim, Bavaria. Lord Lilford (i>). 11. 1. 1. 67-69.

{Woltcrstofff.)

9. Niesliy, Silesia. Dr. E. Hamilton (r). 97. 12. 4. 20.

2. Csallok'oz-Somor j a, Budapest Museum (e). 91.3.1.29.

Pressburg, Hungary.
6. Trefail,Steiermark,Hun- Lord Lilford (p). 11.1.1.90.

gary.
6 6. Haida, Bohemia. Lord Lilford (p). 11. 1. 1. 61-66.

(Wolterstorff.)

1. Karlsbad, Bohemia. Lord Odo Russell 44. 9. 7. 3.

(c & p).
6. Hatszog, Hunycad, Trail- C. G. Danford (c). 3. 11. S. 20.
sylvania.

2. Zubcrer. BudapestMuscum(B). 91. 3. 1. 31-32.
3 iJ, 2 9. Bustenari.Prahova, Rou- Lord Lilford (p). 4. 4. 6. 67-71.

mania. [W. Dodson.)
2 o', 2 9. St. Galleu, Switzerland. O. Thomas (p). 4. 4. 5. 35-38.

(Zollikofcr.)
2 9. Untorvatz, Grisons. O. Thomas (p). 4. 4. 5. 39-40.

{Zollikofcr.)
i, 6 juv. Davcsco, Ticino. O. Thomas (p). 2. 8. 4. 28-29.

(Zollikofcr.)
6. Switzerland. E. R. .\lston (p). 79. 9. 25. 30.

SCIURUS VULtiAPaS ITALICUS Buiuipaite.

1838. Sciurus italicus Bonaparte, Iconogr. Faun. Ital., i, fasc. 23 (Italy).
1857. Sciurus vulgaris Blasius, Siiugethiero Dcutschlands, p. 272 (part).

1906. Sciurus vulgaris italicus Trouessart, Bull. ilus. d'Hist. Nat. Paris,

XII, p. 364 (part).

1907. Sciurus 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 italicus Trouessart, Faunc ^lamm. d'Europc,
p. 122.

2\i/p,' /(/ra/////.— Italy.

GeiKjraphical tJistrihation. — Italy except tlic region iioith of
the Apennines.

Di(upwnis. — Ked phase H<ihter and more ochraceous than in
Sri II run vuhjaris fuscoater ; brown phase less dark than in
fnscodter, the tail with broad grizzled median area ; back rarely
if ever showing any evident trace of a darker median line, and
sides rarely with any noticeable greyish 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-bufT; underparts from throat to
base of tail buflfy white ; interramia tinged vath grey ; inner
side of legs paler and less red than outer sirle, closely resenabhng

SClURtJS 013

mediiin 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 jjredominates, and in which there is
frecjuently 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,
Home, 2.

6. Quiesa, Lucca, Italy. Dr. E. Hamilton (p). 98. 10. 2. 9.

i. San Casciano, Florence. Dr. E. Hamilton (p). 98. 10. 2. 10.

2. Siena. [Brogi.) Paris Museum (e). G. 7. 15. 1-2.

?. Arsoli, Rome. {Coli.) G. Barrett-Hamilton 11. 1. 2. 36.

6. Viterbo, Rome. {Coli.) G. Barrett-Hamilton 11. 1. 2. 35.
(P)-

SciURUS VULGARIS LiL^EUS Miller.

1906. Scittrus vulgaris italicns Trouessart, Bull. Mus. d'Hist. Nat. Paris,

XII, p. 364 (part).

1907. Sciurus vulgaris lilnms Miller, Ann. and Mag. Nat. Hist., 7th ser.,

XX, p. 429, November, 1907. Type in British Museum.
1910. Sciurus vulgaris Hiatus Trouessart, Fauna Mamm. d'Europe, p. 123.

Ti/pe locality. — Agoriani, north side of Lyakura (Parnassus)
Mountains, Greece.

Geographical distrihution. — 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 back with the hair-brown
shoulders and neck ; red phase not known and probably rare or
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 outei- surface of hind
legs. Inner surface of hind legs and line along sides of body
bordering white of underparts tawny-ochraceous, becoming

3 N

914 RODENTIA

lighter and duller anterioi'ly 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.

Measurements. — 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 localit}'.

1. Agoriani, Lyakura Mts., Greece. Lord Lilford (p). 7. 9. 8. 1.

[SclilUter.) {Type of subspecies.)

1. Parnassus. Lord Lilford (p). 8. 10. 17. 1.

SciURUs VULGARIS ALPiNUs Desmarest.

1822. Sciurus alpinns Desmarest, Mammalogie, ii, p. 543 (Pyrenees).
1842. Sciurus alpimis P. Cuvier, Hist. Nat. des Mamm., iv, tabl. gen., p. 4

(described and figured in fasc. 24, pi. 237, 1821).
1857. Sciurus vulgaris Blasius, Saugethiere Deutschlands, p. 272 (part).
1905. Sciurus alpinns Cabrera, Bol. Real Soc. Espafi. Hist. Nat., v, p. 230,

April, 1905.
190G. Sciurus vulgaris alpinus Trouossart, Bull. Mus. d'Hist. Nat., Paris,

XII, p. 3G3.
1910. Sciurus vulgaris alpinus Trouossart. Faune ]Mamm. d'Europe, p. 119.

Type localitij. — Pyrenees.

Geographlcnl distrihuiion. — Pyrenees ; limits of range not
known.

Diagnosis. — Size essentially as in Seiurus vulgaris itaUrus ;
colour in dark phase similar to that of S. v. russus and »S'. v.
fuscoafer ; skull with I'ostral poi-tion noticeablj' broadened and
shortened.

Measurements. — Adult male from Espot, Lurida, 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.JI. and
Cabrera).

BemnrJi-s. — The status of this race is not clearly understood.

Sciurus vulgaris numantius Miller.

1905. Slciunisl sp. Cabrera, Bol. Real Soc. Espafi. Hist. Nat., ^Madrid, iv,

"pp. 224,231, April, 1905.
1905. S\ciurus] rufus Cabrera, Bol. Real Soc. Espan. Hist. Nat., iladrid,

IV, p. 225, April, 1905 (not of Kerr, 1792).
1907. Sciurus vulgaris numantius Miller, Ann. and ]\Iag. Nat. Hist., 7th

ser., sx, p. 428, November, 1907. Tj'pe in British Museum.
1910. Sciurus rulgaris mimantius Trouessart, Faune Mamm. d'Europe,

p. 120.

Ti/pe localitij. — Pinares de (^Hiintanar do la Sierra, Burgos,
Spain.

SCIURUS 915

Gi'0(/rnp]iica] disirihiition. — Northern and north-central Spain
(provinces of Leon, Vitijria, Huesca and Burgos) ; limits of I'auge
not known.

Diagnosis. — Size greater than in the central European forms,
nearly equal to that of S. v. infiiscatus ; 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 brown, 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-butFand clay-colour. Cheeks drab. Ear-tufts
blackish brown, sides of neck pale dull wood-brown. Inner
surface of legs and ill-defined stripe along sides of belly dull light
hazel. Eeet 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
annulations. 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 suftusion. 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.

Shdl 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, 23-5 and 250 ; tail, 190
and 210; hind foot, 63 and 64; ear, 34 and 34. Adult malo
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 :^
Prance : Solf^riuo, Landes, 1 ; St. Jean de Luz, Basses-Pyrenees, 1.

(These specimens intermediate between nnmantius and ritssits.)

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.) ;

i'inares de Quintanar, Burgos, 7.

?. S o 1 f e r i n o, Landes, O. Thomas (p). 6. 4. 1. .38.

France. (A. Robert.)

S. 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).

* ]\Iisprinted 230 in original description.

3 N 2

916 RODENTIA

?. Pajares, Leon. O. Thomas (p). 8. 2. 9. Gl.

(iV. Gonzalez.)

2 6. Arrechavaleta, Vitoria. O. Thomas (p). 8. 2. 9. 65-66.
(N. Gonzalez.)

2 i, ?. Paiiticosa, Huesca. 0. 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 subspecies.)

SciURUS vuLOARis ixFuscATUS Cabrera.

1905. Sciiirus mfuscaius Cabrera, Bel. Real Soc. Espau. Hist. Nat., v,
p. 227, April, 1907. Type in U.S. National Museum.

1909. Sciurits vulgaris infiiscatus Miller, .\nn. and IMag. Nat. Hist., Stb

ser., Ill, p. 418, May, 1908.

1910. Scktrus vulgaris infuscatus Trouessart, Faune ^Maram. d'Europe,

p. 121.

Tijpe locality. — Las Navas, Avila, Spain.

Geographical distribution. — Central Spain.

DiacjnoHin. — Largest of the European squirrels (condylobasal
length of skull, 52 • 4 to 55 mm.) ; colour very dark, the two
jahases if present not well defined ; tail rusty rod, 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, tlie
general effect nearly the mummy-brown of Ridgway, the posterioi-
half of back darker and less grizzled than shoulders and neck ;
underfur slate-grey, the tips of tlie hairs russet ; face rather
abruptly russet, the muzzle tinged with buff; cheeks below and
behind eyes an indefinite drab brown, noticeably contrasted with
surrounding parts, especially with white of throat ; ears russet,
the longer hairs dusky ; feet rufous, somewhat darker than that
of Ridgway, this colour suffusing outer surface of fore leg and
extending uji inner surface of hind leg to thigh, though in both
in.stances duller and more brownish on leg than on foot ; imder-
parts buffy white, the extreme bases of hairs inconspicuously
greyish, 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 whitisli, often very
noticeable median area below.

Sladl and teeth. — Except for their larger size tlie skull and
teeth do not differ appreciably from those of the central European
forms.

Measurp.ments. — Type (adult male) : head and body, 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.

f

sciuRUs 917

Specimens examined. — Five, from tho following localities in centra!
Spain: Las Navas, Avila, 1 (U.S.N. INI.) ; La Granja, Segovia, 3 (B.M. and
Cabrera) ; no exact locality, 1.

6. Old Castile, Spain. Lord Lilford (r). 94. 6. 11. R.

i. La Granja, Segovia. A. Cabrera (r). 7. 10. 17. 1.

{Escalcra.)

SnuKUS vuLUAUis siiGuiLE Miller.

1909. Sciurus vulgaris segurn' Miller, Ann. and Mag. Nat. Hist., 8th ser.,

Ill, p. 418, May, 1909. Type in British ]\Iuseuni.

1910. Sciurus vulgaris segur;e Trouessart, Faune ]\Iamm. d'Europe, p. 121.

Ti/jje locoJity. — Molinicos, Sierfa de Segura, Albacete, iSpaiii.

Geoijraplikal distribntion. — Known only from the Sierra de
Segura, provinces of Albacete and Jaen, south-eastern Spain.

Diagnosi><. — Related to Sciurus vulgaris infuscatus, but back
less blackish, its uuderfur 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.

Colour. — 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
.sj^ecimens the light element is more nearly russet and the black
is essentially absent ; underfur pale ecru-drab, the hairs with
faintl}^ 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
(jf throat ; underfur of head, back, sides, and limbs pale eci-u-
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 buliy 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.

Slull and teeth. — The skull and teeth do not difl'er appreciably
from those of Sciurus vulgaris infuscatus.

Measurements. — Type (adult female) : head and body, 245 ;
tail (probably injured in life), 195 ; hind foot, 61. Average 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 do Segura.

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11. 1. 1.72

95.4. 18.9

95. 4. 18. 10
95. 4. 18. 11

95. 4. 18. 7*

95. 4. IS. 8
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4. 4. G. G7
4. 4. (!. G8
4. 4. G. 70
5G38 Field

115217
4.4.5.36

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S. vulgaris fuscoater.

Germany: Ingelheim, Rheiu-l
hessen . . 1

Ummcrstadt, Thii-i
ringen . . (

Schwarzburg, Thii- [
ringen . . (

Rudolstadt, Thii-'l
ringen . ./

Niesky, Silesia
Roumania : Bustenari .

Switzerland : Brunig, Born .
St. Gallen

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sciuEus 923

Itemarlcs. — While tliis animal is readily diHliuguishable from
the squirrels of central and northern Spain its distinctness from
the imperfectly known Scixnis viihiaris hseticus is perhaps open to
doTibt. In an obviously immature individual, however, the hind
foot is 60 mm. in length as opposed to 45 mm. in the type of
hxticnti.

2 (5, 4 ? Sierra de Segura, Albacete, M. de la Escalera (c) 8. 9. 24. 1-6.
Spain. (8. 9. 23. 3 Type of subspecies.)

SciUKUS VULGARIS B.ETicus Cabrera.

1905. Sciurus hivticus Cabrera, Bol. Real Soc. Espaii. Hist. Nat., v, p. 228,
April, 1905. Type iii museum of University of Seville.

1910. Sciurus vulgaris bxticus Cabrera, Asoc. Espau. Progr. Cien., Congr.
Zaragosa, 1908, p. 11, June, 1910.

1910. Sciurus vulgaris b;rticus Trouessart, Faune Blamm. d'Europe, p. 122.

Ti/pe localHij. — Alanis, Province of Seville, Spain.

Geograpliical distribation. — At j^resent known from the type
locality only.

Diagnosis. — In general much like Sciurus vulgaris scgune, but
size considerably less (hind foot only about 50 mm.) and white
area on underparts unusually narrow.

BemarJcs. — This scjuirrel, of which I have seen no specimens,
appears to be similar to Sciurus vulgaris segurse 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 Sciums ; 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 infuscatus, 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 surface. 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 haeticus.

* (from infuscatus).

924

KODEXTIA

Genus CITELLUS Oken.

181G. CitcUus Oken, Lchrb. der Naturgcscb., Th. iii, Abth., ii, p. 824

{citcUus).
1825. Spcrmophihis P. Cuvier, Dents cles jMammiforcs, p. IGO {citellits).
1827-34. Gitillus Lichtenstein, Darstell. ncucr oder wcnig bekannter

Siiugotb., pi. XXXI (citcUus).
1857. Spcrmophilus Blasius, Siiugetbicrc Dcutscblands, p. 275.
l'J02. Citdlus Allen, Bull. Amer. Mus. Nat. Hist., xvi, p. 375, October 11,

1902.

Tjipe xjx'cics. — Mhs ritelliig Linn;i'us.

Gcor/rapliical dhirihitiion. — Northern H'eniispliero fri)ni
Hungary eastward through Asia and North America to the
central United States.

Characters. — Strictly terrestrial Srliiridie of medium size and
slightly modified Sciurine aspect, the tail flattened but relatively
shorter and less bushy than in Sciitriis ; cheek pouches present ;
skull more massive than in ScinrKS, 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 (Jshaped pattern in moderately worn
teeth ; anterior upper premolar well developed.

Hemarls. — Though the limits of the genus CitrJhi)- are not at
present well understood, owing chietly to the difficulty of com-
paring the Old AVorld and American forms, the group may be
regarded as containing about eight Pala-aix'tic species, two of
which occur in central Europe, llemains of extinct members of
tlie genus are found as far west as England.

KEY TO THE EUROPEAN SPECIES OP CITELLUS.

Back Nvitbout distinct spots ; skull with incisive fora-
mina normal, their length greater than diameter of
alveolus of upper incisor (Hungary in general, and
eastward) C. citcUus,]). 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 east of the Carpathians, and
eastward) C. suslica, p. 929.

CITELLUS CITELLUS Liniueus.

1766. [iLTi^s] citcUus Linnfeus, Syst. Nat. i, 12th ed., p. 80 (Austria).
1778. M21S citillus Pallas, Nov. Sp. Quadr. Glir. Ord., p. 19.
1857. Spcrmophilus citcUus Blasius, Saugethiere Deutschlands, p. 275.
1904. [CitcUus'] citcUus Trouessart, Catal. Mam. Yiv. Foss., Suppl., p. 339.
1910. CitcUus [CitcUus) citcUus Trouessart, Fauna Mamm. d'Europe,
p. 127.

Type localiiy. — Austria.

Geographical distribution. — From Silesia and Bohemia, east-

CITELLUS . 925

ward thr<tui;h llus.sia into Asia; eastern limits of range not
known.

Dingiiosia. — Size medium (head and body, 195 to 1^20 mm. ;
tail, 60 to 75 j 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 buff and dark brown so arranged
as to produce an evident effect of light mottling on a dark
ground ; skull with ratlier low, broad brain-case ; incisive
foramina not unusually reduced in size ; incisor teeth both above
and below exhiljiting the minimum degree of compression.

External cliaracters. — General form less slender than in
Sciurus vulgaris, the short, narrow tail extending little beyond
outstretched hind feet, the ear short and without tuft, the fur
thin, coarse and lying close to body. Ear low, obscurely pointed
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 thumb
rudimentary but with rather evident, compressed nail ; hind
foot with third digit longest, second and fourth sub-equal 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 1 5 mm. long. Mammae ; p 1 — 1 ; a 2 — 2 ; « 2 — 2 = 10.

Colour. — Back and sides usually cream-buff but occasionally
more yellowish, in some specimens apjjroaching the buff yellow
of Ridgway ; sides inconspicuously " lined " with black ; entire
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 efiect of obscure light mottling, the spots
about 5 mm. in diameter ; crown and upjjer half of cheeks
grizzled, the light element paler than on body, the dark relatively
more evident ; an ill-defined eye-ring, whitish or yellowi.sh accord-
ing to general colouring of body ; muzzle sometimes with a
rusty tinge ; sides of muzzle and lower half of cheeks clear
whitish or huffy, continuous with the similarly coloured pale
area covering throat and fore part of chest and inner side of fore
leg ; belly washed with a buft* usually somewhat more yellow
than that of sides ; feet like ground colour of back ; tail without
definite colour pattern, the ujjper surface grizzled, essentially
like back, somewhat darker at tip, the pencil with light margin ;
under surface of tail a nearly clear dull buff'.

Shidl. — Tlie skull is smallei- than that oi Sciurus vulgaris,

92G . EODENTIA

and its general form clitfers notably in the cuneate 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 tf> lambdoid ridge, sometimes with a slight flattening
in interorljital 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. 1S8.
Citflhit: cltelius. Xat. size.

surticiently to be just visible from above. Brain-case very
broadly ovate when viewed from above, its greatest width
equal to length, its posterior width obviously exceeding than
that at postorbital constriction ; surface less rounded than in
Scinrus vulgaris, and lyrate figure formed by parietal ridges
much narrower ; lamljdoid crest essentially as in S. viihjari,^, but
constriction just in front of occipital region less evident. Occiput
low, its general form as in Sciurus vnhjaris, but contrast between
its height and that of middle of brain-case not especially mai-ked ;
protuberances on posterior surface of occiput less noticeable than

CITELLUS 92/

in S. ritJgiiris, aud exposed portion of petrosal noticeably smaller ;
paroccipital processes rather large, their extiemities reaching
beyond level of lower edge of condyle and often to that of
lower edge of bulla ; basioccipital essentially like that of Sciurus
ndgaris in general form, but anterior portion of outer margin
raised into a conspicuous plate-like process applied to inner side
of bulla ; auditory bulla essentially as in Sciurus vulgaris but more
evenly though somewhat less inflated. Tnteroibital 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 supraorbital notch inconspicuous ; postorbital processes
I'elatively shorter than in Sciurvs vulgaris, the space lying
betfween 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
230sterior limit of orbit ; anterior zygomatic root with conspicuous
projecting ridge extending along its entire anterior border ;
anteorbital foramen sub-circular in outline, 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 ; mesopterygoid
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. Mandible 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 veiy 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 fiom antero-internal extremity nearly

928

half-way along anterior and outer curvature, its surface smooth,
yellowish white. Lower incisor with root extending- slightly
beyond m.j ; 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 vidgaris 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. ridcjaris, 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-
ing a slight cusp, its abruptly
rounded inner termination lying
on anterior surface of base (jf
main tubercle ; thii'd ridge re-
l^resented by a minute cusp
between outer extremities of IJ-
shaped figure, the space between
the limbs of the \J occupied
by a smooth dejaression ; 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
J///*-', second limb of \J, with its outer cusp somewhat reduced ;
in «(■'•, .second limb of U obsolete and distorted, not extending to
outer bordei- 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
i-idge 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.

Measurements. — Averatje and extremes of four adults from

Fig. 180.
Clfdlux clti'lliis. rheek-teeth.

CITELLUS 929

Hirschberg, Bohemia: head and body, 200 (195-215); tail, 6(5
(60-73); hind foot, 36 '6 (35 -4-38 -4). Average and extremes
of six adults from Znaim, Mahren, Austria- Hungary : head and
body, 209-1 (200-220); tail, 63-7 (58-70); hind foot, 36-4
(35 • 6-37). For cranial measurements see Table, p. 930.

Specimens examined. — Thirty-nine, from tlie following localities : —

AusTRiA-HuNGABY : Haida, Arva, Bohemia, 1 ; Hirschberg, Bohemia, 5 ;
Znaim, Mahren, 7 ; Csallokoz-Somorja, western Hungary, 4 ; central
Hungary, 1 ; Sarvar, Eisenburg, Hungary, 1 (U.S.N.M.) ; Monos Petrie,
Bihar, 2 ; Csehtelek, Bihar, G.

RouMANiA : Kustendje, 2 ; Dobrudscha, 3 (B.M., U.S.N.M. and Genoa) ;
Moldavia, 3.

Bulgaria : Bogoroff, Sofia, 1 ; Sofia, 2 (Andersen).

Turkey : Ob-Meidan, 1.

2 (J, 2?. Hirschberg, Bohemia, Lord Lilford (p). 8.9.10.1-4.

Austria-Hungary.
6, 3 ?. Znaim, Mahren. Lord Lilford (p). 8. 9. 10. 1-4.

4. Csallokoz-Somorja, Press- Budapest Museum (e). 94. 3. 1. 33-3C.

burg, 400 ft.
2 9. Monos Petrie, Bihar. Hon. N. C. Roths- 9. 7. 23. 1-2.

child (c & p).
2 (i, 4 9. Csehtelek, Bihar. Hon. Mrs. Chas. 10. 9. 14. 2-7.

Rothschild (c & p).
2. Kustendje, Roumania. E. Cavendish Taylor 5. 5. 6. lG-17.

(c & p).
1. Dobrudscha. Purchased (Pruliere). 86. 4. 2. 3.

1. Moldavia. Dr. E. Rtippell. 42. 9. 4. 1.

1. Moldavia. Purchased (Lead- 43. 9. 11. 2.

beater).
1. Moldavia. Dr. J. E. Gray (p). 123. a.

9. Bogoroff, Sofia, Bulgaria. Dr. A. Richards 2. 10. 5. 1.

(c & p).

CITELLUS SUSLICA Gueldenstaedt.

1770. Mus suslica Gueldenstaedt, Nov. Comm. Acad. Sci. Imp. Petrop.,

XIV, pt. I, p. 389 (Voronezh, Russia).
1770. [Mus citellus] var. gutfatus Pallas, Nov. Comm. Acad. Sci. Imp.

Petrop., XIV, pt. I, p. 506, pi. 21, fig. 2 (Doubtfully a name).
1792. [Arctomijs] citellus S Icucopictus Donndorff, Zool. Beytrage, i, p. 486

(Renaming of guttatus).
1845. Sp\ennophilus'\ guttulatus Schinz. Synop. Mamm., ii, p. 70 (Renaming

of guttatus).
1910. Citellus guttatus Trouessart, Fauna ]\Iamm. d'Europe, p. 128.

Type locality. — Voronezh, Chernigof, Russia.
Geographical distrihution. — 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 incisive 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

3 0

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MARMOTA 931

biitfy white, the spots roundish, varying 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-buflf 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-buff"; heel and
ankle clear russet. Tail a dull buffy russet below, fringed with
cream-buff', a grizzle of blackish and cream-buff" above.

SJcull and teeth. — The only skull examined is imperfect. It
is smaller than that of C. cifellus, and apparently less deep at
middle, though dorsal profile is similarly convex throughout.
Rostrum narrower and not so deep as in C. citellus, 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. citellus, 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 citellus.

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 cxamiyied. — 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 =

marmot a).

1780. Arctoniys 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 (hudsonhis =

monax).

1857. Arctomys Blasius, Saugethiere Deutschlands, p. 278.

1904. Marmota Trouessarfc, 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 Jliller,
Ann. and Mag. Nat. Hist., 7th ser., xvi, pp. 461-464, October,
1905).

Type species. — -Marmota alpina Blumenbach = Mits marmota
Linnceus.

Geographical distribution. — Northern Hemisphere from the

3 o 2

932 EODENTIA

Alps eastward through Asia and North America to the Atlantic
coast of the United States.

Characters. — Strictly terrestrial Sciuridse, 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 CiteUus, 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.

Bemarks. — About two dozen species of Marmoia 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 EUEOPEAN SPECIES OF MABMOTA.

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. marmoia, p. 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 pale muzzle, the tail inconspicuously
blackish at tip ; skull with posterior border of
postorbital iJrocess nearly over narrowest portion
of postorbital constriction (eastern Hungary) ... M. bobak, p. 937.

MARMOTA MARMOTA Linna?US.

1758. [Miis] marmota Linnaeus, Syst. Nat., i, 10th ed., p. 60.

1779. [^Mannota'] alpina Blumenbach, Handb. der Naturgesch., i, p. 80
(Substitute for Marvwta).

1801. A[rctoviys] m[armota'] tigrina Bechstein, Gemeinn. Naturgesch.
Deutschlands, i, 2d ed., p. 1029.

1801. A[i-ctomys] m[an7iota^ alba Bechstein, Gemeinn. Naturgesch. Deutsch-
lands, I, 2d ed., p. 1030.

1801. A[i-ct07nys'] vi[annota] nigra Bechstein, Gemeinn. Naturgesch.
Deutschlands, i, 2d ed., p. 1030.

1857. Arctomys marmoia Blasius, Siiugethiere Deutschlands, p. 280.

1904. [Marmoial marmotia Trouessart, Catal. ]\Iamm. Viv. Foss., SuppL,
p. 343.

1910. Marmoia marmoia Trouessart, Faune Mamm. d'Europe, p. 129.

Type localifj/. — Alps.

Geograpliical distrihufion. — Alps and Carpathians. Frequently
referred to as occurring in the Pyrenees, but this probably an
error.*

Diaiinoxh. — Size medium (head and body about r).")0, tail

* See Trutat, Bull. Soc. d'Hist. Nat. de Toulouse, xi, p. 110, 1877.

MARMOTA

933

about 150, hind foot ahout i)3 inni. ; condylobasal leiigtli of fully
adult skulls. 92 to 98 nun.) ; 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 cliararters. — 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 nairow 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 jaads 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 phalanx 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. Mammaj :j^1 — 1; «2 — 2; i 2 — 2 = 10.

Colour. — Upper parts a grizzle of black and light bufl', the
exact proportions of the two sufliciently variable to produce
considerable individual differences in the general effect, but top
of head usually blackish finely ticked with whitish bufl" 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 bufiy 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 buffy white ; underparts and inner surface of legs
a clear yellowish tawny-buff, this colour suffusing outer surface
of legs and sides of body, also a conspicuous area extending
upward immediately behind shoulder and partly separating dark

* 'S'erified in twenty skins.

934

i-egion of neck from that of back ; feet buff", somewhat liglitei-
than outer surface of legs ; tail black or blackish brown, sufiused
to a varying degree with buff, the light colour most conspicuous
at and near base.

Shill. — Apart from its much greater size the skull of Marmota
marmota resembles that of Citellus 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 result that dorsal profile while equally convex in

Fic;. 190.
Mannola marmota. Slightly reduced.

front is noticeably flattened posteriorly as compared 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
Citellus 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

Fig. 191.
Marmota marmota. Slightly reduced.

Flo. 192.
Mannota iiianiwta. .Sliglitlv reduced.

936

of brain-case ; paroccipital process robust, not very long, its
extremity projecting silghtly beyond level of lower edge of
condyle ; basioccipital broad and short, 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
resi^ects the skull so closely resembles that of Citellus citeUus,
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-shaj^ed 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 vi^ flattened rather than basin-shaped. Lower cheek-teeth
diftering 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

Via. 193.
ilarmota inamnuta.
Cheek-teeth. X 2.

MARMOTA 937

the enamel pattern ; premolar with a small but c%i(leut cingulum
cusp on anterior boi-der.

Mcasureme.Hts. — Adult male from Barcelonnette, Basses- Alpes,
France : 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.

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

France: Near Barcelonnette, Basses-Alpes, 8 (B.M. and Mottaz).

Switzerland: Salanfe (at foot of Dent du Midi), Valais, 1 (Mottaz);
INIeiringen, Bern, 2 ; Engstlen Alp, Unterwalden, 1 ; Engelberg, Unter-
walden, 1 (U.S.N.M.) ; Scliaufigg, Grisons, 3 ; Canton Grisons (no exact
locality), 4 (U.S.N. INI.) ; Rheinwaldhorn, Grisons, 1.

Italy : Vagna Valley, near JIte. Rosa, Novara, 1 (Genoa) ; Aosta
Valley, Turin, 20 (Turin) : Argentera, Cuneo, 3 (Turin) ; llaritime Alps, 1
(Genoa).

Austria-Hungary : Felkaerthal, Tatra Mountains, Hungary, 1.

2(^,2?. Barcelonnette, Basses-Alpes, 0. Thomas (p). 8. 8. 10. 145.

Prance. (C. Mottaz.) 63-65.

Skull. Meiringen, Bern, Switzer- Tomes Collection. 7. 1. 1. 195.
land. [Koeserman.)
1. Engelberg, Unterwalden. Tomes Collection. 7. 1. 1. 131.

{Koeserman.)
S, 2 ?. Schaufigg, Grisons. 0. Thomas (p). 2. 8. 4. 31-33.

[Zollikofer.)
S. Rheinwaldhorn, Grisons. O. Thomas (p). 2. 8. 4. 30.

{Zollikofer.)
1. Felkaerthal, Tatra Moun- Poprad Museum 7. 4. 19. 1.
tains; Hungary. (e).

MARMOTA BOBAK Miiller.

1776. Mas hohak P. L. S. Miiller, Natursyst. Suppl. u. Regist. -Band, p. 40

(Poland).
1778. Mus ardomys Pallas, Nov. Sp. Quadr. Glir. Ord., p. 75 (Poland).
1780. Ardomys hohac Schreber, Siiugethiere, pi. ccix (description, iv,

p. 738, 1782). Renaming of Mus ardomys.
1811. Ardomys baibac Pallas, Zoogr. Rosso-Asiat., p. 155 (Poland).
1857. Ardomys bobac Blasius, Siiugethiere Deutschlauds, p. 283.
1904. [Mar7tiota] bobac Tro^essart, Catal. Mamm. Viv. Foss., Suppl., p. 343.
1910. Marmota bobac Trouessart, Faune Mamm. d'Europe, p. 129.

Type locality. — Poland.

Geographical disfribution. — From Poland and eastern Hungary
(Galicia and Bukowina) eastward into Asia. Exact limits of
range not known.

Diagnosis. — Size essentially as in Marmota marmota ; colours
uniform, the entire animal a yellowi.sh 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

938

EODENTIA

somewhat heavier than those of M. niarmota, especially the lower
molars ; anterior face of incisors whitish or pale orange.

External characters. — Essentially as in Marmota mariiiota, but
thumb present as a small though distinct tubercle bearing an
evident appressed nail. Mammae : p 2 — 2, a 2 — 2, / 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 (jbscuring the

Fig. 194.
.VariHiita hohak. 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.

ShuU 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
less narrowed posteriorly, their combined breadth at level of

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940 RODENTIA

posterior border (jf premaxillary fully three-fourths greatest
combined breadth anteriorly. Orbit lai'ger than in the related
species, the postoibital 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 bullte larger than in Marmota marmota, the area of
each bulla appearing greater than that of basioccipital and con-
dyles instead of about equal to it. Palate relatively narrower
^josteriorly, and incisive foramina wider in proportion to their
length than in M. marmota. Mandible more I'obust than in the
related animal, e.specially 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 exaviincd.—Tv/o, both from Russia.

Family PETAURISTID^.

1855. Ptermnyinn- Brandt, M6m. Acad. Imp. Sci., St. Petersburg, Gth ser.,

Sci. Nat., VII, p. 151.
1857. Sciurina Blasius, Saugethiere Deutschlands, p. 2G6 (part).
1879. Pteromichu Anderson, Anat. and Zool. Researches, Expeds. Yunnan,

p. 278.
1891. Sciurimr Flower and Lydekker, Mammals, living and extinct, p. 450

(part).

Geographical distrihutlon. — 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 stifi" cartilaginous process growing from
wrist, the membrane serving as a parachute by means of which
gliding flight is effected.

Remarks. — The Petanristldse are so sharply difl'erentiated from
the Sciuridce, that it seems preferable to recognize the two groups
as families. Eight genera are known,f one of which is repre-
sented in Europe.

* This difierence 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.

SCIUROPTERUS 941

Genus SCIUROPTERUS F. Cuvier.

1825. Sciuropterus F. Cuvier, Dents des Mammif^res, p. 255.
1857. Pteromys Blasius, Saugethiere Deutschlands, p. 268.

Ty2Je species. — Scitt,rus volans Linnseus - Pteromys russicus
Tiedemann.

Geof/i'aphical distrilmtion. — As in the family.

Characters. — No membrane extending between tail and hind
leg ; structure of molars essentially as in Schirus, the enamel not
thrown conspicuously into wrinkles, the transvei'se ridges well
developed ; posterior upper premolar about as large as first molar,
never decidedly smaller.

Hemarhs. — 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. [Scmr2(s] volans Linnaeus, 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[ero7nys] vulgaris Wagner, Schreber's Saugthiere, Suppl., iii, p. 228

(Substitute for volans).
1857. Pteromys volans Blasius, Saugethiere Deutschlands, p. 269.
1908. Sciuropterxis russicus Allen, Bull. Amer. Mus. Nat. Hist., xix, p. 132

March 31, 1903.
1910. Sciuropterus russicus Trouessart, Fauno 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 difierent 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 very large ; ear low,' rounded,
almost concealed in the fur, the outer margin faintly concave

942 EODENTIA

above, both surfaces clothed with fine short hairs which never
form tuft at tip. Feet relatively shorter and broader than in
S. viilr/aris, 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 claw of second,
fifth a little beyond base of first phalanx of fourth ; palmar and
plantar tuVjercles 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.

Culour. — Upper parts a uniform pale, silvery grey, with a
faint bufi'y cast, the slaty under colour appearing irregularly at
surface when hairs are disarranged, particularly near outer edge
of lateral membrane, and on feet ; e\-ery where 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 narrcjw blackish eye-ring ; underparts and inner
surface of limbs dull bufFy white, everywhere inconspicuously
sprinkled with blackish hairs ; tail more bufiy than body (nearly
the cream-bufi" of Ridgway), the upper and lower surfaces and
the tip much clouded with dusky, the sides almost clear.

Skull. — As compared with that of Sciurns 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 Seiurus rnlgaris, but
nasals abruptly curved downward at tip, occiput slightly C(mcave
owing to the more posterior position of condyles and paroccipital
processes due to enlargement of bulla?, and ventral profile more
nearly parallel with dorsal profile, its posterior portion carried
further downward by the large bulla?. 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 essentialh' 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
temporal fossa at antero-external i-orner of parietal, nmfh as in
Lepus ; lambdoid crest low and rounded ; viewed from the side
the brain-case is more depressed than in Seiurus vulgaris, so that

SCIUROPTEKUS

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 Cifelhis ; auditory bulhe sub-circular in general
outline, their form essentially as in Sciurus vulgaris, but their

Via. 195.
Sciuropterus russicus. Nat. size.

size actually as well as relatively greater, the antero-posterior
diameter equal to distance from front of bulla to front of m^, the
lateral diameter to outer margin of meatus fully IJ times least
width of basioccipital ; anterior upper margin of meatus developed
into a conspicuous forward-curved flange ; petrosal 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 tliat bases of postorbital 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

944 RODENTIA

actually as well as relatively larger than in Sclurus vulgaris, and
less well defined posteriorly owing to aljsence 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 u})per 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 by 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 emarginate 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 dia.stema ; 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
wii, 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.

rct'/^.—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^,

SCIXJROPTERUS

945

cut Iiy ;i deep posterior re-entrant fold, so that 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 m^, iir and m^, a short pi'ocess extends outward
from base of inner cusp into space between first and second
transverse ridges, while in pni^, iii^
and nr a similar but less developed
process lies in space between second
and third ridges ; third and fourth
ridges of in-^ 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 relativ^e sizes
and in general form except that nt.^
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 connecting 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 ; autero-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 cusj:) ; a similar but larger depression behind base of
postero-internal cusjj of m.^, the cingulum rising back of it so
as almost to form a supplemental cusp.

Measurements. — 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. 946.

Scluroptcrus russims.
Cheek-teeth, x 5.

Specimens examined. — Two from Finland (B.M. and U.S.N.M.
one from Russia, and a stufied specimen from unknown locality.

also

1. Finland.
1. Russia.
1 st. Europe.

Dr. H. Schulman (p).
Purchased (Parzudaki)
Purchased (Brandt).

1. 6. 9. 1.
53. 12. 6. 29.
■12. 5. 16. 41.

946

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Teeth moderately worn.

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Locality.

Finland .
Russia .

947

Family CASTORID^.

1821. Casioridn: Gray, London Med. Repos., xv, p. 302, April 1, 1821.

Geoijraijhical distribution. — Forested portions of the Northern
Hemisphere, south to the Mediterranean region and tlie southern
United States.

Characters. — Essentially as in the Sciuridne, but skull without
postorbital processes, and cheek-teeth rootless, with I'e-entrant
enamel folds ; premolars j ; angular portion of mandible short,
broadly rounded ; size large ; form heavy ; tail broad, flattened
from above downwards, its surface scaly ; habits aquatic.

BemarJcs. — Though several extinct members of the group are
known the family Cadoridse is represented by only one living
genus, widely distributed in the northern portion of both the
Old and New AVorkls.

Genus CASTOR Linnreus.

1758. Castor Linn^us, Syst. Nat., i, 10th ed., p. 58 (flher by tautonymy).
1806. Fiber Dumeril, Zoologie Analytique, p. 18, diagnosis on p. 19.
(Substitute for Castor.) Not Fiber Cuvier, 1800.

Ti/pc species. — Castor fiber Linnaeus.

Geographical distribution. — As in the family.

Characters. — General characters as in the family ; upper
molars sub-equal, each with one internal and three external
enamel folds.

EcntarJiS. — 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 Linnjeus.

1758. [Castor] fiber Linnseus, Syst. Nat., i, 10th ed., p. 58 (Sweden).

1792. C[astor] f[iber] albus Kerr, Anim. Kingd., p. 222.

1792. C\_astor] f[iber] solitarius Kerr, Auim. Kingd., p. 221.

1801. Clast07-1 /[iber] variegatus Bechstein, Gemeiun. Naturgesch. Dcutsch-

lands, I, 2d ed., p. 913 (Europe).
1801. C[astor] f\ibcr] fulviis Bechstein, Gemeinn. Naturgesch. Deutsch-

lands, I, 2d ed., p. 913 (Europe^.
1803. Castor galluv Geoffroy, Gatal. ]\Iamm. du Mus. Nat. d'Hist. Nat.,

Paris, p. 168 (Banks of the Rhone, France).

1822. Castor niger Desmarest, Jilammalogie, pt. ii, p. 278 (no exact

locality).
1822. Castor varitis 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. Mamm., p. 287 (Substitute

for gallisv).

948

KODENTIA

1833. Castor inopnus Billberg, Linn. Samf., p. 34 footnote (Substitute for
fiber).

1857. Castor fiber Blasius, Saugethiere Deutschlands, p. 405.

1897. Castor fiber Collett, Bergens ^Museums Aarbog, pp. 3-127. Habits in
Norway.

1907. Castor alhicus Matsohie, Sitz.-Ber. Gesellsch. Naturforsch. Freunde,
Berlin, p. 216, October, 1907. (Dessau, Anbalt, Germany.) See
Lonnberg, Arkiv for Zoologi, v, no. 6, pp. 1-16, 1909.

1907. ? Castor vistulinus Matschie, Sitz.-Ber. Gesellscb. Naturforsch.
Freunde, Berlin, p. 219, October, 1907 (western Poland).

1910. Castor fiber Trouessart, Faune JMamm. d'Europe, p. 130.

Ti/jx' localitij. — Sweden.

Geofirapliical distrihution. — Formerly the entire forested region
of Europe, west to Great Britain ; now exterminated everywhere
except in portions of Scandinavia, and along the courses of some
of the larger rivers of central Europe, as the Rhone, Elbe, and
Danube.

Diai/noiilK. — General features as in the family and genus ;
jiosterior extremity of nasals lying at or slightly behind level
of middle of orbit and decidedly behind termination of nasal
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
liead 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 rounded, 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 biistly,
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 live, 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 conca\e 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, that of fmirtli digit witli a coiispiouous, lionij'.
lateral ly-conipressed supplement fully as large as claw itself
springing from ball of finger beneath claw ; entire sole naked,
the surface coarsely reticulate ; three ek)ngate, ill-detined 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, 4-6-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.

Cohmr. — General colour a peculiar clayey buff produced b}'
the longer hairs, and very uniform throughout, the underparts
a little less yellowish ; underfur light smoke-grey at base,
darkening to hair-brown or bister at tips, this colour rarely
showing through at surface ; muzzle, chin, and lower part of
cheeks greyish but not forming any decided contrast with
suiTounding parts ; feet washed with a drab brown.

Shill. — In general aspect the skull is low, robust and heavily-
built, though not conspicuously 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 bulhe standing out con-
spicuously just in front of mastoid protubei-ances and rising to
level of dorsal surface of brain-case ; sagittal crest moderately
develoj^ed in adults, highest posteriorly, dividing anteriorly into
two low ridges curving forward and outward to rudimentarj'
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 magnvmi ; floor of brain-case with deep pit
occupying entire area between bullae, its edges well defined
throughout, its posterior margin formed by lower lip of foramen
magnum ; auditory bulla flask-shaped, the meatal tube about as

CASTOR 951

long as tmnsvei'se diameter of inflated portion of bulla, con-
sisicuously ridged postero-externally, the main axis of flask
oblique but nearly upright, sloping outward at an angle of only
about 30^ ; portion of bulla apjjearing on lower surface of skull
about li times as long as wide, evenly inflated, the width at
middle about eijual to that of transverse diameter of basioccipital
pit measured inside the rim ; bulla separated posteriorly and
postero-externally from paroccijMtal 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 exjjanded at middle, its upper edge with projection
marking postei-ior limit of orbit, its depth through this jsrojection
rather greater than diameter of orbit, its anterior extremity
firmly 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 first 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 before
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 border slightly behind level of m^, its
median spine well developed ; mesopterygoid sjiace 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
border 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-off angular process.

Teetli. — Upper incisor heavy, its course scarcely indicated
on outer surface of I'ostrum, its root lying behind anteorbital
foramen ; shaft about as deep as wide, the anterior face slightlj^
curved, a little longer than sub-equal outer and inner faces, the
posterior border abruptly rounded ; enamel dark yellowish brown
in colour, essentially smooth but with very obscure longitudinal

952

EODENTIA

wrinklfts ; lower incisor with root extending slightly beyond
in.,, 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
upper 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 maxillary
teeth nearly straight, extending
oljlitjuely forward to come in con-
tact with extremity of first outer
fold, or frequently in pm* to curve
slightly behiiid 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 obli(]uely forward
half-way across crown, or in piii* occasionally about to inner
l)order, the second outer fold curves noticealjly backward and
extends nearly to inner side of crown, and the third outer fold
extends directly inward or somewhat oljliijuely 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 wavj^ 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 liody 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

Via. 108.
Ca.itor filer, riieek-teftli.

Specimens examined. — Six, from the following localities
Norway : Froland, Kristiansand, 1.
England : Norfolk, 1 (skull).

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954 EODENTIA

France : Rhone, near Aiies, 2 ; near Avignon, Vaucluse, 1.
Germany : Elbe Kiver, 1 (U.S.N.M.).
Austria-Hungary : Danube, 1 (skull).

HehiarTis. — 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.

c?, skeleton. Froland, Kristiausand, Dr. R. Collctt (p). 97.10.14.1.
Norway.

Skeleton. Norfolk, England (from W. -T. Angove (c). 85. 8. 4. 1.
peat-bog).

6. Aries, Bouches du Rhone, Lord Lilford (p). 94. 5. .30. 1.

France. [Dr. P. Siepi.)

9. Aries, Bouches du Rhone. Dr. P. Siepi (c). 5. .3. 9. 1.

1. Avignon, Vaucluse. Purchased (Dev- 85. 5. 20. 1.

rolls).

Skull. Danube, Austria-Hungary. Purchased (Brandt). 875. b.

955

Order UNGULATA.

1801. Ungulata Boclistcin, Gemciun. Naturgesch. Dcutsclilands, i, 2iid
ed., p. 182.

Gedgraphlral diatrihiition. — Essentially throughout all o£ the
lari^er laud masses of the world, Austi'alia excepted.

ClttnaQtcvH. — Terrestrial, herbivorous or tminivorous, placental
uiauimals with broad-crowned, tuberculated or ridged molars,
digitigrade feet, and hoofed digits.

liemarlcs. — The exact limit of the order Umjulata is still
a matter of controversy. The group is here considered as
equivalent to the Diphtrthra. Thus restricted it contains two
main groups, the Artiodarfi/la, in which the main axis of the
foot lies between the third and fourth digits, producing the well-
known cloven hoof, and the Perissodacti/la, 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 Artiodacfiila. The genus
Eqmii<, representing the Pcrissodnctyln, probably disappeared as
a natural member of the fauna of western Europe during paleo-
lithic times. The Artlodactyla 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 Unguhda of Europe are so inadequately represented in the
British and other Museums 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 entered at second hand.

ARTIFICIAL KEY TO THE EUROPEAN FAMILIES OF
ABTIODAGTYLA.

Upper incisors present ; lower canine not incisor-like ;

horns absent (Pigs) Suid:r, 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.) Bovida-, p. 986.

Horns (antlers), consisting of a solid bony outgrowth
from the frontal bone, periodically shed and re-
newed, often absent in females (Deer) Cervidsv, p. 962.

956

UNGULATA

Family SUTD^..
1S21. Suichr Gray, London :Med. Eepos., xv, p. 30G, April 1, 1821.

Geographical distribution. — Warmer 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 supported by
a well developed free ossicle (not shown in figs. 199 and 200) ;
inci.sors 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.

Reiiiarhs. — The family Snirlse contains at least six well-
defined living genera. Only one of these, the typical »S'».«, occurs
in Europe.

Genus SUS Linnaeus.

1758. Snx LinnffiUR, Syst. Nat., i, 10th ed., p. 49.

176C. Ajxr Pallas, Miscellanea Zoologica, p. 10 (Substitute for Snx).

1815. A2}er Rafinesque, Analy.se de la Natuie, p. 5G (Substitute for Sns).

1857. Sns Blasius, Siiugethiere Deutschlands, p. 508.

1SG8. Scrofii, Gray, Proc. Zool. Soc, London, p. 38 (Domestic Pig).

Ti/jje .aperies. — Siis scrofa Linna'u.s (by tautonymy).

Geographical distrihntion. — Same as that of the family Suidff,
except that in Africa it is confined to the Mediterranean region.

Characters. — Dental formula : i jjE;!;, r ,3,, pm ^E;, m '.^'.. = 44 ;
incisors and premolars tending to remain functional throughout
life, though the small /'', piti'^ and pm^ are not infrec[uently lost :
bunodont structure of molars showing no special modification ;
carunes small in females, moderately large and conspicuf)Usly
tienchant in males ; skull elongate, high and narrow, typically
suine and without conspicuous modifications.

Remarlcs. — 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 THPJ EUROPEAN SPECIES OF SUS.

Upper length of skull in adult males usually more

than 350 mm. ; ?«^ with third transverse ridge

well developed (Distribution general) S. scrofa, p. 957.

Upper length of skull in adult males usually less

than 300 mm. ; vi^ with third transverse ridge

obsolete (Sardinia) S. mcridionalis, p. 960.

957

SUS SCROFA Linn?eus.

1758. [Sus] scrofa Linnfeus, Syst. Nat. i, 10th ed., p. 49.

1785. [Sus] 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 europnus Pallas, Zoogr. Rosso-Asiat., i, p. '265 (Substitute for

scrofa).
1857. Sus scrofa Blasius, Siiugethiere Deutsehlands, p. 510.
1882. \_Sus scrofa] var. celtica Strobel, Atti Soc. Ital. Sci. Nat., Milauo,

XXV, p. 79, June, 1882 (Renaming of true scrofa).
1910. Sus scrofa Trouessart, Paune, Mamm. d'Europc, p. 225.

Tijije locaUfi/. — Germany.
Geograj^liical distribution

-Soutlicrii and central Eiiropf,

Fig. 199.
Sus scrofa. X j.

formerly we.st to Ireland and north to southern Norway and
Swedeii ] now restricted to that portion of-- the Continent lying

958

UNGULATA

sf)uth of tlic Baltic, aiicl occurring in foreslcd regi<jns only ;
oastern limit of range unknown.

Diagnosis. — Size rather large, the length of head and body
about 1 ■ 5 m. in full-grown males, somewhat less iri 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 different individuals tending to become blackish,
gi'eyish or reddish, the nature of these variations not under-
stood ; face, cheeks and throat grizzled by sprinkling of whitish

Kir,. 20(1.
■ .srro/ft..

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 evei\ywhere 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 crown, 115 ; width of ear, 90; length of hairs at upper
edge of ear, 30. For cranial measurements see Table, p. 961.

sus 959

Specimens exanmicd.— Sixteen, from the following localities:—
Prance : No exact locality, 1.

Germany: Harz Mountaius, 1 skull (U.S.N.M.) ; Waldleinigen,
Odcnwald, N. Baden, 1 ; Wiirtemberg, 2 (skulls) ; no exact locality, 2.

fe:

J'IG. 201.
Sris scrofa. Cheek teeth, luit. size.

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

DNGULATA

HeniarJi's. — Though it seems probabk^ that 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 Sus ncrofa was in type three new forms
have been recognized by Thomas : —

Siis attila Thomas, Abstr. Proc. Zool. 8oc. London, No. 105,
p. I, March 12, 1912. Type locality, Koloszvar, Austria-
Hungary. Distribution, Hungary and Kussia. Much larger
than SiiH 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
Wertheimstein .

/S'jts scrofa rdstiJianiis Thomas, Abstr. Proc. Zool. Soc. London,
No. 105, p. 1, March 12, 1912. Type locality. Province of
Burgos, Spain. Distribution, northern Spain. Smaller than
true Scrofa (uj^per length of skull, 353 mm. ; height at
occiput, 198).

Siis scrofa bseticns Thomas, Abstr. Proc. Zool. Soc. London,
No. 105, p. 2, March 12, 1.912. Type locality, Goto Doilana,
Huelva, Spain. Distiibution, southern .Spain. Smaller than
rastilianu.s (upper length of skull, 324 mm. ; height at occiput,
208).l

1 (juv.;

6.

6 skull.
J skeleton.

i skull.
6.

2 9.

9.

6.

6 skull.

Oden-
Baden,

France.
Waldleiuigcn,

wald, N.

Germany.
Wurtembcrg.
Germany.

Germany.
Province of Burgos.
(liev. S. Gonzalez

Purchased (Lefcbvre). 43. 12. 29. 12.
H.R.H. Grand Duke 92. 8. 3. 1.

Louis of Hesse

(c &r).
Dr. A. Giinther (c).
Zoological Society's

Museum.
Purchased.
Hon. N.C.Rothschild

(I')-
(Type of S. s. castilianus Thomas.)
S. & N. Gonzalez (c). 8. 7. 7. 32-33.

59. 9. 6. 100.
713. 51.

02. 3. 20. G.
11. 10. 5. 3.

Quiutanar de la Sierra

Burgos.
Almonte, Seville, Spain. Lord Lilford (r). 95. 9. 4. IG.

{A. Ruiz.)
Goto Dofiana, Huelva. A. Chapman (c k r). 8. 3. 8. 12.
(Type of S. s. bndicus Thomas.)
Goto Douana. A. Chapman (c & p). 8. 3. 8. 13.

SUS MERIDIONALIS 31ajor.

1881. Sua scrofa mcridionalis Major, Atti Soc. Tosc. Sci. Nat., Pisa, Proc.

Verb., in, p. 119 (Sardinia ; the wild pig). Fully described in
Vol. VI, fasc. 2, pp. 316-362, 1883.

1882. ? Sus scrofa var. sardoiis Strobel, Atti Soc. Ital. Sci. Nat., Milano,

XXV, p. 221 (Sardinia; the domestic pig).
1910. Siis scrofa sardo^ts Trouessart, Fauna ]\Iamm. d'Europc, p. 226 (the
wild pig).

Tiifie locality. — Sardinia.

Gco(jruj)h leal distrihutiun. — Sardinia.

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962 UNGDLATA

Diaijno.sis. — Like Sns scrofa but not attaining so great size,
the upper lengt.-- of skull in adult male about 300 mm., upper
tooth-row including canine 114 to 128 rnm. ; posterior molar
both above and below with third transverse ridge much reduced ;
colour in both adult and young essentially as in Sus scrofa.

3Teasiiremcnts. — For cranial measurements see Table, p. 961.

Specimens examined. — Five skulls (Genoa).

RemarJcu. — The wild boar of Sardinia apf)ears to be a well-
defined local form.

Family CERVIDtE.

1820. Cervidn: Gray, London Med. Repos., xv, p. 307, April 1, 1821.

Geographiral distrlhution. — 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.

Characfrrs. — 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.

Beviarkft. — About sixteen genera of living deer are known, a
number which will probably be inci-cased by more detailed study.
Three of these, Cervns, Alrr.<i and Bam/if er, 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. Fi\e occur in Europe.

KEY TO THE EUROPEAN GENERA OF CEEVIDJ^.

Vomer high, dividing posterior nares into two chambers ;
no marked contrast in size between anterior pre-
molar and m- ; width of upper molars about one-
fourth 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) Ilavrjifer, p. 979.

Vomer low, not dividing posterior nares into two
chambers ; a marked contrast in size between
anterior premolar and vi- ; width of upper molars
about one-third that of palate ; antlers normally
present in males onlj', 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 noticeabh' longer than wide ;
muzzle not entirely hairy ; antlers not beginning to
develop until about a year after birth.

CERVDS 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) Alces, 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) Cervus, p. 963.

Maxillary canines absent, antlers either not terete
or not spreading, the complement of tines not
entire.
Auditory buUse evenly inflated, their surface with-
out conspicuous ridges ; antlers spreading,
without bez tine, the distal portion narrowly
palmate; tail well developed (Fallow deer)... Daww, p. 970.
Auditory bullce collapsed, their surface con-
spicuously ridged ; antlers erect, terete, with-
out brow tine ; tail reduced to an incon-
spicuous papilla (Roe Deer) Capreohts, -p. 973.

Genus CERVUS Linnixms.

1758. Cervus Linnmus, Syst. Nat., i, 10th ed., p. CG.

1827. Elaphus Hamilton Smith, Griffith's Cuvier, Animal Kingdom, v,

p. 307 (Substitute for Cervus).
1857. Cervus Blasius, Saugethiere Deutschlands, p. 438 (part).
1899. Eucervus Acloque, Faune de France, Mamm., p. 71 (not of Gray,

1866). Substitute for Cervus.

Type sjjecies. — Cervus elaphus Linnajus (by tautonymy).

Geograpliical distribution. — North temperate region of l)otli
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 chamloers ;
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 orbital cavity ; tail well
developed, moderate ; muzzle naked ; young spotted with white,
their colour very different from that of adults.

Hemarlcs. — The genus Cervus as thus defined contains the
Red Deer group and its immediate allies. The exact limits of
tbe genus are not definitely known ; Init there* can be no doubt

3 Q 2

964

UNGDLATA

that the name Cervus is currently applied to several groups
which should be recognized as distinct.

It is not possible to attempt any I'evision 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 prol)ability that this course is
final I nm treating all the European forms as geographical races
of Cervvs elajilinfi.

CERVUS ELAPHUS LinnffiUS.

(Synonymy under subspecies.)

Geofira'plncal (lisiribution. — Forested regions (where not ex-
terminaterl) fi'oni Si)aiu and Ii'eland eastward, ajid fi'om Sardinia

KlG. 202.
Cervus elaphus. X \.

cKRvus 965

and the ModitLirraiieiiii coast to northern Seotlaml and central
Sweden ; an isolated race on , the coast of central Norway ;
absent from the peninsula of Italy ; eastern limit of range not
known.

Diagnosis. — 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, gre^'ish browu in winter.

FIG. :;o.3.
Ccrvus claphus. Inuisiform teeth. Nat. size.

the underparts 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 wliich often occurs in region of " cup " in old individuals ;
in fully developed antlers there are normally present a brow tine,
bez tine and more than 5 points.

CERVUS ELAPHUS GERMANicus Desmarest.

1822. Ccrvus claphus germanicus Desmarest, Mammalogio, ii, p. 43i

(Germany).
1822. ? Ccrvus elaphus albus Desmarest, JIammalogie, ii, p. 435 {nomeii

nudum).
1815. Ccrvus claphus albifrons Reichenbach, Vollstiiudigste Naturgesch.

des In- und Auslands, Siiugeth., iii, pi. ill bis, fig. 26 (no exact

locality; see p. 18).
1857. Ccrvus claphus Blasius, Siiugethiere Deutschlands, p. 439.
1874. Ccrvus claphus, varius Fitsiinger, Sitzungsber. kais. Akad. Wissensch.

Wien, Math.-Naturwiss. Classe, lxix, pt. i, p. 674 (Germany).
1874. Ccrvus claphus, albus Pitzinger, Sitzungsber. kais. Akad. Wissensch.

Wien, Math.-Naturwiss. Classe, LXix, pt. i, p. 575 (Germany).
1898. ? Ccrvus claphus viaral Lydckker, Deer of All Lands, p. 79 (part ;

specimens from the Galician Carpathians, Hungary).
1903. Ccrvus vulgaris Botezat, ilorphol. Jahrb., xxxii, p. 115, November 17,

1903 (Renaming of claphus).

96G

UNGULATA

1903. ? Clervus] vulgaris cam.j)cstris Botczat, !Morphol. Jahrb., xxxii, p. 154,
November 17, 1903 (Lowlands and beech forests on lower slopes
of Carpathian jNIountains, Bukowina, Austria-Hungary). Not
Cervus ccmipcstris F. Cuvier, 1817.

1903. ? C[ervus] vulgaris montami.s Botezat, Morphol. Jahrb., xxxii, p. 155,
November 17, 1903 (Carpathian Mountains, Bukowina, Austria-
Hungary).

1906. Cervus claiohus germanicus Lonnberg, Arkiv for Zoologi, iii, No. 9,

p. 14, January 2% 1906.

1907. ? Cervus baUicus INIatschie, Das Waidwerk in Wort und Bild, xvi,

]}. 186, March 15, 1907 (Forest near Liebemiihl, Ostpreussen,

Cxermany).
1907. '? Cervus albicus Matschie, Das Waidwerk in Wort und Bild, xvi,

p. 186, ]\Iarch 15, 1907 (Muskau, Oberlausitz, Silesia, Germany).
1907. ? Cervus rlicnanus Matschie, Das Waidwerk in Wort und Bild, xvi,

p. 186, INIarch 15,1907 (Viernhoim, Hessen-Darmstadt, Germany).
1907. ? Cervus bajovaricus Matschie, Das Waidwerk in Wort und Bild,

XVI, p. i86, March 15, 1907 (Rohner, Konigssee, Oberbayern,

Germany).
1910. Cervus elaphus germanicus Trouessart, Faune Manim. d'Europc,

p. 228.

Type locality. — Germany.

Oeoffraphical distrihutivii. — Continental Europe, limits of range
not known ; animals of this general type occur from the Mediter-
ranean i-egion to the Baltic and from western France to eastern
Hungary, though they pi'obably I'epresent more than one geo-
graphical race.

Diagnosis. — Size large, as in C. elajilru.'s chtplin.^ ; caudal tlisk
conspicuously lighter than Hanks and usually with a definite
black or blackish border.

Mcasurrmculs. — For cranial measurements see Tabic, p. 982.

Specimens examined. — Nine, from the following localities : —

France : No exact locality, 2.

Gebmany : Gohrdc, Hanover, 1 ; south Germany, 1 ; no exact locality, 1.

Austbia-Hdxgaiiy : Bohemia, 2 ; Carpathians, 1.

Italy : Carapo Carnico, Cadore, 1 (Turin).

licmarlis. — 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 balti.cus. — 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, Ostj^reussen,
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. TyjDe locality : Muskau, Ober-

(hornless).

(H.M. Empcroi
William I.)

? skull.

S antlers.

2 i antlors.

S. Germany.

Germany.

Bohemia.

6 antlers.

Carpathians.

CERvus 967

lausitz, Silesia. Kaiige : middle Germany south of the Baltic
divide, west to the Weser divide, and south to tlie Main divide.

CervuH rhenanus. — Antlers with beam bent abruptly inward
at level of trez tine ; points of all tines directed upward. Type
locality : Viernheim, Hessen - Darmstadt. Range : western
Germany from the Harz Mountains to the Danube basin.

Cervus hajovaricns. — Antlers as in C. rhenanus, but with inner
tines directed inward and backward. Type locality : Rohner,
Konigssee, Oberbayern. Distribution : Oberbayern.

One or more forms of Red Deer probably distinct from
C. claplms tjermnnicus occur in eastern Hungary. They lla^'e
been referred to C. claplms maral (type locality, " the Persian
Mountains ") by Lydekker, and have been described as new under
the names carnpcstris and monfanus by Bozetat.

9 & yg. St. France. Purchased (Lefehvre). 43. 12. 29. 5 & 11.

6 frontlet Gohrde^ Hanover. J. E. Harting (p). 88. 6. 12. 1.

Dr. A. Giinther (c). 59. 9. 6. 103.

689. p.
Col. J. Evans (p). 89. 11. 20. 1-2.

H.H. Prince Heinrich 9G. 10. 10. 1.
von Liechtenstein (p).

Cervus elaphus elapiius Liunajus.

1758. iCervus'] claplms Linnaeus, Syst. Nat., i, 10th ed., p. 67.
1906. Cervus elaphus Lonnberg, Arkiv for Zoologi, iii, No. 9, p. 9,
January 22, 1906.

Tjipc localitij. — Southern Sweden.

Geographical distribution. — " The [range] .... extended in
bygone days probably over the greater pai-t of Gotaland. At
present the red deer is in Sweden confined to southern Skaiiia,
and there chiefly found on a few large estates, Hiickeberga,
Of \ edskloster, Borringe, Siifdeborg, Snogeholm, Skabersjij, etc.
The number is quite small, perhaps not more than about 100 in
all " (Lonnberg).

Diagnosis. — Size large ; caudal disk not conspicuously lighter
than flanks and never with a definite black border.

Eciaarlcs. — I have not seen this animal. From Lonnberg's
account it appears to be sufliciently different from the stag of
central Europe to merit recognition by name.

Cervus elaphus atlanticus Lonnberg.

1906. Cervus elaphus atlanticus Lonnberg, Arkiv f5r Zoologi, in, No. 9, p. 9,

January 22, 1906.
1910. Cervus elaphus atlanticus Trouessart, Fauno !Mamm. d'Europe, p. 228.

Type localitij. — Hitteren Island, Trondhjem, Norway.
Geographical dlstrihution. — West coast of Norway fi-om
Stavanger Fjord north to about latitude G5°.

968 UNGULATA

Diagnosis. — Size smaller and colour paler than in true Qtvus
elaj)hts ; caudal disk with distinct dark border.

Measurements. — For cranial measurements see Table, p. 9^2.

Specimens examined. — Two, one from Fredo, Norway, the other from
Gloppen, Nordfjord, Norway (both U.S.N.M.).

BemarJcs. — Though sharply differentiated from the Swedish
and central European forms, the Norwegian red deer closely
resembles that of Scotland."'

Cekvus klaphus scoticus Liinnberg.

190G. Ccrvus claphus scoticus Lonnbcrg, Arkiv for Zoologie, iii, No. 9,

p. 11, January 22, 1906.
1910. Ccrvus ela.phus scoticus Trouessart, Faune Mamm. d'Europe, p. 228.

Type localiti/. — Glenquoich Forest, Inverness-shire, Scotland.

Geographical disfribiition. — Great Britain ; limits of range
not known.

Characters. — Like Ccrrus claphus atlanticus l)ut colour darker
and less grey.

Measurcmeiits. — 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; Woliurn Abbey, Bedfordshire, 1; Exmoor,
Devon, 1 ; England, no exact locality, 4.

Iteniarlfs. — The status of the Bi-itish form of Red Deer is
not well understood. The cranial characters mentioned by
Liinnberg, as distinguishing the animal from atlanticus, are
inconstant, but there appears to be an appreciable difference
in colour between the two races.

6, 9. Balmacaan, Invcr- Bradley Martin (v). 9. 1. 15. 1-2.

ness, Scotland.
2 c5 antlers. Jura, Ardgour, Ar- F. Ilamilton-Lcigh (i>). 11.2.21.1-2

gyllshirc.
^ skull with- Loch Sunart, Argyll- Gen. 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.
(5 stuffed. Alnwick Chase, Duke of Northumbcr- 63.11.16.5.

Northumberland. land (r).
d stuffed. Woburn, Bedford- Duke of Bedford (p). 97. i. 3. 3.
shire.

* For discussion of the meaning of this resemblance see Stejneger,
Smithsonian Miscellaneous Collections, xlviii, pp. 462-469, May 4, 1907,
and Bergens Museums Aarbog, xiv, 1908.

CERVUS

yoi

9 skull.

Exmoor, Devon.

R. A. Sanders (r).

G. 2. 26. 1.

6 antlers.

England.

689. b.

6 antlers.

England.

Purchased (Lead-
heater).

46. 11. 20. 17.

6 skull.

England.

Purchased (Baker).

47. 12. 11. 16.

6 antlers.

(From a pcat-bog.

Jabez Allies (r).

49. 3. 5. 1.

Ckkvus elaphus hispanicus Hilzheimer.

1909. C[crviis] elaphus hispanicus Hilzheimer, Archiv fiir Rassen- und

Gesellschafts-Biologie, 1909, p. 313. (South-western Spain?)

Type in Stuttgart IMuseum.
1911. ? Ccrvus elaphus bolivari Cabrera, Bol. Real Soc. Espafi. Hist.

Nat. XI, p. 559, December, 1911 (El Pardo, Madrid, Spain). Type

in Madrid Museum.

Tyj'C localitij. — Spain, exact locality not known.*

Geographical distribution. — Iberian Peninsula, limits of range
unknown.

Diagnosis. — Size apparently less than in Cervns elaplms
scoticiis ; colour decidedly more greyish ; skull narrower,
particularly in interorbital and palatal regions.

Measurements. — For cranial measurements see Table, p. 982.

Hemarhs. — The Red Deer of Spain are divided by Cabrera
into a smaller southern race (Jiisjianicns) 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. — Pour, from Goto Donana, Huelva, Spain, and
two from Pinares de Quintanar, Burgos, Spain (U.S.N.M.).

d ?. Goto Donana, Huelva, Lord Lilford (i>). 95.9.4.14-15.

Spain. (A. Ruiz).
6. Goto Donana. A. Chapman (c & r). 8. 3. 8. 14.

o skull. Goto Donana. A. Chapman (c & p). S. 3. 8. 15.

Cervus elaphus coksicanus Erxleben.

1777. \Cervus'\ corsicanus Erxleben, Syst. Regni Anim., i, p. 304 (Corsica).
1822. C[ervus] mediterraneus Blainville, Journ. de Phys. Chem. Hist.

Nat., xciv, p. 262 (Corsica).
1855. \_Cervus elaphus] $ minor Wagner, Schreber's Siiugth., Suppl., v,

p. 354 (substitute for corsicanus and mediterraneus),
1910. Cervus elaphus corsicanus Trouessart, Faune Mamm. d'Europe,

p. 229.

Type localiti/. — Cor.sica.

Geographical distribution. — Corsicaf and Sardinia.

Diagnosis. — Size about as in Cervus elaplius hispanicus (height

* Dr. Hilzheimer informs me 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. iii) states that the stag is not native to Corsica.
(The Histories of Polybius translated by Evelyn S. Shuckburgh, ii, p. 80).

970

UXGULATA

at shoulder of mounted male in Tufiii Museum 800 mm.), Init
general colour darker than in any of the small continental forms.
Measurements. — For cranial measurements see Table, p. 984.

Specwiens examined. — Four, all from Corsica (Turin).

Genus DAMA Hamilton Smith.

1827. Dama Hamilton Smith, Griffith's Cuvier, Auim. Kiugrl., v, p. 306.

1844. Platyccros Wagner, SchreLer's Saugth., Suppl., iv, p. 317.

1855. Dactyloceros Wagner, Schreber's Saugth. , Suppl. v, p. 352 (Substitute

for Dama and Flatyceros).
1857. Ccrvus Blasius, Saugethiere Deutsehlands, p. 39 (part).
1893. MacJilis Zittel, Handb. Palaeont. iv, p. 402 (published as a synonym

of Dama with Kaup as authority).
1898. Palmatus Lydelskcr, Deer of all Lands, p. 125. Wrongly attributed

to Giebel, Saugethiere, p. 351 (a group name used in the plural :

" Palmati mit schaufelformigem Gewieh ").

Ti/i)e species. — Cervus dama Linnseus.

Geograpliical distrihution. — Mediterranean region of southern
J]urope and western Asia ; occurs in a condition of semi-
domestication as far ncn'th as the British Islands and southern
Scandinavia.

Characters. — Like Cervus but with skull shorter and relatively
much broader, orbits lai'ger, maxillary canines absent, cheek-
teeth, particularly mr, more bracli3^dont, lower incisors (fig. 20'))
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.*

Bonarks. — The Fallow Deer form a sharply defined group
generically distinct from Ccrvus. Two species are known, one of
which occurs in Europe.

DAMA DAMA Linn;i'us.

1758. [Cervus] dama Linnteus, Syst. Nat., i, 10th ed., p. G7.

1798. Cervus platyccros Cuvier, Tabl. Elem. do I'Hist. Nat. dcs Anim.,

p. IGO (Renaming of dama).
1816. L (sic) [ervus] mauricus F. Cuvier, Bull, des Sci. Soc. Philomath.,

1816, p. 72 (locality not known).

* The extinct genus Alee Blumenbach (Handb. d. Naturgesch., 6th ed.,
p. 697, 1799, type Alec giganteus Blumenbach from the peat bogs of
Ireland = " Ctvi,-z(s mcgnceros" Auct. or " 2Iegaccros Jiibernicus" 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 in the combination of xalesiometacarpalian 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

182'J. [Ccrvits llama] /3. leiicwthiops Fischer, Synops. Mamm., p. 448

(Germany).
1829. [Ccrvus dama] y. maura Fischer, Syuops. Mamm., p. 448 (Renaming

of mauriciis) .
1843. Dmna vulgaris Gray, List Spec. Mamm. Brit. JIus., p. 181

(Renaming of dania).
1857. Ccrvus dama Blasius, Siiugethiero Doutschlands, p. 4,53.
1874. Dama2}latyceros, nigerFitzmger,Sitzniigsher. jjais. Akad. Wissensch.

Wien, Math.-Naturwiss. Classe, lxix, pt.,i, p. 553 (Renaming of

mauriciis) .
1874. Dama platyceros, varius Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lxix, pt. i, p. 555

(Germany).
1874. Dama platyceros, alhus Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lxix, pt. i, p. 555

(Renaming of leucxthiops).
1885. [Dama] dama Lataste, Actes See. Linn, de Bordeaux, xxxix, p. 288.
1910. Dama dama Trouessart, Faune Mamm. d'Europe, p. 229.

Type localitt/. — Sweden (introduced).

Georjraphical distribution. — 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-
fluent ; antlers with brow tine large,

Fig. 204.
Dama dama. X

Dama dama. Incisiforra teeth.
Nat. size.

and with region of trez tine showing no tendency to become
flattened.

3Icasurt'ments. — Adult male : head and body, 15J:0 ; tail,

972

UNGDLATA

190 ; pencil, 100 ; hind foot (with hoof), 435 ; ear from crown,
165 ; width of ear at middle, 90. For cranial measurements see
Table, p. 983.

Specimens exafnined. — 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.)

<5st.

Woburn, Bedford-
shire, England.

Duke of Bedford (f).

96. 9. 24. 1.

<;st.

Tring Park, Hert-

Hon. Walter Roths-

98. 10. IS. 1.

fordshire.

child (p).

(5 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.

Nannan Park,
^Merionethshire.

J. E. Harting (p).

88. 6. 12. 3.

46 antlers.

New Forest.

Mrs. Smyth (p).

50. 2. 5. 1-46

i skull.

England.

Purchased.

50. 11. 13. 15

? skull.

England.

693. k.

2 .5 2 9 skulls.

Lidth de Jeudc Coll.

67. 4. 12. 234
235, 236, 241.

6 St.

Italy.

Purchased (Lefebvre).

43. 12. 29. 4.

Genus CAPREOLUS Gray.

1821. Capreolus Gray, London Med. Repos., xv, p. .307, April 1, 1821.
1837. Cajn-ca Ogilby, Proc. Zool. See. London, 1836, p. 135, June 27, 1837.
1857. Caprcolus Blasius, Siiugcthiere Deutschlands, p. 467.

Tjipc species. — Cervus caprcnJus Linnajus.

(jeograplucal distribution. — Central and soutlua-n Europe from
Great Britain and southern Scandina\ia eastward ; in Asia east
to the Pacitic coast.

Cliararters. — Telemetacarpalian deer of small size with narrow,
elongate hoofs, maxillary canines normally ab.sent,* lower incisors
distinctly though not excessively difierentiated in size and form,
lachrymal vacuity large, the pit reduced to a shallow, incon.spicuous
depression ; vomer low posteriorly, showing no tendency to divide
the po.sterior nares into two chamljers ; antler small,! ^^st 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 anteri(jr base extending over orbital
cavity ; tail reduced to an inconspicuous papilla ; muzzlt; naked ;
young with spotted coat very ditierent from pelage of adult.

* For account of their occasional occurrence see KoUiker, Wiirzburger
Naturwiss. Zeitschr., vi, p. 82, 1866.

t Sometimes present in females (see Sherren, Field, London, cxiv,
p. 751, October 23, 1909).

CAPREOLUS

973

Remarks. — About a dozen forms of Capreolus have been
described. One species, represented by four geographical races,
occurs in Europe.

CAPREOLUS CAPREOLUS Linnseus.
(Syuonymy under subspecies.)

Geographical distribution. — 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 1-50 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. 206.
Capreohis capreoltm

Fig. 207.

Capreolus capreoliis.

Incisiforni 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 conspicuous contrast.

Remarlis. — In western Europe Capreolus capreolus is repre-
sented by four well-defined local races,

KEY TO THE EUROPEAN FORMS OP CAPREOLUS
CAPREOL US.

General colour of face distinctly darker than that

of body (Great Britain) C. capreolus thotti, p. 975.

General colour of face not darker than that of body.

Pale throat patch and neck patch in winter

pelage sharply defined whitish (South-eastern

Europe) C. capreolua traii.tsylvani-

cws, 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. capreolus capreolus,

p. 974.
General colour of winter pelage a coarsely
grizzled grey without decided yellowish
tinge (Spain) C. capreolus canuR. p. 975.

974

UNGULATA

Capreolus capreolus capreolus Linna?us.

1758. [Cervtis] capreolus LiiiDaeus, Syst. Nat. i, 10th ed., p. G3 (Sweden).
1792. C[erV'Us] capreolus albus Kerr, Anim. Kingd., p. 302 (Franche Comt^,

France).
1829. ? Capreolus dorcas Burnett, Quart. Journ. Sci. Lit. Art, 1829, pt. 2,

p. 353 (nomen nudum).

1843. Capreolus capm'a Gray, List. Spec. Mamm. Brit. Mus., p. 17G

(Renaming of capreolus).

1844. C\jipreolus'] europiius Sundevall, Ofversigt af Kongl. Veteusk.-Akad.

Forhandl., Stockholm, i, p. 184 (Renaming of capreolus).

1845. C[erviis] capreolus pluinbeus Reichenhach, A'ollstandigste Natur-

gesch. der In- uud Auslands, Siiugeth., in, pi. in bis, fig. 53

(Germany).
1857. Capreohts capreolus Blasius, Silugethiere Deutschlands, p. 457.
18G0. Capreolus vulgaris Fitzinger, Wissensch.-pop. Naturgesch. der

Saugethiere, iv, p. 192 (Renaming of capreolus).
1874. Capreolus vulgaris, niger Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, IMath.-Naturwiss. Classe, lxx, Abth. i, p. 247

(Germany).
1874. Capreolus vulgaris, varius Fitzinger, Sitzungsber. kais. Akad.

Wissensch. Wien, Math.-Naturwiss. Classe, lxx, Abth. i, p. 247

(Germany).
1910. Capreolus capreolus Trouessart, Faune Mamm. d'Europe, p. 2.33.

Ti/pe localifi/. — Sweden.

Gt'ogru'phical disfrihutiov. — Originally confined to southei'n
Sweden but now somewhat extended artificially to the noilh
and west.

Diagnosis. — Throat patch indistinct ; general colour in winter
pelage grey decidedly tinged with yellow.

Measurements. — For cranial measurements .see Table, p. 983.

<5st.

France.

2 <5, 1 9.

Perri^res,

Seine-

Hon. N. 0. Roths-

10. 11. 17. 1.*

et-Marue.

child (p).

11. 11. 18. 1.*

12. 1. 17. 1.*

S.

Armandvilliers,

Hon. N. C. Roths-

11. 12. 5. 1.*

Seine-et-Marne.

child (p).

9.

Skabersjo, Sweden.

Dr. Einar Li'mn-

10. 10. 18. 1.

i'. st. (melauo),

?, 1 separate

skull.

Westphalia.

berg (p).
Purchased

(Gerrard).

7G. 5. 4. 1-3.*

3 skulls.

S. Germany

Dr.A. Gunther(c).

59. 9. C. 107-109

15 pairs of

Bavaria.

F. N. A. Fleisch-

11. 9. 13. 1-15.*

antlers.

mann (p).

i head st.

Austria.

Col. R. W. Shipway

98. 10. 12. 1.*

2 9.

Csehtelek,

Bihar

Hon. Mrs. Charles

10. 12. 4. 1.*

Comitat,

Huu-

Rothschild (p).

11. 1. 14. 11.*

7 skulls.

gary.

Lidth de Jeude

G7. 4. 12. 22f

Collection.

231.*

Not determined by the author. — O. T,

CAPREOLDS

975

Caprholus capreolus transsylvanicus Matschie.

1907. Capreolus transsylvaninis Matschie, Das Wcidwerk in Wort und
Bild, svi, p. 224, March 15, 1907.

Type locality. — Bana, Roumania.

Geographical disfrihufion. — Eastern Europe, westward to the
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.).

BemarJcs. — 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 canus by the sharply defined white throat patch.

2 9 Padola, Cadore, Venetian Alps. Turin Museum (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 distribution. — 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 (5, 2 9. Pinares de Quintanar S. & N. Gonzalez (c). 8. 7. 7. 27-31.
de la Sierra, Burgos, (8. 7. 7. 28. T]jpe o\ subspecies.)

Spain.

Capreolus capreolus tiiotti Lonnberg.

1910. Capreolus capreolus tliotti Lonnberg, Ann. and Mag. Nat. Hist.,
8th F.er., vi, p. 297, September, 1910. Type in British Museum.

Type locality.— AvivliWj, Craig EUachie, Morayshire, Scot-
land*.

Geograpjlrical distrihution. — Great Britain.

* The locality of the type specimen was accidentally recorded as
Aberfeldy in the original description. — O. T.

976

UNGULATA

Diagnosis. — Differs from C. capreolus capreolus 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, 3.30; ear, 127. For cranial measurements see Table, p. 983.

6 head sfc. Nairn, Scotland. Earl Cawdor (p). 93. 1. 3. 1.

6, 9. Nairn. Earl Cawdor (p). 8. 8. 18. 1-2.

S skeleton. Nairn. Earl Cawdor (p). 85. 10. 6. 1.

9 skull. Nairn. Earl Cawdor (p). 85. 10. 6. 2.

i, 9. Arndilly, Craig Ella- W. S. Menzies (p). 8. 11. 22. 1-2.

chie, Morayshire. (8. 11. 22. 1. Type of subspecies.)

<?, 9 heads st. Poltalloch, Argyll- Col. E. D. Malcolm 7. 6. 6. 1-2.
shire. (p).

i, 9. Thornhill, Dum- H. S. Gladstone (p). 11. 2. 22. 1-2.

friesshire.

S St. Scotland. Earl of Derby (p). GO. k.

Frontlet and Scotland. Gen. Hardwicke (p). 088. d.

antlers.

2 frontlets Scotland. G88. a-b.
and antlers.

i St. Purchased (R. Ward). 97. 12. 11. 2.

i St. Whatcombe, Bland- -J. C. Mansel Pleydell 97. 8. 21. 1.

ford, Dorset. (p).

Genus ALCES Gray.

1821. Alces Gray, London Med. Repos., xv, p. .307, April 1, 1821.

1841. xilcelapMis Gloger, Hand- u. Hilfsbuch der Naturgesch, i, p. 143

(Substitute for Alecs). Not of Blaiuville, 1810.
1857. Alces Blasius, Siiugethiere Deutschlands, p. 434.
1902. Paralccs Allen, Bull. Amer. Mus. Nat. Hist., xvi, p. 100, July 1,

1902 (Substitute for Alces assumed to be a homonym of Alee

Blumenbach, 1799).

Ti/2)e species. — Cermis alces Linnajus.

Geographical distrihutlon. — Northern forested portions of lioth
hemispheres ; in the Old World west to Norway and south to
eastern Germany.

Characteis. — Telemetaearpalian deer of lai'gest size with
narrow, elongate hoofs, maxillai'y canines al^sent 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 luisal 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 Lounberg, Zool.
Anzeiger, xxviii, pp. 448-449, January 17, 1905).

97^

Fig. 208.

Alces dices. X 1.

3 R

978

UXGDLATA

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.

RemarJcs. — The genus Alces contains four currently recognized
species or geographical foi'ms, two in America and two in the
Old World. Only one occurs in Europe.

ALCES ALCES Linnaeus.

1758. [Cervus] alces Linnseus, Syst. Nat., i, lOth ed., p. 66.

1829. ? Alces eurcypxits Burnett, Quart. Journ. Sci. Lit. Art, 1829, 353

(nomen nudum).
1836. Alces machlis Ogilby, Proc. Zool. Soc. London, p. 135 (Renaming of

alces).

1842. Alces antiqucn-um Riippell, Museum Senckenbergianum, iii, p. 183

(Renaming of alces).

1843. Alces pahnatus Gray, List Spec. Mamm. Brit. jMus., p. 182 (Re-

naming of alces).
1857. Alces pahnatus Blasius, Saugethiere Deutschlands, p. 434.
1860. Alces jubata Fitzinger, Wissench.-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.

Geographical 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.

Fig. 20
Alces alces. Incisiform teeth. Nat. size.

Diagnosis. — General characters as in the genus Alces ; height
at shoulder about two meters ; general colour brown, lighter and

RANGIFER 979

approaching wood-brown on head, back, and legs, darker, almost
blackish on tail, throat, and underside of body.

Measurements. — Adult male from Stenkjter, Trondhjem,
Norway : head and body about 2900 ; tail, 95 ; hind foot, 855 ;
ear, 330 ; height at shoulder, 1900. For cranial measurements
see Table, p. 984.

Specimens examined. — Twelve, from the following localities : —
Norway: Stenkjser, 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 Famise 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, Vollstandigste Naturgesch. des In- und Auslands,

Saugeth., iii, p. 12 (Alternative for Tarandus).

Type species. — Cervus tarandus Linnseus.

Geographical distribution. — Northern forests and barren
grounds of both hemispheres ; in the Old World north to
Spitzbergen ; Novaya Zemlaya ?

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

9S0

UNGULATA

KEY TO THE EUROPEAN SPECIES OF RANGIFER.

Size small, upper length of skull less than

225 mm. (Spitzbergen) R. platyrhynchus, p. 985.

Size large, upper length of skull about 270-

300 mm.

Upper length of skuU 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,^. 980.

Upper length of skull in adult male about
300 mm. ; upper tooth-row about 85 mm.
(wooded portions of northern Fiim^land
and formerly of northern Sweden} R. fennicus, p. 981.

RANGIFER TARANDUS Linmeus.

1758. \Ce)'vus] tarandus Linnaeus, Syst. Nat., i, 10th ed., p. 67.

1789. [Cervus tarandus] a rangifer Gmelin, Syst. Nat., i, 13th ed., p. 177.

1827. Tarandus lapj}onu7n Billberg, Synopsis Faunae Scandinaviee, i, p. 22

(Renaming of C. tarandus).
1842. Tarandus borealis Rii^jpeU, Museum Senckenbergianum, iii, p, 183

(Renaming of C. tarandus).

Fio. 210.
Rangifer taramlu.t

RANGIFER 981

1852. Taranchis furcifer Baird, Kep. Coram. Patents, 1851, ii, Agric, p. lO'J

(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, Paune Mamm. d'Europe, p. 231.

Type locality. — Alpine region of Swedish Lapland.

Geographical diatribution. — 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 1 150 mm. ;
upper length of skull 270-290 mm.) ;
skull with 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. — Adult male from Heimdalen, Norway : head
and body, 2000±, tail, 150 ; hind foot, 450± ; ear, 100±. For
cranial measurements see Table, p. 984.

Specimens examined. — 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.

Beviarls. — The matei-ial examined is too limited to form the
basis of any discussion of the various forms of wild and tame
reindeer.

cisifonii teeth

(5 St. LoerdalMts.,Sogue Sir W. J. Ingram 87.9.20.1.

Fjord, Norway. (c & p).

J & ? antlers. Loerdal Mts. Sir W. J. Ingram 87. 9. 20. 2-3.

(c & p).

6 St. Fillefjeld, Norway. Chas. Ingram (c &p). 79. 10. 9. 1.
& skeleton.

9 St. FiUefjeld, Norway. Sir W. J. Ingram 81. 9. 28. 2.

& skeleton. (c & p).

?. Norway. Chas. Ingram (c & p). 75.10.30.1.

? (ad. & yg.). Norway. J. C. Ingram (c & p). 83. 7. 28. 1-2.

RANGIFER FENNICUS Lonnberg.

1909. Rangifer tarandus fcnnicus Lonnberg, Arkiv for Zoologi, vi, No. 4,
p. 10, July 14, 1909.

Ty2)e locality. — Torne, Lappmark (" thus probably in Enon-
tekis," Jide Lonnberg), Finnland.

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RANGIFER 985

GecKjrapMcal 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 Bangifer tarandas (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-rovv^ about 85 mm. ;
lower tooth-row about 90 mm.).

Measure merits . — For cranial measurements .see Table opposite.

Bemarlcs. — -This animal, which I know only from Lonnberg's
descrijjtion and figures, appears to be specifically distinct from
Bangifer tarandus.

RANGIFER PLATYRHYNCHUS Vrolik.

1829. Cervus (Tarandus) platyrhynchus Vrolik, Nieuwe Verhandel. van 't

Kroninke Nederl. Inst., Eerste Klasse, pt. 2, p. 160 (Spitzbergen,

see Vrolik, I.e., p. 239).
1862. Cervus tarandus forma spetshcrgcnsis Anderson, Ofversigt af Kongl.

Vetensk.-Akad. Porhandl., Stockholm, xix, p. 457, October 8, 1862.
1866. \_Eayigifcr arctictis} var. spitzbergcnsis Murray, Geogr. Distrib.

Mamm., p. 334, described on p. 155.
1902. Bangifer spitzbergcnsis Camerano, Mem. Reale Accad. Sci. Torino,

2d ser., li, pp. 159-240, 1902.
1910. Bangifer spitzbergcnsis Trouessart, Faune Mamm. d'Europe, p. 232.

Type locality. — Spitzbergen.

Geographical distribution. — Spitzbergen.

Diagnosis. — Size much less than in Bangifer tarandus and
B. fennicvs (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 B. tarandus.

Measurements. — ^For cranial measurements see Table opposite.

Specimens examined. — Two mounted individuals and six skulls
(U.S.N.M. and Turin).

Beviarlcs. — The Spitzbergen Reindeer is sharply and com-
pletely differentiated from both of the species occurring in
continental Europe.

DbG DNGULATA

Family BOVID^.

1821. Bovidn- Gray, London Med. Rcpos., xv, p. 308, April 1, 1821.

Geof/rajjlu'cal distrihution. — 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
occui'ring in Loth sexes, always, when present, in the form of
permanent horns supported by bony cores arising from the frontal
bones ; molars usually (always in European members of the
family) hypsodont ; lateral digits rudimentary or absent, usually
I'ei^resented by the hoofs alone.

licmarlcs. — 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, Ocis, Capra and
Hupicapra, now occur naturally in western Europe.*

Genus OVIS Linnseus.

1758. Ovis Linnaeus, 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 (il/. asiaticus).

1798. Musmon Schrank, Fauna Boica, i, p. 81 (Substitute for Ovis).

1795. Aries Link, Zool. Beytrage, i, pt. ii, p. 96 (part).

181G. Amnion Blainville, Bull. Soc. Philomathique, Paris, p. 76, Jlay, 1816

(Alternative name for Ovis).
1857. Ovis Blasius, Siiugethiere Deutschlands, p. 466.

Ti/jic species. — Oris aries Linnpeus (by tautonymy).

Geofjrajihical 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 Mediteri-anean portion of Europe.

Characters.— Bovidte 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
genei'al foi'm ; lachrymal pit present ; nasal branch of premaxillary
not wedged between nasal and maxillary ; horns aKvays present
in males, usually absent or rudimentary in females, very robust

* The genera Bos and Bison have become extinct within historic times ;
a species of Bubalus, introduced in the 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 an
outwardly directed spiral ; teeth strongly hypsodont, the upper
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 ^J when unworn not more than height.

BemarJcs. — The genus Ovis contains about forty described
sj^ecies, mostly Asiatic. Only one is definitely known to occur
wild in Europe.

OVIS MUSIMON Pallas.

1811. JEgoccros musinion Pallas, Zoogr. Rosso-Asiat., i, p. 230 (Sardinia).
1829. [Ovis nmsiinon] b var. occidentalis Braudt and Ratzeburg, Getreue

Darstellimg und Beschreibung der Thiere, p. 55 (Corsica).
1841. Ovis musmon Bonaparte, Iconogr. Faun. Ital., Indice distrib.
1857. Ovis musimon Blasius, Saugcthiere Deutschlands, p. 471.
1905. ? Ovis matschiei Duerst, Martin Wilokens Crrundziige der Naturgescb.

der Haustiere, 2nd ed., p. 180 (Corsica and Sardinia).
1910. Ovis musimon Trouessart, Fauna Mamm. d'Europe, p. 242.

Tjipe locality. — Sardinia.

Geographical distribution. — Corsica and Sardinia.

Diagnosis. — Size decidedly less than in ordinary domestic
sheep (head and body of rams about 1 300 ; 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 separated from that of belly by a black stiipe
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.

Sixcimens examined. — 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 Lydekker, Field, London, ex, pp. 147, 197, and Bernard, ibid.,
p. 147.

988

UNGULATA

Ocis musimon : foxy red with brown stripe on back, white
spots (Flecken) on head and flanks ; horns light coloured, curved
in a single plane.

Ovis matschiei : more brownish grey in general colour, the
face ash-grey rarely marked with white ; horns dark brown,
curved in a sjjiral 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.

<5st.
S, 9.

yg-

6 St.
2?.

S skeleton.

5 skull.

6 horns.

Gennargentu Mts.,

Sardinia.
Gennargentu Mts.

(P. Bonovii.)
Sardinia.

(W. Ewer.
Corsica.

Ford G. Barclay 95.4.4.1.

(c & P).
E. N. Buxton (p). 95. 4. 16. 6-7.

A. B. Bay ley- 5.3.6.2.

Worthingtou (c & p).
Zoological Society.
Zoological Society.

53. 8. 29. 49.

61. 3. 24. 2.

62. 12. 22. 2.
60. 4. 23. 1.
55. 12. 26. 162.

Zoological Society.
Zoological Society.
Dr. C.I. Forsyth Major 9. 1. 11. 1.

(c).

Genus CAPRA Linnteus.

1758. Ca-pra Linnaeus, Syst. Nat., i, 10th ed., p. 68 {hircus, by tautonymy).

1762. Hircus Brisson, Regn. Anim. in Classis ix distrib., 2nd ed., p. 12

[Hircus Brisson = Capra hircus Linnaeus).

1766. Ibex Pallas, Spicil. Zoologica, ii, fasc. 11, p. 81 (sibiricus).

1795. Aries Link, Zool. Beytriige, i, pt. 2, p. 96 (Substitute for Capra).

1798. Tragus Schrank, Fauna Boica, i, p. 80 (Substitute for Capra).

1811. JEgoceros Pallas, Zoogr. Rosso-Asiat., i, p. 224.

1857. Capra Blasius, Siiugethiere Deutschlauds, p. 474.

Type species. — Capra hircus Linnaeus.

Geo(jra2)Mcal (listrihution. — 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.

Cltaracters. — Like Ovis externally but hind feet without
glands, males bearded, and horns ne\-er 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.

BemarJcs. — About thirty-five sjiecies are currently referred to
the genus Capra. Half a dozen of those occur in the Mediter-
ranean region of Europe.

989

CAPRA IBEX Linnpeus.

1758. [Ca2)ra] ibex Linnaeus, Syst. Nat., i, lOtb ed., p. 68.

1786. Capra alpina Girtauner, Observ. et M6m. sur la Phys., I'Hist. Nat.

et les Arts, xxviii, p. 224, March, 1786.
1857. Capra ibex Blasius, Siiugethiere Deutschlands, p. 475.
1906. Ilhex} ibex Camerano, Mem. Accad. Reale Sci. Nat., Torino, 1905-

1906, p. 284.
1910. Capra ibex Trouessart, Faune ]\Iamm. d'Europe, p. 230.
1912. Capra ibex graicus Matschie, Deutsche Jagu. Zeitung, lix, no. 8,
April, 1912.

Type locality. — Swiss Alps.

Geographical 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).

i st. Italian side of Monte Purchased (Rowland 97. 12. 11. 1.

Rosa. Ward).

i St. Alps. Purchased (Parreys). 41. 596.

6 St. Alps. Mrs. A. G. Campbell 650. 2.

(P)-

• S skull. Alps. Messrs. Murray and 18.35.

Brockedon (p).

c5 horns. Chamonix. Purchased (Faery). 45. 12. 16. 1.

2 S horns. 650. a. b.

? skull. Purchased (Warwick). 57. 3. 18. 2.

CAPRA PYRENAICA Schinz.

(Synonymy under subspecies.)

Geographical disfrihution. — 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, outwai'd and backward,
distinctly compressed laterally, the antei'ior surface without
evident transverse ridges, the antero-external border raised and
rib-like ; colour paler than in Capra ibex, and dark areas sharply
defined. f

Remarks. — Four local races of Capra pyrenaica 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 Socage.

Capra pyrenaica pyrenaica Schinz.

1838. Capra fiyrcnaica 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. Capi-a pyrenaica 2}yrenaica Cabrera, Proc. Zool. Soc, London, p. 966,

December, 1911.

Type locality. — Spanish Pyrenees.

Geographical distribution. — Formerly the Pyrenees and ea-stern
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.

c5 St. & 9 skull. Pyrenees. Purchased (Boubee). 48. 2. 5. 4-5.

? head St. Spain. Capt. J. Marriott (c & p). 0.10.28.1.

<5 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.

+ " The species, 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 pelage 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, Loudon, 1911, p. 967, December, 1911).

991

Capra pyrenaica lusitanica Franca.

190.9. Capra lusitanica Francja, 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, xjp. 1-20, xjIs. 1-2. Articles separately paged. Bocage
wrongly cited as authority for the name lusitanica. Type in
Lisbon Museum.

1911. Capra pyrenaica, peculiar subspecies, Cabrera, Proc. Zool. Soc,
London, p. 966, December, 1911.

Type locality. — 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 victoria 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. pyrenaica and C. p. liispanica,
rather browner than Mspanica 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. Capra hispanica Schimper, Comptes-Rend. Acad. Sci., Paris, xsvi,
p. 318, March, 1848.

1910. Capra pyrenaica hispanica Trouessart, Fauno Mamm. d'Europe,

p. 237.

1911. Capra pyrenaica hispanica Cabrera, Proc. Zool. Soc, London, p. 96C,

December, 1911.

Type locality. — Sierra Nevada (" Picacho de Yeleta et ]Mula-
hacen "), Spain.

Geographical distribution. — -Mountains of southern and eastern
Spain (Sierra de Ronda, Sierra Nevada, Sierra Morena, Sierra
Martes and Sierra de Cardo).

Diagnosis. — Dark markings minimum for the species, the
latei'al 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 Cabrei'a) : head and body,
1190 ; hind foot with hoofs, 305.

6 st. S. Spain. Purchased (Parzudaki). 55. 11. 26. 14.

CAPRA iEGAGRUS Erxleben.

1777. [Cap^-a] ifgagrus Erxleben, Syst. Regni Anim., i, p. 260 (Caucasus).

1857. Capra xgagrus Blasius, Saugethiere Deutschlands, p. 485.

1858. ? JBgocerus pictus Erhard, Fauna der Cycladen, Wirbelthiere, p. 32

(Syra, Cyclades, Greece).
1888. ? Capra dorcas Reiclienow, Zool. Jahrb., Syst., iii, p. 594, May 31,

1888. Island of Joura (Gyaros), Greece.
1899. ? Clapi-a] wgagrus var. jourcnsis Ivrea, Proc. Zool. Soc, London,

X3. 599. Nomen nudum.
1399. ? Capra cretensis Lorenz-Liburnau, Wissensch. Mitth. aus Bosnien

und der Hercegovina, vi, p. 865 (Crete).
1910. Capra .rgagrus picta, C. xgagrus dorcas and C. ngagrus cretensis

Trouessart, Faune Mamm. d'Europe, pp. 238-239.

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. Rupicapra Blainville, Bull. Soc. Philomathique, Paris, p. 75, May,
1816.

1840. Capella Keyserling and Blasius, Wirbelthiere Europas, p. iv.

1841. Cemas Gloger, Gomeinn. Hand- u. Hilfsbuch der Naturgesch., I, p. 153

(not of Oken, 1816).
1857. Capella Blasius, Siiugethierc Deutschlands, p. 487.

Type species. — Capra rupicapra Linnasus.

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 j)osteriorly ; lachrymal pit absent ; nasal
branch of premaxillary not in contact witli 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.

Remarks. — Four members of the genus Rupicapra are now
recognized, a number which will probably be increased when
adecjuate material is bi-ought together.

RUPICAPEA

993

RUPICAPRA RUPICAPRA LiliniBUS.

1758. [Capra] rupicapra Linuaeus, Syst. Nat., i, 10th ed., p. G8 (Switzerland).
1829. ? Rupicapra hamulicornis Burnett, Quart. Joum. Sci. Lit. Art.,

1829, pt. II, p. 353 (noinen nudum).
1843. Rupicapra tragus Gray, List Spec. Mamm. Brit. Mus., p. 167

(Renaming of rupicapra).
1845. Rlupicainri] capella Bonaparte, Atti della sesta Riunione degli

Scienziati Italiani, Milano, 1844, p. 337 (Renaming of rupicapra).
1847. \_Capra rupicapra^ a sylvatica Sundevall, Kongl. Vetensk.-Akad.

Handl., Stockliolm, 1845, p. 284 (Wooded portions of Alps).
1847. [Capra rupicapra'] fi alpina Sundevall, Kongl. Vetensk.-Akad.

Handl., Stockholm, 1845, p. 284 (Higher portions of Alps).
1857. Capella rupicapra Blasius, Siiugethiere Deutschlands, p. 488.
1870. Rupicapra europea Cornalia, Fauna d'ltalia, pt. i, p. 53 (Substitute

for rupicapra ; " Cuv." cited as authority).
1897. [Rupicapra'] dorcas Schulze, Abhandl. u. Vortrage Gesammtb. Natur-

wisseusch., iv. No. 10, p. 9 (Substitute for rupicain-a).
1910. Rupicapra rupicapra Trouessart, Paune Mamm. d'Europe, p. 235.

Type locality. — Switzerland.

GeograpMcal 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,
Xi.

Rupicapra rupicapra.
Incisiform teeth (iiat. 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

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 fi-ontal between them.

Measurements. — Adult male (mounted) : head and Vjody, 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.) ; Transylvania;
S. Carpathians, 2.
No History : Five.

6 St.

Tyrol.

H.R.H. Crown Prince SI. 9. 21. 1.
Rudolph of Austria

ad. and yg.

Tyrol.

Prince Rudolph of

78. 6. 20. 1-2.

skeletons.

Austria (p).

2 6 skulls.

Alps.

Gen. Hardwicke (p).

6:31. a. b.

3 frontlets

Alps.

Dr. J. E. Gray (p).

46. 10. 13. 31-38

and horns.

6 skull.

Alps.

J. Gould (p).

46. 7. 7. 2.

6 skull.

Alps.

Dr. A. Giiuther (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.

Trausyl-

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. ilus. Civ. Stor. Nat. Geneva,

XX, p. 347, December 20, 1899. Type in Genoa Museum.
1910. Bupicapra oriiata Trouessart, Fauue ilamm. d'Europe, p. 235.

Type locality. — Barrea, near Alfedena, Province of Aquila,
Italy.

Oeographical distribution. — Abruzzi Mountains, Italy.

Diagnosis. — General colour as in Bupicapra rupicapra, 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 jB. rupjicap)ra, and area of ujjjier 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 type locality (B.M. and
Genoa).

6. Abruzzi Mts., Italy.

Genoa Museum (e).

4. 2. 29. 1.

RUPICAPRA 995

RUPICAPRA PYRENAICA Bonaparte.

1845. B[upicapra] pyrcnaica Bonaparte, Atti della sesta Riuuiono degli

Scienziati italiani, Milauo, 18i4, p. 337.
1878. Antilopc rupicapra var. Riitimeyer in Trutat, Bull. Soc. Hist. Nat.

Toulouse, XII, p. 117.
1910. Rupicapra rupicapra Trouessart, Faune Mamm. d'Europe, p. 235

(Part ; in synonymy only).
1910. Rupicapra rupicapra pyrenaica Cabrera, Proc. Zool. Soc. London,

p. 998. December, 1910.

Type localiiy. — Pyrenees.

Geographical distrihution. — Pyrenees.

Diagnosis. — Colour of winter pelage distinctly less dark than
in Rupicapra rupicapra ; markings on neck essentially as in
R. ornata ; horns about perpendicular, placed more closely
together than in the other species, the very narrow 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.).

Remarks. — This very 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 R. parva), it appears to have been completely
overlooked."}"

1 st. Pyrenees. Purchased (Boub6e). 48. 2. 5. 7.

RUPICAPRA PARVA Cabrera.

1910. Rupicapra rupicapra parva Cabrera, Proo. Zool. Soc. London, p. 999,
December, 1910. Type in Madrid Museum.

Type locality. — Picos de Europa, Santander, Spain.

Geographical distrihution. — Cantabrian Mountains, Spain.

Diagnosis. — Described as smaller and darker than Rupicapra
pyrenaica, and with throat-patch very ill-defined, scarcely paler
than general colour of body.

Measurements. — Type (adult female) : height at shoulder,
570 mm. ; length of horn along anterior curve, 146 (Cabrera).

6 St. Picos de Europa, Cantabrian J. W. E. Lee (c & p). 97. 1. 7. 8.
Alps.

* Bull. Soc. Hist. Nat. Toulouse, xii, pp. 115-117.

t See Lydekker, Great Game of Europe, Asia and America, p. 185, 1901.
Trouessart places the name pyrenaica in the synonymy of rupicapra.

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998 APPENDIX

APPENDIX.

Aflditional material shows the distinctness of the Sardinian
(7roc?'fZMra mentioned on pp. 101 and 111-112. The animal has
been described as : —

CROCIDURA ICHNUS.ffi Festa.

1912. Crocidura ichmis.f Festa, Boll. Mus. Zool. Anat. Comp., Torino,
xxvir, No. 648, p. 1, March 9, 1912. Type in Turin Museum.

Ti/2)e locality. — Piscina, Lanusei, Sardinia.

Geograjjkicnl distribution. — Sardinia.

Characters. — Like Crocidnra sicuJa, but unicuspid teeth both
above and below full}' as large as in C. russula, and third upper
unicuspid more conspicuou.sly exceeding second than in either of
the related species. Brain-case apparently as deep in proportion
to its width as in C. 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 (ll-8-12-2j. 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; tvpe) ; Ovile Seardu, 3 (Genoa); Zinnigas, 2
(Genoa) ; Assuni, 1 (U.S.N.:\I.).

The cranial measurements of Evotomys glareolus norvegicns
were accidentally omitted from Table, pp. 652-653, and are as
follows : —

999

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1000 APPENDIX

Genus Spalax (pp. 887-896). Tlie living forms recognized
by Mehely (Species Generis Spalax. A Foldi Kutyak Fajai
Szarmazas- es Rendszertani Tekintetben, Budapest, 1909) as
occurring in Europe west of Russia, are as follows :* —

Sub-genus MesosjMlax (p. 22). New sub-genus to contain the
species monticola and hungaricus. No type designated.

Spalax monticola monticola Nehring. Type locality : Kupres,
Bosnia.l (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.

Sjjalax 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.^ (T3^e in Agricultural
High School, Berlin.)

Spalax monticola ayrmiensis Mehely (p. 30). Type locality :
0-Pazua, Szerem, Slavonia, Austria-Hungary. (Type in Agram
Museum.)

Spalax monticola i^crhirus Mehely (p. 31). Type locality :
Servia. (Type in Hungarian National Museum.)

Spalax hungaricus hungaricus Nehring.

Spalax hungaricus transsylvamcus Mehely (p. 31). Type
locality : Transylvania. (Type in Hungarian National Museum.)

Sub-genus Macrospalax (p. 23). New name for sub-genus
Spalax.

Spalax gnrcus grsccus 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 polonictis Mehely (p. 35). Type locality : vicinity of
Lemberg, Galicia, Austria-Hungary. (Type in Lemberg Museum.)
As synonyms are cited : Podolian Marmot Pennant, Sj'nopsis,
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 (h) from Herzegovina. Neither is desig-
nated as type. As a is young and h adult, and Nehring expressly notes
that the characters of the species are particularly well shown by b, I
assumed (p. 894) that b should be regarded as the type. Mehely states
(p. 121) that a is the type of monticola. He bases his description of
hercegovinetisis on Nehring's specimen b exclusively.

INDEX

[Principal references are indicated by the use of heavy type.]

abraraus, Pipistrellus, 213
abramus, Vesperugo, 213
Acanthomys, 883
acervator, Mus, 878
acervifex, Mus, 878
Achlis, 979
Acomys, 883

,, minous, 883
Acosmiiithus, 883
Acromys, 863
Adelonj^cteris, 225
sedilis, Vespertilio, 184
Eegagrus, Capra, 992
^goceros, 988
Aeorestes, 167

sestiva, Mustela erminea, 389
afra, Geuetta, 451
africana, Mustela, 412
africanus, Putorius, 412
africauus, Putorius uivalis, 412
agilis, Miororays, 841
agilis, IMus (Apodemus) minutus, 844
agilis, Mus miuutus, 844
agilis, Vespertilio, 220
agrarius, Apodemus, 836
agrarius, JIus, 836
agrarius, IMus (Apodemus), 836
agrestis, Arvicola, 668, 671, 672, 673,

675
agrestis, INIicrotus, 662, 668, 671, 683
agrestis, Microtus agrestis, 668
agrestis, ]\Ius, 668
Agricola, 659
agrius, Felis, 470
agrius, Felis ocreata, 470
alba, Arctomys marmota, 932
alba, Canis vulpes, 331
alba, Hystrix cristata, 543
alba, Lepus timidus, 526
alba. Lynx vulgaris, 471
alba, Mustela foina, 375
alba, Mustela lutreola, 415
alba, Talpa europjea, 4
albicans, Mus musoulus, 869
albicans, Vesperugo kuhlii, 216
albicus. Castor, 948

albicus, Cervus, 966

albida, Talpa europjea, 4

albifrons, Cervus elapbus, 965

albigularis, Vesperus (Marsipolffinius),
238

albipes, Sorex, 88

albirostris, Mus agrarius, 836

albiventris, Crocidura aranea, 101

albiventris, Sorex fodiens, 70

albolimbatus, Vespertilio, 215

albo-maculata, Talpa europsea, 4

albo-notatus, Sciurus vulgaris, 905

albus. Castor fiber, 947

albus, Cervus 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, 428

albus, Rattus domesticus, 858

albus, Sciurus vulgaris, 905

albus, Sorex fodiens, 69

albus, Ursus arctos, 286

Aloelapbus, 976

Alces, 976

„ alces, 978

alces, Alces, 978

alces, Cervus, 978

alcythoe, Vespertilio, 215

alexandrino rattus, Mus, 853

alexandrlnus, Epimys rattus, 854

alexandrinus, Mus, 854

alexandrlnus, Mus (Epimys) rattus,
854

algidus, Lepus, 526

algirus, Erinaceus, 130

algirus, Erinaceus algirus, 131

Alopex, 318

,, lagopus, 319

,, spitzbergenensis, 324

alopex, Canis, 330

1002

alpina, Capra, 989

alpina, Capra rupicapra, 993

alpina, Marmota, 932

alpina, Sciurus vulgaris, 910

alpinns, Arvicola, 716

alpinus, Lepus timidus, 526

alpinus, Myofcis murinus, 193

alpinus, Putorius, 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

Ainblyotus, 225

americana, Arvicola, 730

Amniomys, 752

Ammon, 986

amori, Eliomys, 560

amori, Mioxus nitela, 560

ampliibius, Arvicola, 725, 730, 738,
744-740, 749

amphibius, Arvicola amphibius, 730

amphibius, Mus, 730

amphibius, Sorex, 70

Amphisorex, 29, 65

anglicus, Muscardinus avellanarius,
584

angiilaris, Microtus, 706

angustifrons, Arvicola agrestis, 671

angustifrons, Lutra, 356

Animalivora, 134

annulatus, Ursus arctos, 286

anomalus, Neomys, 81

anomalus, Neomys fodiens, 81

Anotis, 887

antinorii, Sorex, 02

antipfB, Crocidura, 95

antipai, Crocidura, 95

anfciquorum, Alces, 978

Aper, 956

aper, Sus setosus, 957

Aphrontis, 898

Apodemus, 791

,, agrarius, 836
,, cpimelas, 794

flavicollis, 828
„ „ flavicollis, 829

,, ,, wintoni, 831

,, fridariensis, 825

,, hebridensis, 824

,, hirtensis, 825

,, sylvaticua, 797

,, ,, callipides, 809

,, ,, creticus, 813

„ ,, dicbrurus, 810

,, ,. sylvaticus, 803

aquatica, Arvicola, 730

aquaticus, Hypudasus paludosus, 738

aquaticus, Lemmus, 730

aquaticus, Sorex, 69

Aquias, 137

aquilonius, Lepus, 508

aquilonius, Lepus medius, 508

aquitanius, Microtus (Ghionomys) ni-
valis, 717

aquitanius, Microtus nivalis, 717

araneus, Crocidura, 101

araneus, Sorex, 31, 101

araneus, Sorex araneus, 35

arcalus, Males, 352

arctica, Vulpes, 319

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, IMus, 846

arvalis, Arvicola, 683

arvalis, Lemmus, 032, 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, 733

„ ,, tenebricus, 735

,, scherman, 744

,, ,, exitus, 746

„ ,, monticola, 749

,, ,, scherman, 745

,, terrestris, 738

Aspalax, 887

assimilis, Arvicola arvalis, 683

asturianus, Microtus, 693

ater, Arvicola arvalis, 083

ator, Ilypudanis terrestris, 738

ater, Mus (Kpimys) rattus, 863

ater, Mus rattus, 853

ater, Rattus domesticus, 853

atlanticus, Cervus elaphus, 907

atlanticus, Euryalus, 155

1003

atlanticus, Rhinolophus euryale, 155
atratus, Amblyotus, 231
atticus, Crioetulus, 593
atfcila, Sus, 960

aureolus, Mustela putorius, 495
aureolus, 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, Mus, 583
avellanarius, Muscardinus, 583
avellanarius, Myoxus, 583
avellanus, INIyosus, 577
avenarius, Mus, 844
azoreura, Nyctalus, 254
azoreum, Pterygistes, 254
azoricus, Mus, 871
azoricus, Mus musculus, 871

badius, Ursus, 286
basticus, Sciurus, 923
bffiticus, Sciurus vulgaris, 923
bseticus, Sus scrofa, 960
baibac, Arctomys, 937
bailloni, Arvicola, 672
bailloni, Microtiis 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
barbastelle, Vespertilio, 263
Barbastellus, 263
barbastellus, Barbastella, 263
barbastellus, Synotus, 263
barbastellus, Vespertilio, 263
bechsteinii, Myotis, 179
bechsteinii, Vespertilio, 179
bergensis, Sorex araneus, 41
bicolor, Myodes, 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, IMus, 937
bocoamela, INIustela, 405
boccamela, INIustela nivalis, 405
bocoamela, Putorius (Ictis), 405
boccamela, Putorius nivalis, 405
bolivari, Cervus elaphus, 969

bonapartii, Vespertilio, 219

borealis, Pelis, 471

borealis, Gulo, 434

borealis, Lenimus, 615

borealis, Lepus, 526

borealis, Tarandus, 980

borealis, Vespertilio, 234

borealis, Vesperugo, 234

boscai, Eptesicus, 226

boscai, Vespertilio, 226

Bovidfe, 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
I brunneus, Pitymys pyrenaicus, 772

brunneus, Ursus arctos, 286

budapestiensis, Myotis ciliata, 177

buffonii, Lemmus arvalis, 745

bunites, Martes, 380

bunites, Mustela foina, 380

cabrerfe, Euryalus, 155

cabreree, Microtus, 701, 703

cabrerai, ]\Iicrotus, 701

cabrerai, Rhinolophus euryale, 155

cajca, Talpa, 15

caarulea, Canis vulpes, 319

caesarius, Evotomys, 645

calcarata, Romicia, 215

callipides, Apodemus sylvaticus, 809

callipides, ]\Iicromys sylvaticus, 809

callipides, IMus sylvaticus, 809

campestris, Arvicola, 683

campestris, Cervus vulgaris, 966

campestris, Mus, 844

campestris, Mus (Apodemus) minu-

tus, 844
campestris, jSIus minutus, 844
Campicola, 659
campicola, Lepus, 502
canaliculatus, 55
candidus, Mus sylvaticus, 803
candidus. Sorex araneus, 101
caneoe, Crocidura, 109
canescens, Cricetus cricetus, 603
canescens, Cricetus vulgaris, 603
canescens, Lepus, 526
caniceps, Erinaceus, 120
canicularius, ]Mus, 878
caniculator, Mus, 878
Cauidte, 303

caninus, Erinaceus, 120
caninus, Meles communis, 348

1004

Canis, 304

„ aureus, 315

„ lupus, 305

,, ,, dietanus, 315

„ lupus, 313

,, ,, signatus, 314

cantabra, Crocidura, 99
cantabra, Crocidura mimula, 99
eanus, Canis lupus, 313
canus, Capreolus capreolus, 975
canus, Mus ampliibius, 745
capaccinii, IMyotis, 187
capaccinii, Myotis (Leuconoe), 187
capaccinii, Vespertilio, 187
Capaccinius, 166
Capella, 992
capella, Rupicapra, 993
Capra, 988

,, fegagrus, 992

„ ibex, 989

,, pyrenaica, 989

,, „ hispanica, 991

,, ,, lusitanica, 991

,, ,, pyrenaica, 990

,, ,, Victoria, 991

capraea, Capreolus, 974
Caprea, 972
Capreolus, 972

,, capreolus, 973
,, ,, canus, 975

„ ,, capreolus, 974

„ „ thotti, 975

,, ,, transsylvanicus,

975
capreolus, Capreolus, 973, 974
capreolus, Capreolus capreolus, 974
capreolus, Cervus, 974
Caprios, 21

capucinellus, Vespertilio, 184
capucinus, Pitymys subterraneus, 760
carinatus, Sorex, 69
Carnivora, 283

carpathicus, Sorex araneus, 43
carpetanus, Rhinoloplius, 159
caspius, ilus, 858
castaneus, Arvicola terrestris, 746
castaneus, Sorex araneus, 37
castaneus, Sorex tetragonurus, 37
castilianus, Sus scrofa, 960
Castor, 947

„ fiber, 947
Castorida;, 947
Cateorus, 225
Catolynx, 456
Catus, 456
catus, Felis, 462
caudata, Crocidura, 110
caudata, Talpa, 3
cellarius, Mus, 829 ,
celtica, Sus scrofa, 957
celticus, !Mus sylvaticus, 803
Cemas, 992

centralis, Pitymys ibericus, 782

Cervaria, 471

cervaria, Felis, 472

Cervidffi, 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

Chionobates, 495

Chionomys, 712

Chiroptera, 134

chrysothorax, Sores, 101

ciliatus, Neomys fodiens, 73

ciliatus, Sorex, 73

ciliatus, Vespertilio, 177

cincticauda, Eliomys, 559

cinerea, Canis vulpes, 331

cinerea, Sciurus vulgaris, 910

cinerea, Talpa europcea, 4

cinereo-maculatus, Mus musculus,869

cinereus, Lepus timidus, 502

cinereus, Sciurus vulgaris, 906

cinereus, Sorex araneus, 101

cintrifi, Crocidura russula, 108

Citellus, 924

citellus, 924
,, suslica, 929

citellus, Citellus, 924

citellus, Citellus (Citellus), 924

citellus, Mus, 924

citellus, Spermophilus, 924

CitiUus, 924

citillus, Mus, 924

clivosus, Rhinolophus, 162

Cnephffius, 224

cnossius, Oryctolagus cuniculus, 491

Coelophyllus, 137

coUaris, Sorex, 69

collaris, Vespertilio, 169

coUinus, Lepus borealis, 526

collinus, Lepus timidus, 527

Comastes, 167

communis, BarbasteUus, 263

communis, Genetta, 451

communis, Meles, 348

communis, Plecotus, 256

communis, Vulpes, 330

concinnus, Sorex, 35

consolei, 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, 969
corsicanus, Cervus elaphus, 'JG'J
corsicanus, Lepus, 507
corsicanus, Lepus europseus, 507
corsicanus, Lepus mediterraneus, 507
corsicanus, Putorius nivalis, 412
Corsira, 29
Craseomys, 624

crassicaudatus, Sorex vulgaris, 43
crassidens, Lemmus lemmus, 621
cretensis, Capra, 992
cretensis, Capra isgagrus, 992
creticus, Apodemus sylvaticus, 813
creticus, Lepus, 512
creticus, Lepus europseus, 512
CricetinEe, 592
Cricetulus, 593

,, atticus, 593

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

,, caneee, 109

,, caudata, 110

,, cyrnensis, 111

,, ichnusfe, 998

,, leucodon, 88

,, mimula, 94

„ ,, cantabra, 99

„ ,, iculisma, 98

,, ,, miniula, 95

,, russula, 99

„ ,, cintrse, 108

,, ,, pulchra, 103

,, „ russula, 101

,, sicula, 108

Crossopus, 65
crucigera, Cauis, 331
crucigera, Vulpes vulpes, 331
cunioularius, Arvicola, 683
Cuniculus, 484
cuniculus, Lepus, 490
cuniculus, Oryctolagus, 485, 490
cuniculus, Oryctolagus cuniculus, 490
cylindricornis, Rangifer 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

danubicus, Erinaceus, 127

danubicus, Erinaceus europa;us, 127

dasycarpos, Vespertilio, 252

dasycneme, Myotis, 189

dasycneme, Vespertilio, 189

dasj'pus, Vespertilio, 187

daubentonii, Barbastellus, 263

daubeutonii, 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

Desmaninse, 20

Desmanus, 21

destructor, Arvicola, 744

dichrurus, Musculus, 810

dicbrurus, Apodemus sylvaticus, 810

dichrurus, Mus sylvaticus, 810

Dieba, 304

dietanus, Canis lupus, 315

discolor, Vespertilio, 238

discolor, Vesperugo, 238

dolbrogese, Spalax, 889

dolbrogese, Si^alax monticola, 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

druentius, Pitymys, 762

dryas, Eliomys, 570

dryas, Myoxus, 568

Drymomvs, 863

Dryomys", 566, 863

duodecimcostatus, Arvicola, 784

duodecimcostatus, Microtus, 784

duodecimcostatus, Pitymys, 784

duplicata, Arvicola arvalis, 686

duplicatus, Microtus arvalis, 686

Duplicidentata, 483

Dyromys, 566

,, nitedula, 567
,, ,, intermedins, 669

,, ,, 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
„ querciuus, 551
„ sardus, 560

Elius, 572

emargiuatus, 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

ereniita, Sorex, 69

Erinaceidse, 114

Erinaceus, 114

algirus, 130, 131
,, ,, algirus, 131

,, ,, vagans, 133

,, europeeus, 115

„ ,, consolei, 126

,, ,, europteus, 120

,, ,, hispanicus, 122

,, ,, italicus, 123

,, nesiotes, 129
,, roumanicus, 127

erinaceus, Hystris, 120

erminea, Pcetorius, 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

euronotus, Sorex araneus, 41

europsea, Hystrix cristata, 543

europfea, Lutreola, 415

europsea, Talpa, 3

europeeus, Alces, 978

europseus, Canis Vulpes alopex, 331

europseus, Capreolus, 974

europeeus, Erinaceus, 115, 120, 123

europseus, Erinaceus europtpus, 120

europseus, Lepus, 498, 502, 508

europasus, Lepus europteus, 502

europaeus, Macrotus, 250

europaeus, Meles, 348

europaeus, Sus, 957

europea, Rupicapra, 993

euryale, Rhinolophus, 155

Euryalus, 137

euryrhinus, Ursus, 286

Euvespertilio, 167

Euvesperugo, 203, 243

eversmanni, Myrmarctos, 287

Evotomys, 623

,, cassarius, 645

glareolus, 626

britannicus, 634
glareolus, 632
hallucalis, 643
helveticus, 640
istericus, 037
nageri, 641
norvegicus, 638
suecicus, 636
vasconise, 639
,, rufocanus, 648
,, rutilus, 646

,, skomerensis, 644

exilis, Arvicola scherman, 747

exilis, Sorex, 55

exitus, Arvicola scherman, 740

Exochurus, 167

exsul, Microtus agrestis, 669

fferoensis, Mus, 875

fasroensis, Mus musculus, 875

fagorum, Mustela martes, 875

falciger, Ursus, 286

fatioi, Pitymys, 763

fatioi, Pitymys multiplex, 763

favonicus, Myotis bechsteinii, 179

Felidte, 455

Pelis, 456

,, agrius, 470

,, sarda, 468

,, silvestris, 457

,, ,, grampia, 464

,, ,, silvestris, 462

„ ,, tartessia, 405

feunicus, Rangifer, 981

fennicus, Rangifer tarandus, 981

ferox, Catus, 462

ferruginea, Herpestes ichneumon, 441

ferrugiueus, 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
Hava, Arvicola arvalis, 6'JO
ilavescens, j\Ius musculus, 870, 878
llavoscens, Nanuugo pipistrellus, 205
Ilavescens, Talpa europrea, 4
fiavicans, INIustela putorius, 423
flavicollis, Apodemus, 828
flavioollis, Apodemus flavicollis, 829
flavicollis, Mus, 829, 831
flavus, Cauis lupus, 813
flavus, Castor, 947
flavus, Lepus timidus, 502
flavus, Mus musculus, 869
fluviatilis, Lutra, 356
fluviatilis, Sorex, 69
fodiens, Grossopus, 70, 73
fodions, Cuniculus, 490
fodiens, Neomys, 66
fodieus, Neomys fodiens, 69
fodiens, Sorex, 42, 69, 70
foetens, Viverra, 423
fostidus, Putorius, 423
Fa3torius, 384, 415, 418, 428
foiua, jMartes, 374
foina, Martes foina, 375
foiua, Mustela, 375
formicarius, Ursus, 286, 297
formioarius, Ursus arctos, 287
fretalis, Sorex araneus, 45
fridarieusis, 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, 884

gale, Mustela, 402

Gale my s, 20, 65

,, pyreuaicus, 21

,, ,, pyrenaicus, 26

„ „ rufulus, 26

galinthias, Mustela, 414

galinthias, Putorius (Ictis) nivalis, 414

galinthias, Putorius nivalis, 414

gallaacius, Lepus granatensis, 517

galliardi, Arvicola arvalis, 683

gallia3. Castor, 947

gallica, Viverra Geuetta, 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 geuetta, 451
genetta, Viverra, 451
gerbei, Arvicola (Microtus), 773
gerbei, Pitymys, 773
gerbii, Arvicola (Microtus), 773
gerbii, Pitymys, 773
germauicus, Cervus elaplius, 965
germanicus, Mus cricetus, 602
germanicus, Rhinolophus ferrum-

equinum, 142
ghidinii, Vespertilio, 179
ghidinii, Vespertilio bechsteinii, 179
glareolus, Arvicola, 632, 641, 043
glareolus, Evotomys, 626, 645
glareolus, Evotomys glareolus, 632
glareolus, Hypudaeus, 632
glareolus, Mus, 632
Glires, 481
Gliridse, 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, 677
gottbardi, Sciurus vulgaris, 910
graeca, Sciurus vulgaris, 910
graecus, Canis, 315
grseous, Spalax, 695
grsecus, Spalax greecus, 1000
graicus, Capra ibex, 989
grampia, Felis, 464
grampia, Pelis silvestris, 464
granarius, Sorex araneus, 52
granatensis, Lepus, 515, 516
granatensis, Lepus europajus, 516
granatensis, Lepus granatensis, 516
graudis, Ursus arctos, 286
gregarius, Mus, 668
grisea, Herpestes ichneumon, 441
grisea, Talpa europsea, 4

1008

griseogularis, Crossopus ciliatus, 70
griseus, Ursus arctos, 286
groenlandicus, Canis, 324
Gulo, 433

„ gulo, 434
gulo, Gulo, 434
gulo, Mustela, 434
Guloninse, 432
guttatus, Citellus, 829
guttatus, Mus citellus, 929
gutfculatus, Spermophilus, 929
gymnesicus, Eliomys, 558
gymnesicus, Eliomys quercinus, 558

hallucalis, Evotomys glareolus, 643

hallucalis, Evotomys nageri, 643

hamiltonii, Eliomys, 551

Hamster, 596

hamulicornis, Eupicapra, 993

hartingi, Microtus, 704

bayi, Mus sylvaticus, 810

hebridensis, Apodemus, 824

hebridensis, Mus, 824

bebridensis, Mus sylvaticus, 824

Heliomys, 596

bellenicus, Spalax monticola, 1000

belvetica, Ebinolophus euryale, 150

belveticus, Evotomys glareolus, 640

helveticus, Evotomys bercynicus, 640

belvolus, Mus musoulus, 869

Hemiotomys, 659, 723

bercegovinensis, Spalax monticola,
1000

bercynicus, Evotomys bercvnicus,
632, 637

bercynicus, Hypudreus, 632

bercynicus, Sorex alpinus, 63

Herinaceus, 114

bermanni, Sorex, 42, 70

berminea, Mustela, 387

Herpestes, 440

bibericus, Putorius, 407

bibernica, Mustela, 398

bibernicus, Lepus, 531

bibernicus, Lepus timidus, 531

bibernicus, Mus, 858

bibernicus Mus (Epimys) norvegicus,
858

bibernicus, Putorius, 398

bibernicus, Putorius (Ictis), 398

bibernicus, Sorex rusticus, 55

bippocrepis, Vespertilio, 142

bipposideros, Rbinolopbus, 147, 150

bipposideros, Rbinolopbus bippo-
sideros, 149

bipposideros, Vespertilio, 149

bipposiderus, Rbinolopbus, 150

Hircus, 988

birta, Arvicola, 673

birtensis, Apodemus, 825

birtensis, Mus, 825

birtensis, Mus sylvaticus, 825

birtus, Microtus agrestis, 673
bispanica, Capra, 991
bispanica, Capra pyrenaica, 991
bispanica, Viverra genetta, 451
bispanicus, Cervus elapbus, 969
bispanicus, Erinaceus europseus, 122
bispanicus, Lepus, 516
bispanicus, Mus spicilegus, 879
Histrix, 542
Homalurus, 29
bomorodalmasiensis, Rbinolopbus

unibastatus, 142
bomorodensis, Rbinolopbus ferrum-

equinum, 142
bortualis, Eliomys, 551
bumeralis, Vespertilio, 169
bungaricus, jNIesospalax, 1000
buugaricus, Spalax, 889, 894
bungaricus, Spalax bungaricus, 1000
bungaricus, Spalax typblus, 894
buxleyi, Lepus, 491
buxleyi, Oryctolagus cuniculus, 491
byberna, Mustela erminea, 387
bybridus, Lepus, 503
bybridus, Lepus europaeus, 508
bybridus, Mus decumanus, 858
Hydrogale, 65
Hydromustela, 415
bydropbilus, Sorex, 69
Hydrosorex, 65
bydruntina, Crocidura, 101
bypomelas, Vulpes, 331
Hypsugo, 202
Hypudaeus, 623
Hystricida3, 542
Hystridte, 542
Hystrix, 542

„ cristata, 543

iberica, Mustela nivalis, 407

iberica, Putorius nivalis, 407

ibericus, Arvicola, 782

ibericus, Arvicola (Microtus), 782

ibericus, Microtus, 782

ibericus, Pitymys, 7S0, 782

ibericus, I'itymys ibericus, 782

ibericus, Putorius nivalis, 407

Ibex, 988

ibex, Capra, 989

ibex. Ibex, 989

ichnusse, Crocidura, 998

icbnusffi, Vulpes, 336

icbnusae, Vulpes vulpes, 336

Ictis, 384

iculisma, Crocidura mimula, 98

ignotus, Crossopus on Sorex, 43, 70

illyricus, Arvicola, 741

illyricus, Microtus musignani, 741

litis, Mustela, 423

incertus, Arvicola, 690

incertus, Microtus, 690

incertus, Microtus arvalis, 691

1009

iucortus, Pitymys, 691

incisivus, Vespertilio, 226

infectus, Putorius, 423

infuscatus, Sciurus, 916

iiifuscatus, Sciurus vulgaris, 916

inodorus, Sorex, 101

lusectivora, 1, 134

insul, Microtus agrestis, 669

iusulauus, Rhinolophus ferruiu-equi-

num, 147
insularis, Eptesicus serotinus, 226
insularis, Glis, 578
insularis, Lemmus, 668
intermedia, Arvicola, 672
intermedins, Dyromys uitedula, 569
intermedins, l\Ius, 803
intermedins, j\Ius(Epimys) rattus,854
intermedins, Mus sylvatieus, 803
intermedins, ;\Ius rattus, 854
intermedius, Myoxus, 569
intermedins, Sorex, 62, 70
iuterstriatus, IMus, 539
interzouus, ^Mus, 539
isabellinus, Vespertilio, 226
islandicus, Mus, 803
Isotus, 167

istericus, Evotomys glareolus, 637
istricus, Spalax, 1000
italiea, Arvicola amphibius, 740
italicus, Arvicola, 740
italicus, Erinaceus europa^ui, 123
italicus, Glis, 578
italicus, Glis glis, 578
italicus, Putorius nivalis, 405
italicus, lihinolophus ferrum-cqui-

num, 142
italicus, Sciurus, 912
italicus, Sciurus vulgaris, 912, 913
iturissius, Lcpus granatensis, 518

jenaeusis, Tbalassaretos, 298
jcudii, Heliomys, 602
jonrensis, Capra icgagrus, 992
jubata, Alcos, 978

kattlo, Felis, 471
kirscbbaumii, Plecotus, 257
kisnyiresiensis, Rbinolopbus biha-

status, 150
kublii,-Pipistrellus, 215
kuhlii, Vespertilio, 215, 234
kuhlii, Vesperngo, 216
kuhli, Pipistrellns, 216

labiosus, Sorex, 35
Lagidic, 484
Lagomys, 931
lagopuii, Canis, 319
lagopus, Vulpes, 319
Lagos, 495

lanatus, Vespertilio, 184
lappouum, Tarandus, 98U
lardarius, Vespertilio, 245

lasiopterus, Vespertilio, 244
Latax, 354
Lataxiua, 354

latifrons, Arvicola agrestis, 671
latinornm, Martes martes, 373
latiuorum, Mustela martes, 373
latipinnis, Vespertilio, 193
lebruni, Jlicrotus (Cbionomys), 719
lebrunii, Arvicola, 719
lebrunii, Jlicrotus, 718
lebrunii, Microtus lebrunii, 719
Ijeggada, 863
leisleri, Nyctalus, 252
leisleri, Vespertilio, 252
leisleri, Vesperngo, 252
Lemmus, 614

,, lemmus, 615

crasaidens, 621
lemmus, Lemmus, 615
lemmus, IMus, 615
lemmus, IMyodes, 615
leponticus, Microtus, 764
Leporidcfi, 484
Leporina, 484
Lepus, 484, 495

cretieus, 512
em-opaus, 498

,, corsicanus, 507
,. europaeus, 502
bybridub, 508
meridiei, 506
oeeidentalis, 504
pyreuaicus, 506
■ ,. transsylvanicus, 503
granatensis, 515

gallbceius, 517
granatensis, 510
iturissius, 51b
liibcruicns, 531
meditcrraneus, 513
parnassius, 519
timidus, 522

!, scoticus, 529
,, timidus, 526
,, varronis, 528
leucitthiops, Cervus dama, 971
leucippe, Vespertilio, 219
leucocephalus, Mus sylvatieus, 803
Leucocyou, 318
Leucodon, 86
leucodon, Crocidura, 88
leucodon, Crocidura russula, 83
leucodon, !Sorcx, 88
leucodus, Crocidura, 88
leucogastcr, jMus, 854
Lenconoe, 166

leucopictus, Arctomys citellus, 929
Leucorbyuchus, 65
leucotis, Sorex fodiens, 70
lencourus, Sciurus vulgaris, 907
leucurus, Arvicola, 722, 719
leueurus, Microtus lebrunii, 722

3 X

1010

leucurus, Microtus (Cliionomys) le-

bruni, 722
leucurus, Sciurus, 907
leucurus, Sciurus vulgaris, 'J07
leucurus, Sorex, 101
leveruedii, Arvicola, G71
Icveruedii, Microtus agrcstis, G71
Icvis, Microtus, 687
levis, Microtus arvalis, 687
lilffius, Sciurus vulgaris, 913
lilfordi, Lepus, 51G
limbatus, Nannugo pipistrellus, 205
limnophilus, Vespertilio, 189
lineatus, Canis vulpus, 33U
lineatus, Sorex, 69
linneana, Amphisorex, 70
Lipura, 931
lirieaudatus, Sorex, 69
littoralis, Hypudccus paludosus, 738
loiigobarda, Sorex alpiuus, 62
longobardus, Sorex alpiuus, 70
Lontra, 354
loriger, Sicista, 537
loriger, Sininthus, 537
lucanius, Sorex iniuutus, 60
lugubris, Vespertilio, 169
lupulinus, Felis, 471
Lupus, 304

lupus, Canis, 305, 313
lupus, Canis lupus, 313
luscus, Gulo, 434
lusitanica, Capra, 991
lusitanica, Capra pyrenaica, 991
lusitanica, Eliomys uitela, 560.
lusitanicus, Arvicola (Microtus), 776
lusitanicuh, Eliomys, 560
lusitanicus, Eliomys quercinus, 560
lusitanicus, ;\licrotus (Pitymy^^), 776
lusitanicus, T\Ius spicilegus, 882
lusitanicus, Pitymys, 776
lutoa, Canis vulpcs, 331
lutea, Talpa curopica, 4
lutcsccns, Lcpus timidus, 531
Lutra, 354

Intra, 355
lutra, Lutra, 355
lutra, ]\Iustcla, 355
Lutreola, 415
lutreola, Fcetorius, 415
lutreola, Mustela, 415
lutreola, Putorius (Lutreola), 415
Lutrinffi, 354
Lutris, 354
Lutrix, 354
Lutrogale, 355
Lutronectes, 355
lycaon, Canis lupus, 313
Lyciscus, 304
Lyuceus, 470
Lynchus, 471
lyncula, Felis, 471
Lynx, 470

Lynx lynx, 471

,, pardellus, 475
lynx, Felis, 471
lynx. Lynx, 471

.Macblis, 970

machlis, Alces, 978

macropterus, Vesperugo pipistrellus,

204
Macrospalax, 1000
macrotricbus, Sorex, 35
IMacrotus, 256
macrourus, Sorex, 70
maculata, Mustela erminea, 387
maculata, Talpa em'opsea, 4
maculatus, Lepus timidus, 502
maculatus, Mus agrarius, 836
maculatus, Mus musculus, 869
magnus, Vespertilio, 245
major, Canis lupus, 313
major, Crocidura, 101
major, Fcetorius pusillus, 405
major, !Mus sylvaticus, 829
major, ^lustela erminea, 389
major, Rhinolopbus, 142
major, Ursus arctos, 286
major, Vespertilio, 245
major, Vespertilio ferrum-equiuum,

142
majori, Vespertilio, 187
Mangustu, 440

manium, Putorius putorius, 423
maral, Cervus elaphus, 965
marginatus, Pipistrellus, 215
marginatus, Vespertilio, 215
marine, Microtus (Pitymj's), 777
luaricC, Pitj^mys, 777
marianensis, Jleles meles, 352
inarianeusis, Jleles taxus, 352
marinus, Lutra vulgaris, 356
marinus, Ursus, 298
maritimus, Tbalarctos, 298
maritimus, Ursus, 298
maritimus, Ursus (Thalassarctos), 298
]\Larmota, 931

bobak, 937
marmota, 932
marmota, Marmota, 932
marmota, ilus, 932
marmotta, Arctomys, 932
JIarsipoliemus, 238
Martes, 365, 440

,, bunites, 380

,, foina, 374

foina, 375
mediterranea, 380
martes, 306

,, .. latinorum, 378

martes, 372
martes, IMartes, 366, 372
martes, !Martus martes, 372
martes, ]Mustela, 372

101

matechiei, Ovis, 987
maura, Cervus dama, 971
mauricus, Cervus, 970
maurus, Vesperugo, 220
maxima, Vesperugo nocfcula, 244
raaximus, Nyctalus, 244
maximus, Pterjgistes, 244
Mediocricetus, G05
mediterranea, Felis, 4G8
mediterranean ^Nlartes foina, 380
mediterranea, Mustela, 380
mediterraneus, Cervus, 969
mediterraneus, Lepus, 513
mediterraneus, ^leles males, 352
mediterraneus, Pipistrellus pipis-

trellus, 205
raedius, Arvicola, 70S
medius, Lepus, 502, 508
megalotos, Plecotus, 257
mogapodius, Vospertilio, 187
mehelyi, Rhinolophus, 159
raehelyi, Rhinolophus euryale, 159
melanodon, Sorex, 35
melanogaster, Canis, 331
nielanogaster, Vulpes vulpes, 331
melanopterus, Vespertilio, 204
melas, Genotta, 451
melas, Genetta genetta, 451
Meles, 341

,, arcalus, 352

,, meles, 343

,, ,, marianensis, 352

„ meles, 348
meles, ileles meles, 348
meles, MeJes, 343, 348
meles, Taxus, 348
meles, Ursus, 348
Melesium, 341
melina. Lynx \iilgaris, 471
Melintp, 341
melonii, Glis, 579
melonii, Cllis glis, 579
meridianus, ^licrotus arvalis, 080
meridiei, Lepus europiPus, 500
meridionalis, Lepus, 503, 510
meridionalis, ]Mus, 844
meridionalis, ilus (Apodemus), 844
meridionalis, I'utorius (letis) nivalis,

405
meridionalis, Putorius vulgaris, 405
meridionalis, Sc.iurus, 912
meridionalis, Sus, 960
meridionalis, Sus serofa, 960
meridionalis, Vulpes vulgai-is, 331
!Mesocricetus, 605

,, newtoni, 606

Mesospalax, 1000
messorius, Mus, 844
Meteorus, 225, 238
iliccQchiroptera, 134
Micromys, 791, 840

,, mjnvitus, 841

Micromys minutus pratcnsis, 846
,, ,, Boricinus, 841

]\Iicrospalax, 888
!\Iicrotinpe, 610
:\Iicrotus, 658, 659, 712, 752
,, agrestis, 662

,, ,, agrestis, 668

,, ,, bailloni, 672

,, exsul, 609
,, . ., hirtus, 673

,, levernedii, 671
,, ,, negleetus, 675

,, „ rozianus, 680

,, angularis, 706
,, arvalis, 681

arvalis, 683
duplicatus, 686
,, .. levis, 087

,. ,, meridianus, 686

asturianus, 093
cabrerne, 701
dentatus, 703
,, hartingi, 704
,, incertus, 090

,, lehrunii, 718
,, ,, lebruuii, 719

,, ,, leucurus, 722

,. nivalis, 713

,, ,, aquitauius, 717

nivalis, 716
orcadensis, 694
,, ratticeps, 708

,, sandayensis, 696

„ ,, sandayensis,

697
,, westrae, 698

sarnius, 700
ulpius, 723
microurus, Leucodon, 88
Micrurus, 752
midas, Dysopes, 277
milleri, Neomys, 78
mimula, Crocidura, 94, 95
mimula, Crocidura mimula, 95
miniatus, iNIus, 844
minima, Vesperugo noctula, 245
minimus, Mus, 844
minimus, ]\Ius minutus, 844
minimus, Mus (Apodemus) minutus,

844
minimus, Rhinolophus, 151
minimus, Rhinolophus hipposiderus,

151
minimus, Rorex, 55
Miniopteri, 268
]Miniopterina>, 268
Miuiopteris, 268
Miniopterus, 268

schreibersii, 269
Minneopterus, 268
minor, Arvicola amphibius, 740
minor, Canis lupus, 313

1012

minor, Cervus elaphus, 0C9

minor, Crocidura aranea, 95, 101

minor, Lutra, 415

minor, i\Ius rutilus, 632

minor, Mustola, 402

minor, Neomys fodiens, 70

minor, Rhinolophus, 149

minor, Spalax, 745

minor, Ursus arctos, 2SCt

minor, Vespertilio ferrum-equinum,

149
minous, Acomys, 883
minous, Acomys dimidiatus, 883
minuta, Crocidura, 95
minuta, Phyllorhina, 149
minutellus, Vespertilio, 184
minutissimus, Vespertilio, 204
miiintus, j\Iicromys, 841
minutus, ]\Ius, 844
minutus, Putorius, 402
minutus, Putorius (lotis) vulgaris, 403
minutus, Rhinolophus hipposiderus,

154
miuutus, Sorex, 53, 55
minutus, Sorex minutus, 55
minutus, Vespertilio, 154
ilinyopterus, 2riS
mollis, Sorex vulgaris, 43
molliBsimus, Musculus, 871
Molossi, 276
Molossidte, 276

montanus, Cervus, vulgaris, 966
montanus, Plecotus auritus, 257
montieola, Arvicola, 749
monticola, Arvicola scberman, 749
montieola, il^sospalax, lOOU
monticola, Putorius nivalis, 403
montieola, Spalax, 894
monticola, Spalax monticola, KXJO
Mops, 276

moreotica, Canis aureus, 315
moscbata, Crocidura, 101
multiplex, Arvicola, 7(!4
multiplex, Pitvmvs, 764
Mungo, 440
JIungos, 440

,, widdringtonii, 441
muralis, !Mus, H74
muralis, Mus musculus, 874
iluridoe, 591
!Murina, 530
jNIurinfe, 791

murinus, Vespertilio. 192, 238
:\rus, 791, 840, 848, 863

., {n=roensis, 875

,, muralis, 874

,, musculus, 865

,, ., azoricus, 871

musculus, 869

,, spicilegns, 877

hispanicus, 879

,, ,, lusitanicus, 882

Mus spicilegus spicilegus, S7S
musaraneus, Sorex, 101
^luscardinidjc, 549
Muscardiuus, 5S3

,, avellauarius, 583

muscardinus, Myoxus, 583
Muscardiuus pulcher, 590
Musculus, 863
musculus, :\Ius, 865, 869
musculus, Mus musculus, 869
musculus, Sorex, 70
musignani, Arvicola, 744
musignanoi, Arvicola, 744
:\Iusimon, 986
musimon, /F^goceros, 987
musimon, Ovis, 987
musiniani, ^licrotus, 735
Musmon, 986
musmon, Ovis, 987
IVIustela, 365, 381, 384
africana, 412
,, erminea, 385

ffistiva, 389
,, .. erminea, 387

,, ., ricinrp, 397

stal)ilis, 390
,, galintliias, 414
liibernica, 398
,, lutreola, 415
nivalis, 401

boccamela, 405
,, ., iberica, 407

,, nivalis, 402
,, putorius, 418

aureolus, 425
,, ., putorius, 423

Mustelidsp, 340
Mustelina, 384
:\rusteliu», 364
:\rygale, 21
Mygaliua, 20
Mvnomes, 659
Myodes, 614, 623
Myogale, 21
Myogalea, 21 «

IMyopus, 611

schisticolor, 611
Myosictis, 65
myosotis, lilyotis, 193
myosotis, Vespertilio, 192
Myospalax, 887
INIyotis, 166

bechsteinii, 179
,, capaccinii, 187
,. ilasycneme, 189
,, daubentonii, 184
emarginatus, 177
inyotis, 192
mystaeinus, 169
nattereri, 174
nxyguatbus, 199
myotis, Myotis, 192, 193

1013

myotis, Vespertilio, 102
]Myoxida5, 5-i9
INIyosiiia, 54'J
]\Iyoxub, 5(3G, 572
Myrmarctos, 285
myrmephagus, Ursus, 280
mystacinus, Mus, 704
mystaciniis, Myotis, 1G9
mystacinus, Vespertilio, 1G9, 189

nageri, Evotomys, 641

nageri, Evotomys glareolus, 641

nageri, HypudiBus, 641

naiaa, Neomys fodiens, 70

Nannomys, 863

Nanoospalax, 888

Nannugo, 203

nanus, Neomys fodiens, 71

natans, Sorex, 70

nathusii, Pipistrellus, 213

nathusii, Vespertilio, 213

nathusii, Vesporugo, 213

nattereri, ^lyotis, 174

nattereri, Vespertilio, 174

nebrodensis, Arvicola, 770

nebrodensis, Pitymys, 770

nebrodensis, Pitymys savii, 770

jieglectus, Arvicola, 075

neglectus, ]\Iicrotus agrestia, 672, 673,

675
neglectus, Vespertilio, 177
nehringi, Cricetus, 605
nehringi, Cricetus cricetus, 605
nemoralis, Mus, 854
Neocyon, 304
Neogale, 384
Neomys, 65

,, anomalus, 81
fodiens, 66, 70
., bicolor, 73
fodiens, 09

,, milleri, 78
nesiotes, Erinaceus, 129
nesiotes, Erinaceus europrous, 129
newtoni, Cricetus, 006
newtoni, Cricetus (^Nlesocricetus), 000
newtoni, ilesoericetus, 000
niger, Arvicola terrestris, 740
niger, Canis lupus, 313
niger, Capreolus vulgaris, 974
niger. Castor, 947
niger, Cricetus cricetus, 602
niger, Cricetus vulgaris, 602
niger, Dama platyeeros, 971
niger, I^epus timidns, 502
niger, ilus musculus, 809
niger, Jfus sylvaticus, 803
niger, Sciurus vulgaris, 905
niger, Ursus arctos, 285
nigra, Arctomys marmota, 932
nigra, Arvicola agrestis, 071
nigra, Canis vulpes, 331

nigra, Sorex vulgaris, 43

nigra, Talpa eucopfpa, 4

nigra, Vespertilio pipistrellus, 204

nigrans, Vespertilio, 220

nigrosceus, Sciurus vulgaris, 910

nigricans, Arvicola amphibius, 730

nigricans, Brachyotus mystacinus,

169
nigricans. Hamster, 602
nigricans, Lepus timidus, 502
nigricans, ]\Iub arvalis, 668
nigricans, Nannugo pipistrellus, 205
nigricans, Pipistrellus, 204
nigricans, Sorex fodiens, 70
nigricans, Vespertilio mystacinus, 109
nigripos, Sorex, 70
uigro-argenteus, Canis, 330
nigro-fuscus, Vespertilio mystacinus,

109
nigrocaudatus, Canis vulpus, 330
nilssoni, Eptesicus, 234
nilssoni, Vesperugo, 234
nilssonii, Eptesicus, 234
nilssonii, Vespertilio, 234
nilssonii, Vesperugo, 234
nitedula, Dyromys, 567, 508
nitedula, Dyromys nitedula, 508
nitedula, Mus, 551, 568
nitedulus, Eliomys, 508
nitela, Myoxus, 551
nivalis, Arvicola, 716, 719, 722
nivalis, Microtus, 713, 716
nivalis, Microtus (CMonomys), 716
nivalis. Microtus nivalis, 716
nivalis, Mustela, 401, 402
nivalis, ilustela nivalis, 402
nivalis, Putorius, 403
nivalis, Putorius (Ictis), 403
niveus, iMus musculus, 869
nivicola, Arvicola, 710
Noetula, 225
noctula, Nyctalus, 245
nootula, Ptorygistes, 245
noctula, Vespertilio, 245
noctula, Vesperugo, 245
Noctulinia, 242
nordmanni, Sminthus, 537
normalis, Ursus arctos, 286
norvegicus, Epimys, 858
norvegicus, Evotomys, 638
norvegicus, Evotomys glareolus, 638
norvegicus, Mus, 858
norvegicus, ]Mus (Epimys), 858
norvegicus, Ursus, 280
norwegicus, Epimys, 858
norwegicus, Lemmus, 015
nuda, Sorex vulgaris, 42
nudipes, Lutra, 350
numantius, Sciurus vulgaris, 914
Nutria, 355
Nyctalus, 242

,, azoreum, 254

1014

N,yctalus leisleri, 252

,, maximus, 244

„ noctula, 245
Nyctinoma, 276
Nyctinomes, 276
Nyctinomia, 276
Nyctinoraops, 276
Nyctinomus, 276

,, teniotis, 277

Nyctiptenus, 225
Nystactes, 166

obscurus, Rhinolopbus ferrum-equi-

num, 143
occideiitalis, Erinaceus europspus, 120
occidentalis, Lepus europspus, 504
occidentalis, Ovis musimon, 987
occidentalis, Talpa, 15
occidcutalis, Talpa crmca, 15
Ocbetomys, 723

ocbromixtus, Pipistrellus savii, 220
ocbromixtus, Vespertilio, 220
Odiufclurus, 446
CEdocepbalus, 542
okenii, Vespertilio, 226
Ommatostei-gus, S88
orcadonsis, ^licrotus, 094
oi'ieutalis, Lupus, 313
oruata, Tlupieapra, 994
orsiuii, Vespertilio, 269
Oryctolagus, 484

funiculus, 485
,. ,, cuniculus, 490

,, huxleyi, 491
oryzivorus, Mus, 844
Otisorex, 29
otus, Vespertilio, 256
Ovis, 986

,, musimon, 987
oxygnatbus, Mjotis, 199
oxygnathus, ^lyotis myotis, 199
oxygnatbus, Vespertilio, 199
Oxygons, 304
Oxyrbin, 29

pachygnatlius, Vespertilio, 252
Pacbyomus, 225
Pacbyura, 81

,, etrusca, S3

pacbyurus, Sorex, 83
pallidus, Eliomys, 559
pallidas, Rbinolophus bipposideros,

151
pallidus, Sorex araneus, 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 (Eucervaria), 475
pardellus. Lynx, 475
Pardina, 470
pardina, Felis, 475
parnassius, Lepus, 519
parnassius, Lepus europ.ipus, 519
parva, Rupicapra, 995
parva, Rupicapra rupicapra, 995
parvulus, Drymomys, 869
parvulus, ]\Ius, 844
parvus, :\Ius sylvaticus, 803
pascuus, Pitymys ibericus, 783
Paurodus, 86
peechioli, Mus, 810
pelandonins, Pitymys, 778
pellucens, Vespertilio, 187
pendulinus, Mus, 844
peninsulffi, Oenetta, 451
peninsulfP, Genetta genetta, 451
ponnanti, Ampbisorex, 73
peregusna, Mustela, 429 •
peregusna, Vormcla, 429
pcrsonatus, Sorex, 35
perspicillatus, Vespertilio, 142
portinax, Arvicola, 740
Petauristidie. 940
petropbilus, llypudmus, 716
pbasma, Rbinolopbus, 151
Pbvllostomata, 136
Pbyllotis, 137
picta, Capra segagrus, 992
picteti, Mus, 854
pictus, ^ICgoceros, 992
Pinalia, 65
Pinemys, 752

pipistrelle, Vespertilio, 204
Pipistrellus, 202

kublii, 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, 760
,, depressus, 779
druentius, 762'
duodeciracostatus, 784
fatioi, 763
gerbii, 773
ibericus, 780

centralis, 782
ibericus, 782
pascuus, 783
,> ,, regulus, 784

,, lusitanieus, 776

marise, 777
,, multiplex, 764

1015

Pitymys ncbrodeusis, 770
„ pelandonius, 778
„ planiceps, 772
„ proviiicialis, 785
„ pyreiiaicus, 770
,, „ brunueus, 772

„ ,, pyrouaicus, 771

,, savii, 768
,, subterraneus, 755
,, ,, capuciiius, 7C0

„ „ subterraueus,

758
„ thomasi, 786
planiceps, Pitymys, 772
Platyceros, 970
platyoeros, Cervus, 970
platyrhynchus, Cervus (Taramlus),

985
platyrhynchus, liaugifer, 985
Plecotus, 256

,, auritus, 256
Plerodus, 81

plumbeus, Cervus capreolus, 97i
polaris, Ursus, 298
poliogaster, Crocidura, 101
polonicus, Spalax, 1000
polonicus, Ursus arctos, 286
poschiavinus, IMus, 869
poschiaviuus, Iilus musculus, 870
pratcnsis, Arvicola, 632
pratensis, Micromys minutus, 846
pratensis, Mus, 816
pratensis, Mus minutus, 846
pratensis, Mus (Apodemus), minutus,

846
Praticola, 659, 712, 723
princeps, jMus sylvaticus, 829
principalis, Arvicola arvalis, 683
Procerus, 979
Procervus, 979
proprius. Castor, 918
proterus, Ycspei'tilio, 215
proviiicialis, Pitymys, 785
Psammomys, 752
Pscudoconomys, 863
psilurus, Sorcx, 70
Pternopterus, 167
Pteromidffi, 940
Pteromyinte, 940
Pteromys, 941
Pterygistes, 242
pulcher, Muscardiuus, 590
pulchra, Crocidura russula, 103
pullatus, Vesperugo kuhlii, 216
pumilio, Sorex, 55
pumilus, Mus, 844
pumilus. Sores, 55
pusillus, Foetorius, 403
pusillus, Vcspertilio, 204
Putorius, 418
putorius, Ftctorius, 123
putorius, ilustehi, 418, 123

putorius, Mustela putorius, 423
putorius, Putorius, 423
pygniicus, Sorex, 55
pygmicus, Vcspertilio, 204
pyreuccus, Ursus, 286
pyrenaica, Capra, 989, 990, 991
pyrcnaica, Capra pyrenaica, 990
pyrenaica, Mygale, 26
pyrenaica, Myogale, 26
pyrenaica, Rupicapra, 995
pyrenaica, Rupicapra rupicapra, 995
pyrenaicus, Arvicola, 771
pyrenaicus, Galemys, 21
pyrenaicus, Galemys pyrenaicus, 26
pyrenaicus, Glis glis, 582
pyrenaicus, Lepus europseus, 506
pyrenaicus, Pitymys, 770, 771
pyrenaicus, Pitymys pyrenaicus, 771
pyrenaicus, Sorex araneus, 41
pyrenaicus, Ursus, 286
pyrenaicus, Ursus arctos, 287

quadricaudatus, Sorex, 42
quercinus, Eliomys, 551, 558
quercinus, Mus, 551
quercinus, Myoxus, 551

Rangifer, 979

,, fennicus, 981
,, platyrhynchus, 985
,, tarandus, 980

rangifer, Cervus tarandus, 980
ratticeps, Arvicola, 70S
ratticeps, Microtus, 70S
Rattus, 848

rattus, Epimys, 849, 853
rattus, Epimys rattus, 853
rattus, JMus, 853
rattus, Mus (Epimys), 853
rcgularis, Pitymys ibericus, 784
regulus, Pitymys ibericus, 784
rcniifer, Sorex, 69
rhenauus, Cervus, 966
Rhiuocrepis, 137
Rhiuolophidct", 130
Rhiuolophina, 136
Rhiuolophus, 137

„ blasii, 162

,, euryale, 155

„ ferrum-equinum, 139

fer-
rum-equinum, 142
,y .. insu-

lanus, 147
,. hipposideros, 117

,• .. hipposi-

deros, 149
u J, minimus,

151
I. M miuutua,

154

1016

Rhinolophus mehelyi, 15'J

rhinclophus, Sorex, 35

rhodanica, Genetta, 452

rhodanica, Genetta genetta, 452

ricinfe, INIustela crmiuca, 397

ricime, Putorius erminea, 397

ricina;, Putorius crmincus, 3'J7

ripanus, Arvicola, G34

rivalis, Sorex, 70

robustus, DjTomys, 572

Kodentia, 481

roensis, Liitra, 35G

roniana, Talpa, 18

Roinicia, 202

rossicus, UrsuH arctos, 280

roumanicus, Erinaceus, 127

roumauiuus, Erinaceus curopicus,

127
rnzianui, Arvicola, 680
rozianu.-, IMicrotus agrestis, 080
rubeus, ]Mus, 836

rubidus, Evotomys hercynicus, S32
rubidus, Lemmus, 632
rufa, Arvicola agrestis. G71
rufa, Crocidura, 101
rufa, Talpa europaea, 4
rufesceus, Arvicola, 632
rufesceua, Vcbpertilio, 177, 245
rufescens, Vespertilio pipistrellus, 204
rufeseente-fuscus, Hypudteus, 758
rufocanus, Evotomys, 648
rufocanus, Evotomys (Cra.seomys), 643
rufocanus, Hypudaeus, 648
rufofubcus, Brachvotus myatacinus,

169
rufofubcuti, Hypudtcus. 758
rufula, Jlyogale pyrenaica, 26
I'ufula, JNIyogalea, 26
rufulus, Galemys pyrcnaicus, 26
rufus, Ecpus timidus, 502
rufus, Sciurus, 914
rufus, Sciurus vulgaris, 905, 909, 910
rufus, Ursus arctos, 286
Kupicapra, 992

,, ornata, 994
parva, 995
,, pyrenaica, 995
,, rupicapra, 993

rupicapra, Antilope, 995
rupicapra, Capella, 993
rupicapra, Capra, 993
rupicapra, Rupicapra, 993, 995
rubsicus, Ptcromys, 941
russicus, Sciuroptcrus, 9 1 1
russula, Crocidura, 99, 101
russula, Crocidura russula, 101
russulus, Sorex, 101
russus, Sciurus vulgaris, 909
rusticus, Sorex, 55
rutilaiis, Sciurus vulgaris, 910
rutilus, I'jvotomys, 646
rutilus, !Mus, 646

Sacalius, 304

sandayensis, Z\Iicrotus, 696, 697

sandayensis, Jlicrotus orcadensis, 697

sandayensis, Microtus sandayensis, 697

saatonus, Sorex, 40

santonus, Sorex arancus, 40

sapidus, Arvicola, 732, 733

sapidus, Arvicola sapidus, 733

sarda, Pelis, 468

sarda, Felis libyca, 468

sarda, Felis ocreata, 468

sardous, Sus scrofa, 960

sardus, Eliomys, 560

sarmatica, Slustela, 429

sarmaticus, Fcctorius, 429

sarmaticus, Putorius, 129

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, ]\lus, 745

schcrmaus, Mus, 745

schinzii, Vespertilio, 169

schibticolor, Lemmus, 611

schisticolor, I\Iyodes, 611

schisticolor, Myopus, 611

schrauldi, Vespertilio, 169, 177

schrcibersi, ^liuioptcrus, 269

schrcibersii, Miniopterus, 269

schreibersii, Vespertilio, 269

Sciurida;, 897

Sciuriua, 940

Sciurinte, 940

Sciuropterus, 941

russicus, 911

Sciurus, 898

„ vulgaris, 898
,. ,, alpinus, 914

,, ., ba;ticus, 923

,, ,, fuscoater, 910

,, ,, iufuscatus, 916

,, ,, italicus, 912

,, ,, leucourus, 907

liheus, 913
., ., numantius, 914

russus, 909
., seguric, 917
varius, 906
,, vulgaris, 905

scoticus, Cervus elaphus, 90S

scoticus, Lepus medius, 529

scoticus, Lepus timidus, 529

Scrofa, 966

scrofa, Sus, 957

seguric, Sciurus vulgaris, 917

selysii, Arvicola, 768

1017

selysii, Arvicola (Microtus), 762
Selysius, 166
Semicricetus, 605

septentrionalis, Lepus, 526

serbiciis, Spalax mouticola, 1000

serotina, Noctula, 215

serotine, Vespertilio, 226

serotinus, Eptesicus, 226

serotinus, Vespertilio, 226

serotinus, Vespcrugo, 226

setosus, Sus, 957

sibiricus, Pteromys, 911

Sicista, 536

,, loriger, 537
,, trizona, 539

Sicistiuaj, 536

sicula, Crocidura, 108

siculse, Myoxus, 854

siculus, Eptesicus, 239

siculus, Putorius (Ictis) nivalis, 405

siculus, Putorius nivalis, 405

siculus, Vosporus, 238

signatus, Canis lupus, 314

silacea, Vulpes vulpos, 333

silacous, Vulpos vulpes, 333

silvestris, Felis, 457, 462

silvestris, Eelis (Catus), 462

silvestris, Felis silvestris, 462

simplex, Arvicola arvalis, 683

Simplicidentata, 535

skomerensis, Evotoniys, 644

Sminthi, 536

Sminthinse, 530

Smiutlius, 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

Soricidaj, 28

soricinus, Micromys minutus, 844

soricinus, Mus, 844

sowerbyi, Crossopus, 73

Spalacidte, 887

Spalax, 887, 1000
„ dolbrogeae, 889
,, grsecus, 895

Spalax hungaricus, 894

speciosus, Muscardinus avellanarius,

590
sxieciosus, Myoxus, 590
spelsea, Myotis murina, 193
Spermopbilus, 924
spetsbergeusis, Cervus tarandus, 985
spicilegus, Mus, 877, 878
spicilegus, Mus spicilegus, 878
spitzborgeueusis, Alopcx, 324
spitzbcrgonensis, Canis lagopus, 324
spitzbergensis, Rangifer, 985
spitzbergeusis, Rangifer arcticus, 985
spitzbergensis, Thalassarctos, 298
spitzbergensis, Vulpes lagopus, 324
stabilis, Mustcla ermiuca, 390
stabilis, Putorius crraineus, 390
stabilis, Putorius (Ictis) ermiueus, 390
stagnatilis, Sorex, 70
staufferi, Vespertilio, 184
steuorostris, Ursus arctos, 287
stenotus, Vespertilio, 204
stimmingi, Arvicola (Microtus) rat-

ticeps, 708
stimmingi, IMicrotus ratticeps, 708
striatus, Mus musculus, 869
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, 636
Suidae, 966

suillus, Eriuaceus, 120
surmolottus, Mus, 858
Sus, 956
„ attila, 900
,, meridionalis, 960
,, scrofa, 957
suslica, Citellus, 929
suslica, Mus, 929
sylvatica, Capra rupicapra, 993
sylvatioa, Martes, 372
sylvaticus, Apodemus, 797, 803
sylvaticus, Apodemus sylvaticus, 803
sylvaticus, Lepus borealis, 526
sylvaticus, Mus, 803, 829
sylvestris, Felis, 462
sylvestris, Mus, 854
sylvestris, Mustela, 372
Sylvioola, 659
Synotus, 263
syrmiensis, Spalax monticola, 1000

tseniotis, Nyctinomus, 277
Talpa, 3

1018

Talpa cseca, 15
,, europsea, 3
,, ocoidentalis, 15
,, romana, 18
Talpidfp, 2
Talpince, 2
Talpoides, 887
Tarandus, 979
taraudus, Cervus, 980
tarandus, Eangifer, 980
tartessia, Pelis, 465
tartessia, Felis silvestris, 465
Taxus, 341
taxus, Meles, 348
tectorum, ]\Ius, 854
tenebricus, Arvicola, 735
tenebricus, Arvicola sapidus, 735
teniotis, Cephalotes, 277
teniotis, Nyctinomus, 277
terrestris, Arvicola, 738, 745, 746,

749
terrestris, Mus, 738, 745
Terricola, 752
tetragonurus, Sorex, 42
tetragonurus, Sorex araneus, 42, 43
Tetramerodon, 659
Tbalarctos, 297

,, maritimus, 298

Tbalassarctos, 297
Thalassiarcbus, 297
thomasi, Microtus (Pitymys), 786
tbomasi, Pitymys, 780
tboracinus, Sorex, 101
thotti, Capreolus caprcolus, 975
Tbous, 304

tigrina, Arctomys marmota, 932
timiduR, Lepus, 502, 508, 513, 522

527
timidus, Lepus timidus, 526
toscauus, Euryalus, 155
toscanus, Rhinoloplnis euryale, 155
Tragus, 988
tragus, Rupicapra, 993
Tralatitus, 167

traiissylvanicus, Capreolus, 975
transsylvanicus, Capreolus capreolus,

975
transsylvanicus, Lepus, 509
transsylvanicus, Lepus europaeus, 509
transsylvanicus, Spalax hungaricus,

1000
transsylvanus, Eptesicus serotinus,

220
transsylvanus, Vosporus serotinus, 226
transsylvaticus, Lepus europwus, 509 i
trausylvanus, Vesperus serotinus, 226 \
Trilatitus, 166 I

tripartitus, Mus, 539
tristriatus, Mus, 539
triticeus, Mus, 844 i
trizona, Sicista, 539
trizonus, Mus, 639

troglophilus, Rbinolopbus hipposi-

deros, 150
turcicus, Spalax monticola, 1000
typicus, Lepus mediterraneus, 513
typicus, Lepus timidus, 529
typicus, Meles meles, 348
typicus, Mus sylvaticus, 829
typicus, Putorius nivalis, 403
typicus, Rhinolophus ferrum-equi-

num, 143
typicus, Rhinolophus hipposiderus,

150
typicus, Sciurus vulgaris, 905
typus, Isotus nattereri, 174
typus, Myotis murinus, 193
typus, Pipistrellus, 204
typus, Plecotus auritus, 257
typus, Rhinolophus hipposideros, 150

ulpius, Jlicrotus, 723

ulpius, Microtus (Chionomys), 723

ungula, Vespertilio, 142

Ungulata, 955

unicolor, Sorex, 101

unicolor, Vesperugo nathusii, 213

unifer, Rhinolophus, 142

unihastatus, Rhinolophus, 142

Ursarctos, 286

Ursida;, 284

ursinii, Vespertilio, 209

Ursula, Vespertilio, 210

Ursus, 285

,, arctos, 285
ursus, Ursus, 285

vagans, Erinaceus algirus, 133
vagus, Sminthus, 539
variabilis, Arvicola arvalis, 683
variabilis, Lepus, 526, 528, 5-29, 531
variegata, Talpa europsea, 4
variegatus, Canis vulpus, 330
variegatus, Castor fiber, 947
varius, Capreolus vulgaris, 974
varius, Castor, 947
.varius, Cervus elaphus, 965
varius, Cricetus vulgaris, 602
varius, Dama platyceros, 971
varius, Mus musculus, 869
varius, Mus sylvaticus, 803
varius, Rattus domesticus, 853
varius, Sciurus vulgaris, 906
varronis, Lepus, 528
varronis, Lepus medius, 528
varronis, Lepus timidus, 528
vasconise, Evotoinys, 639
vasconife, 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
VespertilionidsB, 165
Vespertilioninse, 165
victorice, Capra pyronaica, 991
virgata, Folis, 412
virgulosus, Iilus, 539
Vison, 415

vison, Mustela putorius, 423
vispistrellus, Vespertilio, 215
vistulinus, Castor, 948
Viverridce, 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, Fojtorius, 403
vulgaris, Genetta, 451
vulgaris, Geuetta 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
vulgaris, 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, Vulpcs, 330
Vulpcs, 325

ichnusBe, 336

„ vulpes, 326

crucigera, 331

,. ,, silacea, 333

,, ,, vulpes, 330

vulpes, Canis, 330, 331
vulpes, Vulpes, 326, 330, 331
vulpes, Vulpes vulpes, 330
Vulpicanis, 304
vulpinus, Pelis, 471
vulpus, Canis, 330

westrse, Microtus sandayensis, 098
widdriugtoni, Herpestes ichneumon,

441
widdringtonii, Herpestes, 441
vyiddringtonii, Mungos, 441
wiedii, Vespertilio, 226
wingei, Dyromys nitedula, 570
wingei, Myoxus, 570
wintoni, Apodemus flavicoUis, 831
wintoni, Mus sylvaticus, 831

Zapodidse, 535
zemni, Glis, 1000
Zibelliua, 365

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