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Full text of "Catalogue of the mammals of Western Europe (Europe exclusive of Russia) in the collection of the British Museum"

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http://www.archive.org/details/cu31924003031220 



CATALOGUE 

OF THE 

MAMMALS 

OF 

WESTERN EUROPE 

(EUROPE EXCLUSIVE OF RUSSIA) 
IN THE 

COLLECTION 

OF THE 

BRITISH MUSEUM 



BY 

GEEEIT S. MILLEE 



LONDON 

PRINTED BY ORDER OF THE TRUSTEES OF THE 

BRITISH MUSEUM 

SOLD BY 

Longmans, Green & Co., 39, Patebnobteb Row, E.C. 

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

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

AND AT THE 

British Museum (Natural History), Cromwell Road, S.W. 
1912 

All rights reserved 



qL 

B56 "j> 



'9? 



A, 311151 



LONDON : 

PRINTED BT WILLIAM CLOWES AND SONS, LI5IITED, 

DOKE STREET, STAMFORD STREET, S.E., AND GREAT WINDMILL STREET, W. 



V>3 



PREFACE 

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

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

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

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



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

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

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

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

SIDNEY F. HAEMEE, 
Keeper of Zoology. 

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

October, 1912. 



INTKODUCTION 

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



INTRODUCTION 



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

Although unquestionably forming the largest of all collections 
of European mammals the material in the British Museum is 
not sufficient to be made the basis of a monographic study of 
the fauna. Free use has, therefore, been made, throughout the 
preparation of this Catalogue, of specimens in other collections. 
Chief among these are the United States National Museum in 
Washington and the private collection of Charles Mottaz in 
Geneva. The material at Washington, about 4000 specimens, 
is mostly from the following sources : (a) Sweden, Germany, 
Switzerland and Belgium, collected by J. Alden Loring ; (6) 
Sicily, Italy and the region of Barcelonnette, Basses-Alpes, 
France, by Dane Coolidge ; (c) south-western France, by 
Robert T. Young ; (d) north-eastern Germany, the Riesengebirge 
and Harz Mountains, by F. L. J. Boettcher. There are also 
miscellaneous smaller collections from Switzerland (G. S. Miller, 
L. Stejneger, E. H. Zollikofer), Belgium (de Selys-Longchamps), 
Holland (G. S. Miller), Denmark (L. Stejneger), Norway (T. 
Stejneger) and Sweden (Sundevall, Tullberg, Lonnberg). Finally, 
the Merriam collection, now the property of the U.S. National 
Museum but not yet catalogued, contains numerous European 
specimens, for the most part received from de Selys-Longchamps. 
The Mottaz collection, about 3000 specimens, is especially rich 
in series of the smaller mammals of Switzerland and the adjoin- 
ing portions of France ; it also contains useful material from 
Italy and western France (Charente). Other supplemental 
material to which I have been given free access, or which has 
been sent for examination in London or Washington, is contained 
in the museums of Madrid (types of Cabrera), Nimes (types of 
Crespon), Paris (types of Geoffroy and other historic speci- 
mens), Genoa (Italian Bats, Microtines and Ungulates), Turin 
(Italian mammals, especially Ungulates), Naples (type of Myotis 
oxygnathus Monticelli), Geneva (types of Fatio, authentic Swiss 

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



INTRODUCTION 



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

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

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

* In these lists 11,372 specimens are recorded. They are distributed 
as follows : Inseotivora, 1,777 ; Chiroptera, 2,210 ; Carnivora, 877 ; Kodentia 
Duplieidentata, 379, Rodentia Simplioidentata, 5,854 ; Ungulata, 284. 

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

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



INTRODUCTION 



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

The literature of European mammals is so voluminous, 
particularly as regards local lists and special notes on distri- 
bution, and it is for the most part based on conceptions of 
species and local races so different from those underlying the 
present work, that an amount of labour incommensurate with 
the importance of the results would be required to prepare 
extended bibliographical Tables for each form recognized. The 
citations are, therefore, restricted to those which seem of impor- 
tance in giving a clear idea of the systematic history of each 
animal ; that is, to the specific and sub-specific names under 
which it may have been described, to the first use of the actual 
binomial or trinomial here adopted, to the names used in the 
monographic works of Blasius, 1857, and Trouessart, 1910, and 
to any other publication which might seem pertinent to a 
particular case. 

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

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

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



INTRODUCTION i x 

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

G. S. M. 



Washington, 



1, 1912. 



Order. 


Number of 

genera 
recognized. 


Number of 

forms 
recognized. 


Number of 

forms not 

represented 

in B.M. 


Number of 

recognized 

forms not 

seen. 


Insbctivora 


7 


45 


5 





Chiroptera 


10 


33 








Oarnivora 


15 


47 


1 





EODENTIA DUPDICIDENTATA 


2 


19 


1 





„ SlMFI/ICIDENTATA 


26 


139 


6 


2 


Ungulata 


9 


31 


9 


4 


Total 


69 


314 


22 


6 



SYSTEMATIC INDEX 



ORDER INSECTIVORA. 



Family TALPID^E . . . 


2 


Sub-Family 1. Talpin^e . . 


2 


1. Talpa Linnaeus 


3 


1. europaea Linnseus . 


3 


2. caeca Savi .... 


. 15 


3. oocidentalia Cabrera . 


. 15 


4. romana Thomas 


. 18 


Sub-Family 2. Desmanum . 


. 20 


1. Galemys Kaup 


. 20 


1. pyrenaious Geofiroy . 


. 21 


a. pyrenaicus Geofiroy 


. 26 


6. rufulus Graells . 


. 26 


Family SORICIDiE . . . 


. 28 


1. Sorex Linnseus 


. 29 


1. araneus Linnaeus . 


. 31 


a. araneus Linnaeus 


. 35 


b. castaneus Jenyns 


. 37 


c. santonus Mottaz 


. 40 


d. euronotus Miller 


. 41 


e. bergensis Miller . 


. 41 


/. tetragonurus Hermann 


. 42 


g. pyrenaious Miller 


. 44 


h, fretalis Miller 


45 


i. granarius Miller . 


52 


2. minutus Linnaeus . 


53 


a. minutus Linnaeus . 


55 


b. luoanius Miller 


60 


3. alpinus Schinz . 


60 


a. alpinus Schinz 


62 


6. heroynious Miller 


63 


2. Neomys Kaup .... 


65 


1. fodiens Sohreber 


66 


a. fodiens Sohreber. 


69 


b. bioolor Shaw . 


73 


2. milleri Mottaz . 


78 


3. anomalus Cabrera . 


'81 


3. Paohyura de Selys-Long 




champs 


81 


1. etrusca Savi. 


83 


4. Crocidura Wagler. 


86 


1. leucodon Hermann 


88 


2. mimula Miller . . . . 


94 


a. mimula Miller . 


95 


6. iculisma Mottaz . 


98 


c. cantabra Cabrera 


99 



PAGE 

3. russula Hermann ... 99 

a. russula Hermann . . 101 

b. pulchra Cabrera . . . 103 

c. eintrae Miller. . . . 108 

4. sicula Miller .... 108 

5. caneae Miller .... 109 

6. caudata Miller .... 110 

7. cyrnensis Miller . . . Ill 

8. balearica Miller . . . 112 

Family ERINACEIDiE ... 114 

1. Erinaceus Linnaeus . . . 114 

1. europaeus Linnaeus . . . 115 
a. europaeus Linnaeus . . 120 
6. hispanicus Barrett- 
Hamilton 122 

c, italicus Barrett-Hamil- 
ton .. . ... 123 

d. consolei Barrett-Hamil- 
ton 126 

2. roumanicus Barrett- 

Hamilton 127 

3. nesiotes Bate .... 129 

4. algirus Duvernoy and 

Lereboullet .... 130 

a. algirus Duvernoy and 
Lereboullet .... 131 

b. vagans Thomas . . . 133 



ORDER CHIR0PTERA. 

Sub-Obdeb microchiroptera. 

Family RHINOLOPHEDiE . . 136 

1. Rhinolophus Lacepede . . 137 

1. ferrum-equinum Sohreber 139 
a. ferrum-equinum Sohre- 
ber 142 

6. insulanus Barrett- 
Hamilton 147 

2. hipposideros Bechstein . 147 
a. hipposideros Bechstein . 149 
6. minimus Heuglin . 
c. minutus Montagu . 

3. euryale Blasius . 

4. mehelyi Matsohie . 



151 
154 
155 
159 
5. blasii Peters 162 



SYSTEMATIC INDEX 



Order Chieoptera — Sub - Order 

MlCROCHIEOPTEBA — COlltd. 

Family VESPERTILIONIDiE . 165 

SUB-FAMILY VeSPEBTILIONIM! . 

1. Myotis Kaup . . 

1. mystacinus Kuhl 

2. nattereri Kuhl . 

3. emarginatus Geoffroy . 

4. bechsteinii Kuhl 

5. daubentonii Kuhl . 

6. capaccinii Bonaparte . 

7. dasyoneme Boie 

8. myotis Borkhausen 

9. oxygnathus Montioelli 

2. Pipistrellus Kaup 

1. pipistrellus Schreber . 

2. nathusii Keyserling and 

Blasius 

3. kuhlii Kuhl . . . . 

4. savii Bonaparte . 

3. Eptesious Rafinesque 

1. serotinus Schreber 

2. sodalis Barrett-Hamilton. 

3. nilssonii Keyserling and 

Blasius 

4. Vespertilio Linnaeus . 
1. murinus Linnaeus . 

5. Nyctalus Bowdich 

1. maximus Patio. 

2. nootula Schreber 

3. leisleri Kuhl 

4. azoreum Thomas 

6. Pleootus Geoffroy . 
1. auritus Linnaeus 

7. Barhastella Gray . 
1. barbastellus Schreber 

Sub-Family Miniopteein^j: 
1. Miniopterus Bonaparte 
1. schreibersii Kuhl 

Family MOLOSSIDiE . 

1. Nyctinomus Geoffroy 

1. teniotis Rafinesque 



ORDER CARNIVORA. 



165 
166 
169 

174 
177 
179 
184 
187 
189 
192 
199 
202 
204 

213 
215 
219 
224 
226 
231 

234 
238 
238 
242 
244 
245 
252 
254 
256 
256 
263 
263 

268 
208 
269 

276 
276 
277 



Family URSID-&! . 


284 


1. Ursus Linnaeus 


285 


1. arctos Linnaeus . 


285 


2. Thalarctos Gray . 


297 


1. maritimus Phipps . 


298 


Family CANEDiE . . . 


303 


1. Oanis Linnaaus 


304 


1. lupus Linnaeus . 


305 


a. lupus Linnaeus . 


313 


b. signatus Cabrera 


314 


c. deitanus Cabrera 


315 


2. aureus Linnaeus 


315 



paob 

2. Alopex Kaup . . . 318 

1. lagopus Linnasus . . . 319 

2. spitzbergenensis Barrett- 

Hamilton and Bonhote . 324 

3. Vulpes Oken 325 

1. vulpes Linnaeus . . . 326 

a. vulpes Linnaeus . . . 330 

b. crucigera Bechstein . . 331 

c. silacea Miller .... 333 

2. ichnusae Miller . . 336 

Family MUSTELID^l ... 340 

Sub-Family Melin.33 . 341 

1. Meles Brisson. . . ". 341 

1. meles Linnaeus . . 343 
a. meles Linnaeus . . . 348 
6. marianensis Graells. . 352 

2. arcalus Miller .... 352 
Sub-Family Luteins .... 354 

1. Lutra Brisson . . . . 354 

1. lutra Linnaeus . . 355 

Sub-Family Mustelines . . . 364 

1. Martes Pinel ... .365 

1. martes Linnseus . . 366 
a. martes Linnaeus . . 372 
6. latinorum Barrett-Ha- 
milton 373 

2. foina Erxleben . . .374 

a. foina Erxleben . . . 375 

b. mediterranea Barrett- 
Hamilton 380 

3. bunites Bate . . . 380 

2. Mustela Linnaeus. . . . 381 
Sub-genus Mustela Linnaeus . 384 

1. erminea Linnasus . . . 385 

a. erminea Linnaeus . » . 387 

b. sestivaKerr .... 389 

c. stabilis Barrett-Hamil- 
ton 390 

d. ricinae Miller . . .397 

2. hibernica Thomas and 

Barrett-Hamilton . . 398 

3. nivalis Linnaeus . . 401 

a. nivalis Linnaeus . . . 402 

b. boccamela Bechstein . 405 
u. iberica Barrett-Hamilton 407 

4. africana Desmarest . . 412 

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

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

7. putorius Linnaeus . . 419 

a. putorius Linnseus . . 423 

b. aureolus Barrett-Hamil- 

ton 425 

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

Sub-Family Guloninje . . 432 

1. Gulo Storr 433 

1. gulo Linnaeus . . 434 



SYSTEMATIC INDEX 



Order Carnivora — continued. 

Family VIVERRIDjE . . . 

1. Mungos Geoffroy and Cuvier 
1. widdringtonii Gray 

2. Genetta Oken . 

1. genetta Linnaeus . 
a. genetta Linnaeus 
6. balearica Thomas 
c. rhodanica Matsehie 



Family FELID^l . . . 

1. Felis Linnaeus. 

1. silvestris Sohreber . 
a. silvestris Sohreber 
6. grampia Miller . 
c. tartessia Miller 

2. sarda Lataste . 

3. agrius Bate . 

2. Lynx Kerr .... 

1. lynx Linnaeus . 

2. pardellus Miller 



440 
440 
441 
446 
447 
451 
452 
452 

455 
456 
457 
462 
464 
465 
468 
470 
470 
471 
475 



ORDER RODENTIA. 
Sub-Obdbb duplicidentata. 

Family LEPORIDiE .... 484 

1. Oryetolagus Lilljeborg . . 484 
1. eunioulus Linnaeus . 485 

a. eunioulus Linnaeus . . 490 

o. huxleyi Haeokel . . . 491 

2. Lepus Linnaeus .... 495 

1. europaeus Pallas . . 498 

a. europaeus Pallas. . . 502 

b. oooidentalis de Winton. 504 

c. pyrenaious Hilzheimer . 506 

d. meridiei Hilzheimer . 506 

e. oorsioanus de Winton . 507 
/. hybridus Desmarest . . 508 
g. transsylvanicus Mat- 
sehie 509 

2. cretious Barrett-Hamilton 512 

3. mediterraneus Wagner . 513 

4. granatensis Rosenhauer . 515 
a. granatensis Rosenhauer 516 
6. gallaecius Miller ... 517 

c. iturissius Miller . . . 518 

d. parnassius Miller . . 519 

5. timidus Linnaeus . . . 522 

a. timidus Linnaeus . 526 

b. varronis Miller . . . 528 

c. seotious Hilzheimer. . 529 

6. hibernicus Bell. . . . 531 

Sub-Oedeb SIMPLICIDENTATA. 

Family ZAPODIDiE .... 535 



Sub-Family Sicistin.^ 
1. Sicista Gray . 

1. loriger Nathusius 

2. trizona Petenyi . 



536 
536 
537 
539 



Family HYSTRICIDiE 

1. Hystrix Linnaeus .... 
1. cristata Linnaeus . 

Family MUSCARDINIDiE . . 

1. Eliomys Wagner .... 

1. queroinus Linnaeus 

2. gymnesieus Thomas . 

3. pallidus Barrett-Hamilton 

4. sardus Barrett-Hamilton . 

5. lusitanious Reuvens . 

2. Dyromys Thomas. 

1. nitedula Pallas . 

a. nitedula Pallas . 

6. intermedius Nehring . 

c. wingei Nehring . 

2. robustus Miller. 

3. Glis Brisson 

1. glis Linnaeus .... 

a. glis Linnaeus .... 
6. italious Barrett-Hamil- 
ton 

c. melonii Thomas . 

d. pyrenaicus Cabrera . 

4. Muscardinus Kaup . 

1. avellanarius Linnaeus . 

2. pulcher Barrett-Hamilton 

Family MURID.S3 .... 

Sub-Family Cbicetin^; . 

1. Crioetulus Milne-Edwards . 
1. atticus Nehring . . 

2. Crieetus Leske .... 
1. erioetus Linnaeus . 

a. crieetus Linnaeus 

b. oanesoens Nehring . 

c. nehringi Matsohie . 

3. Mesooricetus Nehring . 
1. newtoni Nehring . 

Sub-Family Micbotinje 

1. Myopus Miller .... 
1. schisticolor Lilljeborg. 

2. Lemmas Link 

1. lemmus Linnaeus . 

3. Evotomys Coues . 

1. glareolus Sohreber . 
a. glareolus Sohreber 
6. britannious Miller 

sueoious Miller . 

isterious Miller . 

norvegious Miller 

vaseoniae Miller . 

helvetious Miller 
h. nageri Sohinz 
i. hallucalis Thomas 

2. skomerensis Barrett 
milton .... 

3. oaesarius Miller . 

4. rutilus Pallas . 

5. rufooanus Sundevall 



c. 

d. 
e. 
/■ 



Ha- 



paoe 
542 
542 
543 

549 
550 
551 
558 
559 
560 
560 
566 
567 
568 
569 
570 
572 
572 
573 
577 

578 
579 
582 
583 
583 
590 

591 
592 
593 
593 
596 
597 
602 
603 
605 
605 
606 

610 
611 
611 
614 
615 
623 
626 
632 
634 
636 
637 
638 
639 
640 
641 
643 

644 
645 
646 
648 



SYSTEMATIC INDEX 



Order Kodentia — Sub - Order 
Simplictdentata — Family 
Mubid-e — Sub-Pamily Mi- 
cbotin-se —continued. 

4. Miorotus Schrank . . . 658 
Sub-Genus Miorotus Schrank 659 

1. agrestis Linnaeus . . . 662 

a. agrestis Linnseus . . 668 

b. exsul Miller .... 669 

c. levernedii Crespon . . 671 

d. bailloni de Selys-Long- 
ohamps 672 

e. airbus Bellamy . . 673 
/. negleotus Jenyns . . . 675 
g. rozianus Bocage . . . 680 

2. arvalis Pallas . . . 681 

a. arvalis Pallas . . . 683 

b. meridianus Pallas . . 686 

c. duplioatus Rorig and 
Borner .... 686 

d. levis Miller .... 687 

3. incertus de Selys - Long- 

champs 690 

4. asturianus Miller . . . 693 

5. orcadensis Millais . . . 694 

6. sandayensis Millais . . 696 

a. sandayensis Millais . . 697 

b. westrse Miller . 698 

7. sarnius Miller , . . 700 

8. eabrerse Thomas . . . 701 

9. dentatus Miller. ... 703 

10. hartingi Barrett-Hamilton 704 

11. angularis Miller . . . 706 

12. ratticeps Keyserling and 
Blasius. . ... 708 

Sub-Genus Chionomys Miller 712 

13. nivalis Martins ... 713 

a. nivalis Martins . . . 716 

b. aguitanius Miller . . 717 

14. lebrunii Orespon . . 718 
a. lebrunii Crespon. . 719 
o. leueurus Gertie . . 722 

15. ulpius Miller ... 723 

5. Arvicola Lacepede . . . 723 

1. amphibius Linnseus . . 725 

a. amphibius Linnaeus . 730 

b. reta Miller . 732 

2. sapidus Miller . . . 732 

a. sapidus Miller . . . 733 

b. tenebrieus Miller . . 735 

3. terrestris Linnseus. . 738 

4. italicus Savi 740 

5. illyricus Barrett-Hamilton 741 

6. musignani de Selys-Long- 

champs . . . 744 

7. scherman Shaw . 744 
a. scherman Shaw . . 745 
6. exitus Miller .... 746 

c. monticola de Selys- 
Longchamps .... 749 



6. Pitymys McMurtrie . . . 


752 


1. subterraneus de Selys- 




Longchamps .... 


755 


a. subterraneus de Selys- 




Longchamps .... 


758 


6. capucinus Miller 


760 


2. daeius Miller 


760 


3. druentius Miller 


762 


4. fatioi Mottaz .... 


763 


5. multiplex Patio 


764 


6. savii de Selys-Longehamps 


768 


7. nebrodensis Mina-Palumbo 


770 


8. pyrenaicus de Selys-Long- 




champs 


770 


a. pyrenaicus de Selys- 




Longchamps .... 


771 


6. brunneus Miller . 


772 


9. planiceps Miller . 


772 


10. gerbii Gerbe .... 


773 


11. lusitanicus Gerbe. 


776 


12. marise Major .... 


777 


13. pelandonius Miller . 


778 


14. depressus Miller 


779 


15. ibericus Gerbe 


780 


a. ibericus Gerbe . 


782 


6. centralis Miller 


782 


c. pascuus Miller . 


783 


d. regulus Miller 


784 


16. duodecimcostatus de 




Selys-Longchamps 


784 


17. provincialis Miller 


785 


18. thomasi Barrett-Hamil- 






786 


19. atticus Miller . 


787 


Sub-Family Musing . 


791 


1. Apodemus Kaup . 


791 


1. epimelas Nehring . 


794 


2. sylvaticus Linnseus 


797 


a. sylvaticus Linnseus . 


803 


6. callipides Cabrera 


809 


c. dichrurus Rafinesque . 


810 


d. creticus Miller . 


813 


3. hebridensis de Winton 


824 


4. hirtensis Barrett-Hamil- 




ton .... 


825 


5. fridariensis Kinnear . 


825 


6. flavicollis Melchior 


828 


a. flavicollis Melchior . 


829 


b. wintoni Barrett-Hamil- 




ton . 


831 


7. agrarius Pallas .... 


836 


2. Micromys Dehne . 


840 


1. minutus Pallas . 


841 


a. soricinus Hermann . 


844 


6. pratensis Ockskay . 


846 


3. Epimys Trouessart . 


848 


1. rattus Linnseus . 


849 


a. rattus Linnseus . 


853 


6. alexandrinus Geofiroy . 


854 


2. norvegicus Erxleben . 


858 



SYSTEMATIC INDEX 



Order Rodentia — Sub - Order 
Sihplicidentata — Family 
Mubid^; — Sub-Family Mu- 
BiNai — continued. 

4. Mus Linnaeus 863 

1. musculus Linnaeus . . 865 
a. musculus Linnaeus . . 869 
6. azorious Schinz . . . 871 

2. muralis Barrett-Hamilton 874 

3. faeroensis Clarke . . . 875 

4. spioilegus Petenyi . . 877 
a. spioilegus Petenyi . 878 
6. hispanicus Miller . . 879 
c. lusitanious Miller . 882 

5. Aoomys Geoffroy . . . 883 
1. minous Bate .... 883 



Family SPALACIDJE ... 887 

1. Spalax Gueldenstaedt . . 887 

1. dolbrogeae Miller ... 889 

2. hungaricus Nehring . . 894 

3. graeous Nehring . . . 895 



Family SCIURIDiE .... 897 

1. Sciurus Linnaeus .... 898 
1. vulgaris Linnaeus . . . 898 

a. vulgaris Linnaeus . . 905 

6. varius Gmelin . . . 906 

c. leucourus Kerr . . . 907 

d. russus Miller. ... 909 

e. fuscoater Altum . . . 910 
/. italious Bonaparte . . 912 
g. lilaeus Miller .... 913 
h. alpinus Desmarest . 914 
i. numantius Miller . . 914 
j. infuscatus Cabrera . 916 
k. segurae Miller . . . 917 
I. baetious Cabrera . . . 923 

2. Citellus Oken 924 

1. citellus Linnaeus . . . 924 

2. suslica Gueldenstaedt . . 929 

3. Marmota Blumenbach . . 931 

1. marmota Linnaeus . . 932 

2. bobak Miiller .... 937 



Family PETAUBJSTID^ . . 940 

1. Sciuropterus F. Cuvier . 941 

1. russicus Tiedemann . . 941 



Family CASTORIDJE ... 947 

1. Castor Linnaeus .... 947 

1. fiber Linnaeus .... 947 



OBDEB UNGULATA 

PAGE 

Family SUIDjE 956 

1. Sus Linnaeus 956 

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

2. meridionalis Major . . 960 



Family CERVIDiE .... 

1. Cervus Linnaeus . 

1. elaphus Linnaeus . 

a. germanicus Desmarest . 

b. elaphus Linnaeus 

c. atlanticus Lonnberg 

d. scoticus Lonnberg . 

e. hispanicus Hilzheimer . 
/. corsicanus Erxleben. 

2. Dama Hamilton Smith 
1. dama Linnaeus . 

3. Gapreolus Gray . 

1. capreolus Linnaeus . 

a. capreolus Linnaeus . 

b. transsylvanicus Matschie 

c. canus Miller .... 

d. thotti Lonnberg. 

4. Alces Gray 

1. alces Linnaeus .... 

5. Bangifer Hamilton Smith . 

1. tarandus Linnaeus . 

2. f ennicus Lonnberg . 

3. platyrhynchus Vrolik . 

Family BOVID.-E .... 

1. Ovis Linnaeus 
1. musimon Pallas 

2. Capra Linnaeus . 

1. ibex Linnaeus 

2. pyrenaica Schinz . 

a. pyrenaica Schinz 

b. lusitanica Franca . 

c. victoriae Cabrera. 

d. hispanica Schimper . 

3. aegagrus Erxleben 

3. Rupricapra Blainville 

1. rupicapra Linnaeus 

2. ornata Neumann . 

3. pyrenaica Bonaparte . 

4. parva Cabrera 



962 
963 
964 
965 
967 
967 
968 
969 
969 
970 
970 
972 
973 
974 
975 
975 
975 
976 
978 
979 
980 
981 
9S5 

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



APPENDIX. 

Crocidura ichnusae Festa . . . 998 
Evotomys glareolus norvegicus — 

cranial measurements . . . 999 
Spalax — Forms recognized by 

Mehely 1000 



CATALOGUE 

OP THE 

LAND-MAMMALS OF WESTERN 
EUROPE. 



I 



Order INSECTIVORA. 

1827. Insectivora Gray, Griffith's Guvier, Anim. Kingd., v, p. 100. 

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

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

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

KEY TO THE EUROPEAN FAMILIES OP INSECTIVORA. 

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

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

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

mandibular articulation double ; zygoma absent ; 

floor of brain-case with large lateral vacuities ; 

no auditory bulla ; general form mouse-like, the 

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

B 



£ INSECTIVORA 

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

Desmans) Talpidse, p. 2. 

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

Family TALPIDEE. 

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

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

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

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

Sub-Family TALPINTE. 

Geographical distribution. — Temperate portions of Europe and 
Asia, from England to Japan : in Europe south to the Mediter- 
ranean coast. 



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

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

Genus TALPA Linnaeus. 

1758. Talpa Linnseus, Syst. Nat., i, 10th ed., p. 52. 

Type species. — Talpa europsea Linnseus. 

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

Characters. — Dental formula : i ,-Ej, c j5j, pin jtj, m ^ 3 = 44. 
Upper premolars small, distinctly spaced, showing no tendency 
to become imbricated. Lower canine slightly but evidently 
differentiated from incisors in size and form ; auditory bulla very 
slightly inflated, its outlines usually indistinct, the meatus small, 
sub-circular ; external form strictly talpine ; ear-conch absent ; 
eye minute, often covered by the skin. 

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

KEY TO THE EUROPEAN SPECIES OE TALPA. 

Greatest diameter of m 1 about 4 mm. ; three lower 
molars together about 8 mm. (vicinity of 

Borne) T. romana, p. 18. 

Greatest diameter of m l about 3 mm. ; three lower 
molars together about 7 mm. or less. 
Oondylobasal length of skull 33 to 37 mm. (dis- 
tribution general) T. europsea, p. 3. 

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

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

Posterior border of anteorbital foramen over 

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

TALPA EUROP.ffi:A Linnseus. 

1758. [Talpa} europsea Linnaeus, Syst. Nat., i, 10th ed., p. 52 (Sweden). 
1766. Talpa frisius P. L. S. Miiller, Natursyst. Suppl. u. Regist.-Band, 

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

i, p. 50 (Renaming of europsea). 

B 2 



4 INSECTIVOEA 

1777. [Talpa europsea] a albo-maculata Erxleben, Syst. Begni Anim., I, 

p. 117 (Ostfriesland). 
1785. [Talpa] vulgaris Boddaert, Elenohus Anim., i, p. 126 (Europe). 
1789. [Talpa europsea] |8 variegata Gmelin, Syst. Nat., i, 13th ed., p. 110 

(Sweden). 
1789. [Talpa europsea] y alba Gmelin, Syst. Nat., I, 13th ed., p. HO 

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

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

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

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

Heft v-viu, p. 13 (Southern Prance). 
1836. Talpa europica flavescens Reichenbach, Praeht.-gemoinn. der 

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

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

In- und Auslandes, IV, p. 336 (Germany). 
1857. Talpa europsea Blasius, Saugethiere Deutschlands, p. 109. 
1869. Talpa europsea, flavescens Pitzinger, Sitzungsber. kais. Akad. 

Wissensch. Wien, Math.-Naturwiss. Classe, dix, pt. i, p. 400. 
1869. Talpa europsea, maculata Pitzinger, Sitzungsber. kais. Akad. 

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

(Renaming of albo-maculata). 
1869. Talpa europsea 5 grisea Pitzinger, Sitzunsgber. kais. Akad. Wissensch. 

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

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

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

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

Type locality. — Upsala, Sweden. 

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

Diagnosis. — Size medium (hind foot about 18 mm., condylo- 
basal length of skull rarely less than 33 mm.) ; skull with orbit 
rather long, so that distance from posterior border of orbit to 
posterior border of anteorbitai foramen is about equal to that 
from latter point to front of first premolar or back of canine ; 
teeth of moderate size, the length of upper tooth-row (exclusive 
of incisors) less than 14 mm., the greatest diameter of ro 1 about 
3 mm. ; mesostyle of m 2 and m 3 entire or with apex slightly 
notched. 

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



about 12 mm. long throughout body, shorter on head. Muzzle 
pad well developed, slightly wider than high, its entire margin 
free, the upper edge folded back and with a deep narrow median 
emargination, the whole surface finely and evenly rugose. 
Behind pad the skin of the muzzle is naked and wrinkled along 
median line above, the exact size and form of the naked area 
varying in different individuals, but its length usually about 
7 mm., its breadth anteriorly equal to greatest diameter of pad, 
that posteriorly somewhat less. Under side of upper lip with 
deep median groove extending forward to muzzle-pad. Eyes 
excessively minute, less than 1 mm. in diameter, in some 
individuals completely covered by the integument, in others with 
a pin-hole aperture.* No external ear, the meatus about 2 ■ 5 mm. 
in diameter. Legs so short that scarcely more than the feet 
project beyond general integument of body. This is particularly 
true of the front legs, which are entirely concealed to wrist. 
Front feet very large, the palms orbicular, permanently turned 
outward, their surface naked, finely and evenly tuberculo-reticu- 
late, without trace of pads. Toes, five, each armed with a long, 
slightly-curved nail broadly grooved along under surface, that of 
fifth digit not so large as the others. Third digit longest, second 
and fourth sub-equal and slightly shorter, first and fifth slightly 
shorter than second and fourth. Hind foot short, somewhat 
triangular in outline, broad through base of toes but narrowing 
rapidly toward heel, the five toes with well developed, curved, 
but not flattened claws, much less enlarged than those of front 
foot ; second, third and fourth digits sub-equal and longest, fifth 
and first successively shorter, soles naked, reticulate, with 
five rudimentary tubercles at bases of digits, and a fifth near 
middle of inner margin, its anterior extremity projecting so as to 
suggest a supplemental (clawless) toe. Dorsum of pes thinly 
clothed, the hairs at its edge forming a slight fringe. Tail 
thickened and fleshy, about 1^ times as long as hind foot, 
subterete but somewhat compressed, much constricted basally ; 
scales arranged in rather irregular rings, of which there are 
about twelve to the centimeter at middle ; hairs of tail sparse, 
not concealing rings, sometimes forming a thin pencil. Mammae : 
p2-2; »2-2 = 8. 

Colour. — Fur everywhere dark slaty grey, sometimes almost 
blackish, at others more nearly a dark smoke-grey, the hairs 
everywhere witb a noticeable metallic or purplish iridescence ; 
underparts sometimes a little less dark than back, and occa- 
sionally with a yellowish brown suffusion ; tail concolor with body. 

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

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



INSBCTIVOEA 



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

squarely truncate, its anteropos- 
terior diameter about one-third 
transverse diameter. Its anterior 
and lateral sutures remain visible, 
but the lambdoid suture is nearly 
obliterated early in life, though 
its position is usually marked by 
a slight ridge representing the 
lambdoid crest. Base of brain- 
case smooth, without conspicuous 
ridges, furrows or open spaces, 
the bones for the most part some- 
what inflated ; two ill-defined pits 
in surface of basioccipital in front 
of foramen magnum ; a shallow, 
broadly triangular median furrow 
between the low, flattened bullae, 
the sub-circular, slightly triangu- 
lar outline of which is sometimes 
distinctly indicated, but more 
often very obscure ; auditory 
meatus small, nearly circular. 
Inner pterygoid plate small but well developed, the hamulars 
short, turned slightly outward ; mesopterygoid space much 
narrower posteriorly than anteriorly, its greatest breadth less 
than half length, its length much more than width of palate 
between posterior molars, its anterior border broadly rounded ; 
region outside pterygoid plate inflated and with low but distinct 
longitudinal ridge, convergent anteriorly with that forming edge 
of mesopterygoid space. Interorbital region sub-cylindrical, dis- 
tinctly expanded at middle. Rostrum narrower and somewhat 
abruptly lower than interorbital region, its narrowest point just 
behind canines ; nares with evident posterior emargination. 
Anteorbital foramen moderately large, its posterior border over 
metastyle of to' 2 , the plate forming outer wall of canal usually 
much narrower than foramen Lachrymal foramen above anteor- 




¥10. I. 
Talpa, europsedf. Nat. size. 



bital foramen and slightly in front of its middle ; its orifice over 
metastyle of in 1 . Temporal fossa rather large, the distance from 
its posterior border to posterior border of anteorbital foramen 
equal to that from latter point to front of first premolar or back 
of canine. Palate moderately broad, without special peculiarities 
of form, terminating posteriorly, a little behind m 3 , in a slightly 
raised crescentic ridge ; a vacuity about as large as the minute 
incisive foramen on each side of palate opposite space between 
m 1 and m 2 . Mandible rather slender, the ramus curved downward 
near middle and upward posteriorly, the angular process nearly 
on level with alveolar line. Coronoid process large, its broadly 
rounded off extremity rising considerably above articular level, 
its height above alveolar line about equal to least breadth of 
posterior segment of mandible. Articular process slender, the 
single articular surface small, normal in position. Angular 
process slightly longer than articular process, and distinctly 
broader, forming an obliquely-set plate directed almost horizon- 
tally backward, the extremity slightly hooked upward. 

Teeth. — In proportion to the size of the skull the teeth are 
moderately large, their general 
aspect noticeably trenchant. Upper 
incisors simple, chisel - shaped, 
perpendicular, forming a strongly 
convex row between canines, their 
height equal, but their breadth 
diminishing regularly from first to 
third ; crowns of first and second 
usually in contact, that of third 
separated by a slight space from 
second and from canine. 
Lower incisors projecting 
somewhat obliquely for- 
ward, essentially similar to 
the upper teeth in form, 
but smaller and narrower. 
Upper canine large, two- 
rooted,* the height of the 
shaft greater than that of any 
of the other teeth, and fully 
equal to anterior breadth of 
palate ; shaft wider ante- 
riorly than posteriorly, with 
antero-internal longitudinal 
groove, and highly deve- 
loped, slightly concave pos- 
terior cutting edge. Lower 
fourth incisor, but with shaft conical instead of chisel-shaped, 

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




t'IG. 2. 
Talpa europvea. Teeth X 5. 

canine very small, resembling a 



INSECTIVORA 

and set in the jaw at a slightly different angle posterior 
surface of shaft with well developed longitudinal ridge. First, 
second and third upper premolars two-rooted, small, their 
points on level with those of incisors and inner cusps of 
molars, their crowns separated from each other as well 
as from canine and large premolar by narrow equal spaces ; 
crowns rather higher than long, compressed, with slightly 
developed posterior cutting edge, narrowly triangular in outline 
when viewed from the side, the first more slender than the 
others, the second and third with slight though evident postero- 
external angle on cutting edge. Fourth premolar essentially 
like the other three in form, but much larger and three-rooted, 
the length of crown greater in proportion to height ; no secondary 
cusps. Lower premolars two-rooted, similar to the upper teeth 
in form, the second and third the same size as smaller upper 
premolars, the first nearly as large as pm i , but narrower, 
owing to absence of third root, the fourth intermediate. Upper 
molars with crowns much wider externally than internally, 
and completely divided into two unequal sections by a deep 
longitudinal groove passing between protocone and bases of 
paracone and metacone. Protocone large, its posterior com- 
missure extending in line parallel to sagittal plane, and ending 
abruptly at posterior edge of crown, near which it is slightly 
thickened, its anterior commissure similar though shorter, but 
usually showing some trace of thickening,* particularly in m 2 . 
Paracone and metacone sub-equal in m 2 and m 3 , the metacone 
the larger in the former, the paracone in the latter. In m 1 the 
metacone is about double the size of paracone, and is the 
largest and highest cusp of the upper molar series, its posterior 
cutting edge and long commissure functioning with similarly 
enlarged protoconid of m 2 . Styles and outer commissures well 
developed in m 2 and forming a distinct VV"P a ttern ; mesostyle 
entire or with apex slightly notched. In m 1 the parastyle is 
reduced to a minute though usually evident cusplet on the 
cingulum, and the mesostyle to a thickening or angle in 
commissure connecting the two main cusps ; metastyle well 
developed.f In m 3 the parastyle, mesostyle and their commis- 
sures are well developed, metastyle and fourth commissure 
absent ; mesostyle with apex usually bifid. Middle lower molar 
largest, its protoconid the highest cusp in the series. First lower 
molar broader posteriorly than anteriorly, second and third 
slightly broader anteriorly than posteriorly, the two triangles 
essentially alike in form. Metaconid of m 1 low, scarcely more 

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

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



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

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

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

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

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

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

Belgium: Waremme, Liege, 3 (U.S.N.M.). 

Prance : Guines, Pas-de-Calais, 1 ; Pont-de-Briques, Pas-de-Calais, 1 ; 
Trinity, Jersey, 2; St. Lawrence, Jersey, 1; Barhizon, Seine-et-Marne, 2; 
Melun, Seine-et-Marne, 1 (Mottaz) ; Lignieres, Charente, 1 (Mottazl ; 
Huelgoat, Brittany, 1 ; Cadillac-sur-Garonne, Gironde, 1 (U.S.N.M.) ; 
Poret de Bouconne, Gers, 13; Solferino, Landes, 10; Caterille, Haute- 
Garonne, 10; Legouvin, Haute-Garonne, 3; Luchon, Haute-Garonne, 3; 
Bareges, Hautes-Pyrenees, 5; l'Hospitalet, Ariege, 1; Porte, Pyrenees- 
Orientales, 3; St. Gilles, Gard, 1; Valescure, Var, 1; Agay, Var, 1; 
Etupes, Doubs, 3 (Mottaz) ; Lucinges, Haute-Savoie, 9 ; Montauban, 
Haute-Savoie, 15 ; Cranves-Sales, Haute-Savoie, 12. 

Spain: Pajares, Leon, 2; Castrillo de la Beina, Burgos, 9; Castauares, 
Burgos, 8 ; Lerida, 1 ; Barracas, Castellon, 24 ; Catarroja, Valencia, 4 ; 
Dehesa de Valencia, Valencia, 17. 

Germany : K6nigsberg,4 (U.S.N.M.) ; Moritzburg, Saxony, 8 (U.S.N.M) ; 
Ummerstadt, Thiiringen, 3 ; Brunswick, 15 (U.S.N.M.) ; Aachen, 9 
(U.S.N.M.); Bremen, 1; Ingelheim, Kheinhessen, 4; Strass, near Burg- 
heim, Bavaria, 7; Niesky, Silesia, 1; Kalbe, Saale, 2; Magdeburg, 
Saxony, 1 ; Strassburg, 3. 

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



10 



INSECTIVORA 



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

Bulgaria: Sofia, 1 (Andersen). 

Switzerland : Geneva, 15 (U.S.N.M. and Mottaz) ; Lausanne, Vaud, 1 
(U.S.N.M.) ; Les Plans, Vaud, 2 (U.S.N.M.) ; Chesieres, Vaud, 1 (Mottaz) 
Andermatt, Uri, 2 (U.S.N.M.) ; Mflrren, 1 ; Thurgau, 1 ; Oberhasli Valley 
1; St. Gallen, 3; Bheinthal, St. Gallen, 2; Degersheim, St. Gallen, 3 
Gossau, St. Gallen, 3; Untervatz. Grisons, 1; Breganzona, Tieino, 1 
Cortivallo, Tieino, 6 (B.M. and U.S.N.M.) ; Comano, Tieino, 2 (U.S.N.M.) 
Lugano, Tieino, 4 (B.M. and U.S.N.M.) ; Muzzano, Tieino, 2 (B.M. and 
U.S.N.M.) ; Sorengo, 1 (U.S.N.M.) ; Cremignone, Tieino, 1. 

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

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



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

rf. Stockbriggs, Lanarkshire. 

9. Parsop, Herefordshire, Eng- 
land. 

2 6. Bugby, Warwickshire. 

2. Warwickshire. 

6. Eulbourn, Cambridgeshire. 

2 al. Holloway, Somerset. 

2. Somerset. (Hilgel.) 

2 al. Cobham, Surrey. 

1 al. 1. Egham, Surrey. 

9. Knockholt, Kent. 

2 $. al. Bromley, Kent. 

3 S. Devonshire. 

2 c5. Trinity, Jersey, Channel 
Islands. (R. H. Bunting.) 

9. St. Lawrence, Jersey. 

6. Guines, Pas-de-Calais, 80 ft. 
Prance. 

S. Pont - de - Briques, Pas - de- 
Calais. 

5. Huelgoat, Brittany, 600 ft. 
2d, 2 9. Poret de Bouconne, Gers, 

250 m. (A. Robert.) 
2 <5, 2 9. Solferino, Landes. (A. Ro- 
bert.) 
2 c?, 2 9. Caterille, Haute - Garonne, 
900-1000 m. (A. Robert.) 
2 <5, 9. Luchon, Haute - Garonne. 
(A. Robert.) 
2 6, 1 9. Bareges, Hautes-Pyrenees. 

6. L'Hospitalet, Ariege. (A. 

Robert.) 
S. Porte, Pyrenees-Orientales. 
2 S Porte, Pyrenees - Orientales, 

1600-1700 m. {A. Robert.) 
i. Valesoure, Var. 



W. B. Ogilvie- 

Grant (p). 
W. B. Ogilvie- 

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

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

H. King (p). 
E. B. Alston (p). 
Dr. Leach (p). 
P. Heiss (p). 
W. Blaekwell (p). 
H. E.Bawson (p). 
Oxley Grabham 

O. Thomas (p). 

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



11. 1. 3. 62-63. 

11. 1. 3. 66. 

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

11. 1. 3. 64-65. 

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

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

8. 9. 2. 3-4. 

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



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



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



O. Thomas (p). 

O. Thomas (p). 

0. Thomas (p.) 

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

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



92. 9. 5. 1. 
6. 4. 1. 28-31. 

6. 4. 1. 32-35. 

6. 4. 1. 24-27. 

6. 4. 1. 21-23. 

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

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



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



TALPA 1 1 

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

1. Montaubah, Haute - Savoie, A. Eobert (o & p). 97. 1. 9. 1. 

1000 m. 
3 6, 1 9. Cranves-Sales, Haute-Savoie, A. Eobert (o & p). 5. 4. 4. 1. 5. 4, 

900 m. 9. 3-5. 

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

909-1200 m. Prance. (A. 

Bobert.) 
6, 9. Pajares, Leon, Spain. (N. O. Thomas (p). 8. 2. 9. 26-27. 

Gonzales.) 

2. Castrillo de la Reina, Burgos, G. S. Miller (o). 8.8.4.20-21. 

Spain. 
2 6, 5 9. Castrillo de la Reina, Burgos. N. Gonzalez (c.) 8. 7. 7. 1-7. 
<5. Lerida, Spain. (N. Gonzales.) O. Thomas (p). 8. 2. 9. 28. 
6 6, 6 9. Barraoas, Castellon. (N. Gon- O. Thomas (p). 8. 2. 9. 29-40. 

sales.) 
3 <J, 9. Catarroja, Valencia. (N. Gon- O. Thomas (p). 8.2.9.12-15. 

sales.) 
4<5, 5 9. Dehesa de Valencia, Val- O. Thomas (p). 8.2.9.17-25. 
encia. (N. GonsaUs.) 
3(5,9. Ingelheim,Rheinhessen,Ger- C. Hilgert (c). 8.11.2.11-14. 

many. 
6,9. Ummerstadt, Thiiringen. Lord Lilford (p). 11.1.1.115-116. 
(Schuchardt.) 
6. Ummerstadt, Thiiringen. Lord Lilford (p). 11.1.1.137. 
(Schuchardt.) 
6, 3 9. Strass, Burgheim, Bavaria. Lord Lilford (p). 11. 1. 1. 1-4. 

(KOrbits.) 
2 6, 9. Strass, Burgheim, Bavaria. Lord Lilford (p). 11.1.1.132-134. 
(KOrbits.) 
6. Niesky, Silesia. (W. Boer.) Dr. B. Hamilton 97. 12. 4.18. 

(*)■ 

9. Kalbe, Saale. (W. Bauer.) Lord Lilford (p). 11. 1. 1. 5. 
6. Kalbe, Saale. (W. Bauer!) Lord Lilford M. 11. 1. 1. 136. 
6. Magdeburg, Saxony. (Wolter- Lord Lilford (p). 11. 1. 1. 135. 

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

tas.) 
2. Csall6k<Sz-Somorja, Pressburg, Budapest Museum 94. 3. 1. 27-28. 

HuDgary. (p). 

2(5. Hatszeg, Hunyad, Transyl- C. G. Danford (c). 3.2.2.20-21. 

vania, Hungary. 

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

nia. (W. Dodson.) 

6. Comana, Vlasoa, Roumania. Lord Lilford (p). 4. 4. 6. 14. 
1. Miirren, Switzerland. W. Gartner (p). 92. 10. 5. 3. 
9. Thurgau, 400 m. Switzerland. O. Thomas (p). 4. 4. 5. 26. 

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

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

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

land. (E. H. Zollikofer). 

5. Cortivallo, Ticino. (E. H. 0. Thomas (p). 2. 8. 4. 15. 

Zollikofer). 
9. Lugano, Ticino, 300 m. 0. Thomas (c & p). 2. 7. 1. 1. 

6. Muzzano, Ticino. (E. H. 0. Thomas (p). 2. 8. 4. 16. 

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



12 



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TALPA CMCk Savi. 



1822. Talpa cseca Savi, Nuovo Giom. de' Letterati, Pisa, I, p. 265. 
1857. Talpa cmca Blasius, Saugethiere Deutsehlands, p. 115. 

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

Univ. di Torino, xxi, No. 530, p. 1, June 22, 1906. 
1910. Talpa cseca Trouessart, Paune Mamm. d'Europe, p. 63. 

Type locality. — Mountains near Pisa, Italy. 

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

Diagnosis. — Smaller than Talpa europsea (condylobasal length 
of skull, 29-32 mm. instead of 33-37 mm.) and skull with 
narrower rostrum and palate (breadth of rostrum over canines 
about 4 mm. instead of 4 ■ 6-5 mm.) ; temporal fossa shortened, the 
distance from its posterior margin to posterior margin of anteor- 
bital foramen about equal to distance from latter point to second 
premolar ; anteorbital foramen large, its posterior border over 
front of last molar, the plate forming outer wall of canal a 
slender, terete thread ; teeth small, the length of upper tooth-row 
(exclusive of incisors), 11-12 mm. ; mesostyle of m l and m 3 with 
apex deeply notched (this character not visible in specimens with 
much worn teeth). 

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

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

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

<?. Agmizzo, Ticino, Switzerland. 0. Thomas (p). 2. 8. 4. 18. 

(E. H. Zollikofer.) 
{. Stabio, Ticino. (E. H. Zol- 0. Thomas (p). 2. 8. 4. 19. 
likofer.) 
6, ¥ al. Italy. Dr. Riippell (c). 45. 7. 22. 32-33. 

TALPA OCCIDENTALS Cabrera. 

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

xx, p. 212, September, 1907. Type in Cabrera collection. 
1907. Talpa cseca occidentalis Cabrera, Bol. Ileal. Soc. Espafl. Hist. Nat., 

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

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

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



16 



INSECTIVORA 




Diagnosis. — Like Talpa caeca but skull rather robust, the 
breadth of rostrum over roots of canines usually more than 
4 mm. ; anteorbital foramen contracted, its posterior border over 
mesostyle of m 2 , the plate forming outer wall of canal usually as 
wide as transverse diameter of foramen ; crowns of molars 
appreciably enlarged (length of upper tooth-row, exclusive of 
incisors, frequently more than 13 mm.), but dentition otherwise 
as in Talpa caeca. 

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

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

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

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




Fig. 3. 
Talpa cseca (upper figure), and 
T. occidentalis (lower figure). 
> T at. siz2. 



1 al. Galicia, Spain. 
2 S, 2 Sal. La Granja, Segovia, 
Spain. 
Skull. La Granja, Segovia. 
2 S, 1 al. Cintra, Portugal. 
1 skull. 



Prof.Seoane (c&p). 94.1.1.23. 
M.delaEscalera(c). 8. 7. 30. 19-22. 



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



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



TALPA 



17 



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18 



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TALPA ROMANA Thomas. 

1902. Talpa romana Thomas, Arm. and Mag. Nat. Hist., 7th ser., x, p. 517, 

December, 1902. Type in British Museum. 
1904. Talpa romana Camerano, Mem. Reale Aeoad. Soi. di Torino, 2nd ser., 

liv, p. 81. 
1910. Talpa romana Trouessart, Paune Mamm. d'Europe, p. 64. 

Type locality. — Ostia, Rome, Italy. 

Geographical distribution. — Vicinity of Rome, Italy. • 

Diagnosis. — Externally similar to Talpa europeea but a trifle 
larger ; skull and dentition more robust tban in the related 
animal (greatest diameter of m 1 about 4 mm. ; length of upper 
tooth-row, exclusive of incisors, 14 to .15 mm.) ; mesostyle of all 
three upper molars bifid at tip. 

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





fig. 4. 
Crown of molars in Talpa europsea (a\ and T. romana (b). X 5. 



these four teeth about 10 mm. instead of about 8 mm.). In form 
the teeth are similar to those of the related animal, but the 
mesostyle of m 1 is relatively as well as actually larger, and its tip 
is distinctly bifid, though not so deeply as in the succeeding 
teeth. Cingulum between outer bases of main cusps of lower 
molars better developed than in T. europsea, its edge frequently 
forming an evident cusp, especially in m 2 . 

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



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20 



INSECTIVORA 



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

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

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

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

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

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

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

Sub-Family DESMANINJE. 

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

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

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

Genus GALEMYS Kaup. 

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

p. 118 (Mygale pyrenaica Geoff roy). 
1846. Galomys Agassiz, Nom. Zool., Index Univ., p. 149 (Emendation of 

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

(Mygale pyrenaica Geoff roy). 

Type species. — Mygale pyrenaica Geoffroy. 

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

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



GALEMYS 



21 



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

GALEMYS PYRENAICUS Geoffroy. 

(Synonymy under subspecies.) 

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

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

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

* The synonymy of this genus is as follows : — 
1777. Desmana Gueldenstaedt, " Beschaft. Berliner Gesellsoh. Naturforsch. 
Freunde, in, p. 108." (Castor moschatus Linnseus.) 

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

1800. Mygale Cuvi,er, Lee. d'Anat. Comp., i, Tabl. 1. Same type. (Described 

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

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

1830. Gaprios Wagler, Nat. Syst. Amphibien, p. 14 (Substitute for Mygale). 
1836. Myogale Brandt, Wiegmann's Archiv fur Naturgesch., 1834, i, p. 176. 

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



22 



INSECTIVORA 



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

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




fig. 5. 
Galemys pyrenaicus. Nat. size. 



GALEMYS 23 

a rudimentary dorsal and ventral keel. Mammas : pi — l,al — 1, 
* 2 - 2 = 8. 

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

Skull. — In general the skull resembles that of Talpa europeea, 
but the brain-case is shorter and squarely truncate posteriorly, 
the interorbital region is shorter and narrowed instead of 
widened at middle, and the rostrum is longer. Surface of skull 
smooth except for the same ridges as in Talpa, those at sides of 
brain-case not unusually developed. Brain-case decidedly more 
than half as deep as wide, its outline when viewed from behind 
vaguely pentagonal, its posterior margin nearly straight, though 
with slight median swelling ; condyles not projecting posteriorly, 
completely hidden when viewed from above. Interparietal 
projecting further forward than in Talpa, its anteroposterior 
diameter nearly equal to its width. Base of brain-case with 
deep but broad median furrow, the surface of the bones more 
angular and less inflated than in Talpa ; no pits in basioccipital 
in front of foramen magnum. Bullae low and flattened, less 
perfectly formed than in Talpa, the tympanic bone annular and 
retaining its distinctness, though joined with surrounding parts ; 
meatus large, occupying about one-half surface of bulla. 
Mesopterygoid space short, its length scarcely equal to width of 
palate between posterior molars, its width about one-third length, 
its anterior border double rounded, encroached on by slight 
median spine. Outer pterygoid plate reduced to a small but 
evident ridge. Interorbital region hour-glass shaped, widening 
more rapidly posteriorly than anteriorly, its narrowest region 
slightly behind middle. Zygoma straight, flattened posteriorly, 
compressed anteriorly, its length (measured from posterior border 
of orbit to posterior border of anteorbital foramen) equal to 
distance from posterior border of anteorbital foramen to front of 
canine. Rostrum about as wide as in Talpa, but relatively 
longer, the distance from posterior border of anteorbital foramen 
to gnathion about equal to greatest breadth of brain-case instead 
of much less, its dorsal surface on level with that of interorbital 
region, its outer margins nearly parallel to the squarely truncate 
anterior extremity. Nares scarcely emarginate posteriorly. 
Anterior portion of border of alveolus of large incisor distinctly 
thickened, the thickened region terminating laterally in a small 
but evident wart-like nodule. Lachrymal foramen over middle 
of anteorbital foramen and middle of m 1 . Posterior border of 
anteorbital foramen over parastyle of m 2 . Palate essentially as 
in Talpa, but vacuities smaller and incisive foramina large, their 



24 



INSECTIVORA 



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

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

in general aspect. An- 
terior upper incisor much 
the largest of all the 
teeth, the two together 
closing entire front of 
palate ; shaft triangular 
in cross section, and all 
three faces sub-triangular 
in outline, the posterior 
and antero-external faces 
widest, theformer slightly 
concave, the latter 
slightly convex, the two 
forming a perpendicular 
external cutting edge 
about 4 mm. in length 
along their line of con- 
tact ; a much shorter but 
well developed cutting 
edge along line of contact 
of posterior and antero- 
internal faces, extending 
from acutely triangular- 
pointed apex of tooth to 
point of contact with 
tooth of opposite side, 
a distance of about 
■ 6 mm. ; line of contact 
between antero-external 
and antero-internal faces 
ridge ; height of shaft 





Fig. 6. 
Galemys pyrenaicus. Teeth x 5. 



marked by a slight though evident 

about equal to width of palate ; first and second lower incisors 
small, chisel-shaped, strongly imbricated, the second about 
twice as large as first, their shafts directed forward in line 
with upper portion of symphysis menti, the tips of the four 
teeth together forming a straight transverse cutting edge which 



GALEMYS 25 

acts in opposition to combined posterior surface of large upper 
incisors. Upper unicuspids forming an unbroken row continuous 
posteriorly with series of cheek teeth, but separated anteriorly 
from large incisor by distinct space into which the apex of 
second lower incisor fits when jaws are closed. Two anterior 
unicuspids (i 2 and i 3 ) minute, terete, single-rooted, their axes 
directed backward and falling in same line with those of two 
anterior lower unicuspids when jaws are closed. Third unicuspid 
(canine) two-rooted, its crown perpendicular, compressed, larger 
than those of first and second combined, and distinctly greater 
in height. Fourth unicuspid (pm 1 ) single-rooted, subterete, 
scarcely larger than second. Fifth and sixth unicuspids (pm 2 
and pm 3 ) essentially like canine, their crowns distinctly higher 
than wide, compressed obliquely to the tooth-row, with slightly 
developed anterior and posterior cutting edge. Height of third 
and fifth sub-equal, greater than in the others, their tips about 
on level with main cusps of molars ; sixth lower than fifth but 
with crown longer and posterior ridge better developed. Lower 
unicuspids not unlike pm 2 and pm? but with crowns lower and 
longer, slanting a little forward, each with a faintly developed 
antero-internal lobule. These teeth are slightly imbricated and 
their form approximates that of the unicuspids of the Soricidie. 
First and second (i s and c ) higher than third, their form suggest- 
ing that of anterior lower incisor, all three single-rooted. 
Fourth larger than any of the first three, obscurely two-rooted ; 
fifth slightly smaller, single-rooted ; sixth (pm 4 ) largest of the 
series, distinctly two-rooted, its cusp nearly on level with 
main cusps of molars, its an tero- external lobule (rudiment of 
parastyle) more evident than in the others. Large upper pre- 
molar three-rooted, its crown area about equal to that of third 
molar, its main cusp with well developed anterior and posterior 
cutting edges, its antero-internal cusp small but evident, its 
postero-internal cusp about equal to protocone of m 3 . Upper 
molars with crowns wider and less oblique than in Talpa 
europsea, and main cusps not so high. Transverse groove 
between bases of main cusps converted into a median pit by 
better development of commissures of protocone and larger size 
of protoconule and metaconule. Paracone and metacone about 
equal in height, the latter slightly the more robust. Styles well 
developed, except the reduced parastyle of to 1 , the mesostyle in 
each tooth completely divided into two cusps, the W-pattern thus 
changed into two V-shaped figures. Third upper molar with 
crown area about two-thirds that of second, its metastyle and 
fourth commissure absent. Lower molars essentially as in 
Talpa europsea, but contrast in height of outer and inner 
cusps very slight. 



26 



INSECTIVORA 



Galemys pyeenaicus pyeenaicus Geoffroy. 



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

p. 193. 
1910. Myogale pyrenmca Trouessart, Paune Mamm. d'Europe, p. 60. 

Type locality. — Near Tarbes, Hautes-Pyrenees, France. 

Geographical distribution. — Pyrenees and adjacent portion of 
southern Prance ; probably also north-eastern Spain to the 
Ebro ; Asturias 1 

Diagnosis. — Hind foot, 32 • 4 to 34 • 6 mm. ; condylobasal 
length of skull about 33 to 34 mm. 

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

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



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

Spain : Pajares, Leon, 2. 

2 al. Ax - les - Thermes, Ariege, Toulouse Museum 1. 7. 27. 1-2. 

720 m. Prance. (p). 

<5, 9. Ax-les-Thermes, Ariege. V. Builles (p). 8. 3. 27. 4-5. 

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

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

1 al. Pyrenees. (No history.) 

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

daki). 

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

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

Galemys pyeenaicus eufulus Graells. 

1897. Myogalea rufula Graells, Mem. Eeal. Acad. Sci., Madrid, xvii, p. 460. 
1910. Myogale pyrenaica rufula Trouessart, Paune Mamm. d'Europe, p. 61. 

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

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

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



27 



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28 



INSECTIVORA 



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

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

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

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

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

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

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

Family SORICID^E. 
1821. Soricidie Gray, London Med. Repos., xv, p. 800, April 1, 1821. 

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

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



SORBX 29 

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

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

KEY TO THE EUROPEAN GENERA OP SORIGIDM. 

Posterior lower molar with five cusps ; teeth pigmented 

at tips ; tail without sprinkling of elongated hairs. 

Upper unicuspid teeth 5-5 ; cutting edge of anterior 

lower incisor with more than one lobe ; feet not 

fringed Sorex, p. 29. 

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

Shrews) Neomys, p. 65. 

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

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

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

Genus SOREX Linnaeus. 

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

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

p. 119 (Included the undeterminable Sorex constrictus Hermann 

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

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

tetragonurus animal) Part. 
1838. Oorsira Gray, Proc. Zool. Soc. London, 1837, p. 123. June 14, 1837 

(vulgaris = araneus). 
1842. Otisorex De Kay, Zool. of New York, I, Marnm., p. 22 (platyrhinus 



1857. Sorex Blasius, Saugethiere Deutschlands, p. 124. 
1890. Somalurus Schulze, Schriften Naturwiss. Vereins Harzes in Werni- 
gerode, v, p. 28 (alpinus). 

Type species. — Sorex araneus Linnaaus. 

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



30 



INSECTIVORA 



3-3 
1-1' 



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

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

KEY TO THE EUROPEAN POEMS OP SOBEX. 

Anterior mandibular incisor with low, sometimes ill- 
defined lobes on cutting edge ; first lower uni- 
cuspid two-pointed ; lachrymal foramen over 
point of contact between m' and m 2 ; tail about 
as long as head and body ; colour uniform dark 

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

Condylobasal length of skull 19-4 to 20 - 6 mm.; 
upper tooth-row 8'6 to 9'0 mm. (Alps; east- 
ward to Transylvania) S. a. alpinus, p. 62. 

Condylobasal length of skull 19 to 19 • 6 mm. ; upper 
tooth-row 8 - 2 to 8' 6 mm. (Harz Mountains 

and Riesengebirge) S. a. hereynicm, p. 63. 

Anterior mandibular incisor with high, distinct lobes 
on cutting edge ; first lower unicuspid single- 
pointed ; lachrymal foramen in front of point of 
contact between m' and m 2 ; tail shorter than 
head and body ; colour never uniform dark slaty 
grey, the under parts usually much lighter than 
back. 
Pirst, second and third upper unicuspids sub-equal ; 
condylobasal length of skull 14 • 8 to 16 ■ 6 mm. ; 
bead and body usually about 50 to 60 mm. 

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

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

Molars and anterior upper incisor enlarged 

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

Pirst and second upper unicuspid much larger than 
third ; condylobasal length of skull 1Y • 4 to 
20 mm. ; head and body usually about 65 to 
80 mm. (Common Shrews) S. araneus, p. 31. 



SOEBX 31 

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

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

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

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

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

(Alps and neighbouring regions) S. a. tetragonurus, p. 42. 

Back rarely if ever blackish in summer 

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

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

back (Charente, Prance) 5. u. santonus, p. 40. 

Underparts greyish or brownish, contrasted 
with back except when latter is also 
brown. 
Colour of sides not distinctly contrasted 

with that of back ; belly heavily . 

washed with wood-brown (Plains of 

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

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

Norway) S. a. aranew, p. 35. 

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

SOREX ARANEUS Linnaeus. 
(Synonymy under subspecies.) 

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

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



32 INSECTIVORA 

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

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

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




SOREX 33 

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

Skull. — The skull is slender and lightly built, with no special 
peculiarities of form as compared with that of other shrews. 
Brain-case well marked off from interorbital region, its surface 
smooth except in extreme old age, its main sutures remaining 
open until late in life. It is sub-circular in 
general outline when viewed from above, 
but with antero-external portion of border 
noticeably flattened, so that at point of 
greatest breadth there is usually an evident 
angle ; condyles scarcely visible, causing no 
break in posterior outline. Depth of brain- 
case at middle slightly more than half 
greatest width ; no sagittal crest except in 
extreme old age ; lambdoid crest at first 
confined to lateral portions of occiput, rarely 
extending to median line. Dorsal profile 
usually with evident concavity in inter- 

J . J . Sorex amneus. 

orbital region (more marked than in aorex Nat. size. 

minutus and S. alpinus). Nares broadly 
rounded posteriorly, the lateral margin obtusely angled near 
middle. Anteorbital foramen moderately large, not very con- 
spicuous when skull is viewed from in front. Lachrymal 
foramen over middle of to 1 . Mesopterygoid space nearly parallel- 
sided, less than half as wide as long. 

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

.D 



34 



INSECTIVORA 





FIG. 8. 

Sorex aran&us. Anterior 
teeth in profile. X 5. 



and posterior borders sub-equal, slightly longer than anterior 

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

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

than fourth and closely crowded between it and antero-external 

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

other teeth of the series. First lower unicuspid essentially 

similar to first and second upper, 

but crown longer than broad and 

without distinct crushing surface. 

Second larger than first, not 

conspicuously different from it in 

general form when viewed from the 

side, but cutting edge much longer 

and better developed, distinctly 

angled behind middle, the angle 

clearly representing a rudimentary 

second cusp, and transverse portion 

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

of one of the triangles of a 

molar. Large upper pre- 

molar with protocone 

smaller than in the first 

and second molars, though 

well developed and of 

essentially the same form ; 

paracone smaller than pro- 
tocone, therefore much 

smaller than in first and 

second molars ; hypocone 

small but well developed ; 

posterior margin of crown 

more deeply emarginate 

than that of molars. 

Crowns of first and second molars squarish in outline, though 

somewhat wider posteriorly than anteriorly. Outer re-entrant 

angles deeper in second than in first. Protocone long and rather 




^EHZIu 




Fie. 9. 
Sorex aratieus. Teeth x 10. 



SOBEX 35 

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

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

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

Sorex araneus araneus Linnseus. 

1758. [Sorex] araneus Linnaeus, Syst. Nat., I, 10th ed., p. 53 (Sweden). 

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

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

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

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

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

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

iv, I, p. 45. 

1839. S\orex~\ macrotrichus de Selys-Longchamps, Etudes de Micromamm., 

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

the S. macrotrichus Mehlis MSS. No locality given.) 
1839. S\prex~] labiosus Jenyns, Ann. Nat. Hist., n, p. 326, January, 1839. 

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

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

Type locality. — Upsala, Sweden. 

Geographical distribution. — Western Continental Europe, 

d 2 



36 



INSECTIVORA 



araneus. 


tetragonurus. 


•o% 


.. 38-7% 


22-6% 


.. 93-7% 


21-2% 


.. 88-7% 


45-3% 


.. 100-0 % 


54-6% 


■o% 


•o% 


.. 37-2% 



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

Diagnosis. — Size small (condylobasal length of skull usually 
17-8 to 19 mm., hind foot usually 11 to 13 mm.) ; colour rather 
dark, the back ranging from bister to a deep blackish brown ; 
sides distinctly lighter than back except in specimens representing 
the pallid extreme of colouration ; teeth moderately pigmented, 
the hypocone of m 1 and m 2 usually white to tip. 

Teeth. — The teeth show no special peculiarities of form. In 
pigmentation they represent the extreme of restriction of the dark 
areas. While the area of pigmentation on all of the cusps is less 
extensive than in the Alpine and Pyrenean races, the differences 
are best seen in the hypocones of the three large upper cheek 
teeth and protocone of m 3 , as these small cusps, unlike the larger 
ones, may be completely without brown colour. A comparison 
of seventy-five topotypes of Sorex araneus with eighty Swiss 
specimens of S. araneus tetragonurus gives the following results : — 

Large premolar with pigment on hypocone 
First molar with pigment on hypocone . 
Second molar with pigment on hypocone 
Third molar with pigment on protocone 
None of the small cusps pigmented 
All of the small cusps pigmented . 

Measurements. — Average and extremes of twenty specimens 
from the type locality : head and body, 77 ■ 5 (72-85) ; tail, 40 • 1 
(38-43); hind foot, 12-6 (12-13). Average and extremes of 
nine specimens from Lillehammer, central Gudbrandsdal, Norway 
(dry): hind foot, 12-3 (12 "2-12 '6). Average and extremes of 
four specimens from Holaaker, upper Gudbrandsdal, Norway : 
head and body, 70-2 (69-71); tail, 36-5 (34-39); hind 
foot, 12 • 2 (12-13). Average and extremes of twenty specimens 
from Brunswick, Germany : head and body, 78 ■ 9 (68-85) ; 
tail, 39 • 7 (36-45) ; hind foot, 12 • 7 (12 • 2-13 • 0). Average and 
extremes of ten specimens from Waremme, Liege, Belgium : head 
and body, 68-9 (66-72); tail, 42-3 (38-47); hind foot (dry), 
12 • 2 (11 • 8-12 - 8). For cranial measurements see Table, p. 46. 

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

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

Sweden: Upland, 1; Upsala, 97 (U.S.N.M.); Skaane, 3 (U.S.N.M.). 

Denmark: Hillerbd, Zealand, 6; Nystad, Lolland, 3 (U.S.N.M.); 
Skansen, Lolland, 1 (U.S.N.M.). 

Holland: Oosterbeek, Guelderland, 6; Leiden, 4 (U.S.N.M.). 

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

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



SOREX 



37 



Germany : Brunswick, 35 (B.M. and U.S.N.M.) ; Bodethal, Harz Mts., 
15 (U.S.N.M.) ; Mauseklippe, Harz Mts., 2 (U.S.N.M.) ; Bahrenberg, Harz 
Mts., 9 (U.S.N.M.); Tharand, Saxony, 1; Magdeburg, Saxony, 5; Moritz- 
burg, Saxony, 1 (U.S.N.M.); Ingelheim, Rheinhessen, 3; Nuremberg, 
Bavaria, 10 (U.S.N.M.) ; Marxbeim, near Monheim, Bavaria, 15 ; Strass, 
near Burgheim, Bavaria, 1 ; Eulengrund, Riesengebirge, Silesia, 2 
(U.S.N.M.) ; Wolfsbau, near Sneekoppe, Riesengebirge, Silesia, 8 (U.S.N.M.); 
Niesky, Silesia, 5 ; near Konigsberg, 6 (U.S.N.M.) ; no exact locality, 1. 

Austria-Hungary : Haida, Arva, Bohemia, 9. 



6,9. 

2 <5,2 9. 

i. 

6, 5 9. 

c5al. 

9. 

4<5,2 9. 

i 6, 2 9. 

9. 

3 6, 1 9. 

2 i, 2 9. 

6, 3 9, 2. 

9. 
Sal. 

<5,2 9. 

3 3, 4 9. 

2 6. 

i. 

3<J,2 9. 

2al. 

1. 

3 £, 4 9. 

2. 



Molmen, Gudbrandsdal. R. J. Cuninghame 



Norway. 

Holaaker, Gudbrands- 
dal, 1900 ft. 

Lesjevark, Gudbrands- 
dal. 

Holme, Mandal, 200 ft. 
Norway. 

Upland, Sweden. 
(<?. Kolthqff.) 

Hillerod, Zealand, 10 m. 
Denmark. 

Oosterbeek,Guelderland, O. Thomas (c & p 
10-15 m. Holland. 

Hastiere, Namur, Bel- 
gium. 

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

Barbizon, Seine - et 
Marne. 

Auerum Forest, Bruns- 
wick, Germany. 

Tharandt, Saxony. 

Magdeburg, Saxony. 



R. J. Cuninghame 

(Pi- 
Miller Collection. 

R. J. Cuninghame 

Lord Lilford (p). 

O. Thomas (c&p). 



G. A. Boulenger 

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

G. S. Miller (c). 



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

Ingelheim, Rheinhessen. C. Hilgert (o). 
Marxheim, Bavaria. Lord Lilford (p). 

Bayreuth, Bavaria. Miller Collection. 

Strass, Burgheim, Ba- Lord Lilford (p). 

varia. 

Niesky, Silesia, 200 m. Lord Lilford (p). 

(W. Baer.) 

No exact locality. Zool. Soc. Collec- 
tion. 

Germany. Stockholm Museum 

(■)• 
Haida, Bohemia. Lord Lilford (p). 

Haida, Bohemia. Lord Lilford (p). 



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

7. 7. 7. 4452. 

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

98. 6. 7. 2-7. 

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

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

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

8. 11. 2. 6-8. 

9. 9. 8. 7-13. 

7. 7. 7. 2S68- 
2869. 

8. 9. 8. 20. 

99. 1. .9. 9-13. 

55. 12. 26. 300- 

301. 
46. 6. 2. 36. 

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



Sobex aeaneus castaneus Jenyns. 

1838. S[orex] tetragonurus var. $ S[prex] castaneus Jenyns, Ann. Nat. 

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

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

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



38 



INSECTIVORA 



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

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

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

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

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

Scotland : Black Isle, Cromarty, 16 ; South Sutor, Cromarty, 5 ; 
Nairn, Morayshire, 4 ; Dunphail, Elgin, 2 ; Gordonstown, Elgin, 3 ; Lhan- 
bride, Elgin, 1; Lossiemouth, Elgin, 3; Grantown-on-Spey, Elgin, 26 
(Wilson) ; Kennordy, 1 (Wilson) ; Cortachy, Forfar, 6 (Wilson) ; Cromlix, 
Stirling, 10 ; Islay, 3 ; Dunkeld, Perthshire, 2 ; Loch Earn Head, Perth- 
shire, 1 ; Stockbriggs, Lanarkshire, 2 ; Kirtle Bridge, Dumfriesshire, 4 ; 
Wyseby, Dumfriesshire, 3. 

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

England : Berwick-on-Tweed, Northumberland, 2 ; Biding Mill-on- 
Tyne, Northumberland, 3; Newby Bridge, Lake Windermere, Cumber- 
land, 1 ; Grimsby, Lincolnshire, 17 ; Whitnash, Warwickshire, 1 ; Bugby, 
Warwickshire, 2 ; Piley, Yorkshire, 3 ; Wellersey Hill, Broadway, Wor- 
cestershire, .1 ; West Cheshire, 1 ; Shropshire, 1 ; Staffordshire, 1 ; Swith- 
land, Leicestershire, 10 ; Bishopstoke, Herefordshire, 1 ; Leominster, Here- 
fordshire, 1 ;• Graffconbury, Herefordshire, 19 ; Lilford, Northamptonshire, 
2 ; Drinkstone Park, Bury St. Edmunds, Suffolk, 2 ; Lowestoft, Suffolk, 6 ; 
Wormsley, Oxfordshire, 1 ; Stokenchurch, Oxfordshire, A ; Cambridge- 
shire, 1 ; Kensington Gardens, London, 1 ; Hillingdon, Middlesex, 3 ; 
Bletchingley, Surrey, 4 ; Godalming, Surrey, 2 ; Merstham, Surrey, 1 ; 
Bichmond Park, Surrey, 1 ; Crowborough, Sussex, 3 ; St. Leonard's, 
Sussex, 1 ; Tunbridge Wells, Sussex, 1 ; Bastwell, Kent, 3 ; Lyndhurst 
Boad, Hampshire, 2; New Forest, Hampshire, 17; Basingstoke, Hamp- 
shire, 1 ; Alum Bay, Isle of Wight, 3 ; Clifton Bridge, Gloucestershire, 2 ; 
Leigh Woods, Clifton, Gloucestershire, 3 ; Blandford, Dorsetshire, 1 ; 
Combmartin, Devonshire, 4 ; Chagford, Devonshire, 5 ; Northlew, Devon- 
shire, 6 ; no exact locality, 8. 

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



S0KEX 39 

race the dark extreme is less dark and less frequent, while the 
light extreme is more light and more frequent. The constantly 
small size of this animal as compared with the large Continental 
races (bergensis, tetragonurus and pyrenaicus), is .shown by the 
fact that among 102 British specimens measured by many 
different collectors the hind foot exceeds 13 '2 mm. in only six 
instances. 

9. Black Isle, Cromarty, W. R. Ogilvie-Grant 94.10.6.1. 

Scotland. (c & p). 

9(5. Black Isle, Cromarty. W. R. Ogilvie-Grant 11. 1. 3. 103- 

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

2c5. South Sutor, Cromarty. W.R. Ogilvie-Grant 11. 1. 3. 108, 

(c & p). 113. 

2 (5, 2 9. Nairn, Morayshire. W. R. Ogilvie-Grant 11. 1. 3. 99-102. 

(c & p). 

6. Morayshire. E. R.Alston (c&p). 79.9.25.79. 

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

(c & p). 

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

2 6. Gordonstown, Elgin- W. R. Ogilvie-Grant 11. 1. 3. 95-96. 
shire. (c & p). 

<5. Gordonstown, Elgin- W. R. Ogilvie-Grant 11.1.3.98. 
shire. (c & p). 

3 <J, 7 9. Cromlix, Stirlingshire. W. R. Ogilvie-Grant 11.1.3.114-123. 

(c & p). 

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

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

shire. 

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

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

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

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

Wales. (c & p). 163. 

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

shire. 169. 

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

Northumberland, 

England. 

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

land. 

Grimsby, Lincolnshire. G. H. Caton Haigh 11. 1. 3. 124- 

(c & p). 131. 

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

shire. 138. 

Shropshire. T. 0. Eyton (c & p). 63. 10. 12. 4. 

Bishopatoke, Hereford- S. O. Ridley (c&p). 84.10.6.1. 

shire. 

Graftonbury, Hereford- W. de Winton 11. 1. 3. 72-87. 

shire. (c&p). 

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

shire. (c&p). 

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

Suffolk. 

Lowestoft, Suffolk. O. Thomas (c & p). 11. 1. 3. 139- 

144. 

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

shire. (c&p). 148. 

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

(O. Thomas.) 3597. 



3al. 
6,9. 


6, 3 9. 


3 9. 
5 6,39, 


3 6, 3 9. 


2al. 


lal. 


■5,5 9,2. 


<5,6 9. 


lal. 
9al. 


8 <5, 8 9. 


3 st. 


2al. 


3=5,3 9. 


4 9. 


3 9. 



40 INSECTIVOEA 

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

London. 

4 9. Bletohingley, Surrey. W. R, Ogilvie-Grant 11. 1. 3. 149- 

(p). 152. 

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

(W.R.Ogilvie-Grant.) 

Sal. Richmond Park, Surrey. Prof. Owen (p). 75.9.17.1. 

7 S, 10 9. NewForest,Hampshire. Miller Collection. 7. 7. 7. 2851- 

2867. 3023. 

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

shire. (W. P. Stark.) 

5,2 9. Alum Bay.Isle of Wight. 0. Thomas (c & p). 11.1.8.153-155. 

9 (albino) Winscombe, Somerset. P. A. Knight (o. & p). 4. 8. 9. 1. 

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

(c & p). 174. 

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

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



SoKEX ARANEUS SANTONUS Mottaz. 

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

April 30, 1908. Type in Mottaz Collection. 
1910. Sorex santonus Trouessart, Paune Mamm. d'Europe, p. 54. 

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

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

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

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

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

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

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

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



41 



SOREX ARANEUS EURONOTUS Miller. 



1901. Sorex araneus euronotus Miller, Proo. Biol. Soc, Washington, xiv, 
p. 44, April 25, 1901 (Montrejeau, Haute-Garonne, Prance). Type 
in U.S. National Museum. 

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

Type locality. — Montrejeau, Haute-Garonne, France. 

Geographical distribution. — Probably the plains between the 
Pyrenees and the Garonne. At present known from the type 
locality only. 

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

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

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

SOREX ARANEUS BERGENSIS Miller. 

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

m, p. 416, May, 1909. Type in U.S. National Museum. 

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

Type locality. — Graven, Hardanger, Norway. 

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

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

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

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



42 INSECTIVOKA 

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

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

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

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

and U.S.N.M.). 

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

E. G. B. Meade 5. 7. 1. 2. 

Waldo (c & p). 

0. H. Stephenson 8. 1. 5. 1-5. 

(c & p). 

Miller Collection. 7. 7. 7. 4453- 

4456. 

Miller Collection. 7. 7. 7. 4457-9. 

G. Barrett-Hamil- 8. 9. 21. 1. 

ton (p). 

Soeex araneus tetragonurtjs Hermann. 

1780. Sorex tetragonurus Hermann in Zimmermann, Geogr. Gesch., n, 
p. 383 (Strassburg, Germany). 

1792. Sorex guadricaudatus Kerr, Anim. Kingd., p. 208 (Strassburg, 
Germany). (Based on Pennant's account of S. tetragonurus 
Hermann.) 

1835. Sorex fodiens Duvernoy, Mem. Soc. du Mus. d'Hist. Nat., Stras- 
bourg, ii, p. 17. Part : skull, not animal. 

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

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

1868. ? Sorex araneus, pallidus Fitzinger, Sitzungsber. kais. Akad. Wis- 

sensch., Wien, Math. Naturwiss. Classe, lvii, pt. I, p. 488 
(Based on specimen from unknown locality, probably in Italy, 
figured by Bonaparte, Iconogr. Paun. Ital., fasc. xxix, pi., fig. 5.) 

1869. [Sorex vulgaris] var. nuda Patio, Paune Vert. Suisse, I, p. 127 

(Bernese Oberland, Switzerland). 



1. 


Vefsen, Nordland, Nor- 


,5,4?. 


way. 
Skjserdal, Nordfjord. 


2(5,2 9. 


Opheim, Bergen. 


6,2 9. 

9. 


Bergen. 
Bergen, 2,700 ft. 



SOEBX 43 

1869. [Sorex vulgaris] var. nigra Patio, Faune Vert. Suisse, i, p. 127 
(Lucerne, Switzerland). 

1900. S[orexr\ vulgaris var. vel subsp. mollis Patio, Bev. Suisse de Zool., 

viii, p. 471 (Substitute for nigra). 

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

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

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

1905. Crossopus ou Sorex ignotus Patio, Arch. Sci. Phys. et Nat., Geneve, 
4th ser., xix, p. 202, February 15, 1905 (Switzerland). Part: 
mandible, not skull. Type in Geneva Museum. 

1905. Sorex araneus carpathicus Barrett-Hamilton, Ann. and Mag. Nat. 
Hist., 7th ser., xv, p. 506, May, 1905 (Hatszeg, Huny ad, Hungary) . 
Type in British Museum. 

1910. So7-ex araneus tetragonurus and S. araneus carpathicus Trouessart, 
Paune Mamm. d'Burope, pp. 52, 54. 

Type locality. — Strassburg, Germany. 

Geographical distribution. — Alps and neighbouring portions of 
Germany, Prance and Italy ; eastward through Tirol to the 
mountains of Transylvania. 

Diagnosis. — Similar to Sorex araneus bergensis, but colour in 
summer pelage not so dark, the contrast between back and 
sides often not conspicuous, though back is frequently blackish ; 
pigmentation of teeth more extensive than in the northern races, 
the hypocone of m 1 and m 2 usually brown at tip (see tabular 
comparison with true araneus on p. 36). 

Measurements. — Average and extremes of eight specimens 
from the type locality : head and body, 75-8 (71-79) ; tail, 47-5 
(45-50); hind foot, 13-5 (13 -2-14 '2). Average and extremes 
of twenty specimens from Andermatt, Uri, Switzerland : head 
and body, 73-1 (70-81); tail, 51-1 (50-55); hind foot, 13-3 
(13-14). Measurements of an adult female from Zermatt, Valais, 
Switzerland (in alcohol) : * head and body, 74- 6 ; tail, 51 ; hind 
foot, 13-6. Average and extremes of three specimens from 
Vitznau, Switzerland : f head and body, 67 -3 (62-72); tail, 
46-3 (44-49); hind foot, 13-2 (12-8-13-6). Type of Sorex 
araneus alticola Miller (Meiringen, Switzerland, No. 85,930 
U.S.N.M.) : head and body, 76 ; tail, 55 ; hind foot, 14. Average 
and extremes of ten specimens from Meiringen, Switzerland : J 
head and body, 74-0 (70-77); tail, 52-5 (47-57); hind foot, 
13-3 (13-14). Average and extremes of four specimens from 
Locarno, Ticino, Switzerland: head and body, 76-7 (76-82); 
tail, 48-5 (45-50); hind foot, 13-6 (13-4-13-8). For cranial 
measurements see Table, p. 49. 

* Ootype of Sorex vulgaris crassicaudatus Patio, 
t Essentially topotypes of Sorex vulgaris nigra and mollis Patio. 
X Topotypes of Sorex araneus alticola Miller and essentially topotypes 
of Sorex vulgaris nuda Patio. 



44 



INSBCTIVORA 



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

France : Etupes, Doubs, 10 ; Barcelonnette, Basses-Alpes, 3 ; Cha- 
monix, Haute-Savoie, 10 (U.S.N.M.). 

Germany : Strassburg, 8. 

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

Austria-Hungary : Hatszeg, Hunyad, Transylvania, 10 ; Csall6k8z- 
Somorja, Pressburg, Hungary, 3; Schwaz, Tirol, 2 (U.S.N.M.). 

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

0. Thomas (p). 8. 8. 10. 14-24. 

O. Thomas (p). 8. 8. 10. 25-27. 

0. Thomas (p). 8. 8. 10. 28-32. 

Dr. J. Anderson 91. 10. 15. 26. 

(c & p). 

0. Thomas (o & p). 5. 8. 2. 32-34. 

C. Mosch (p). 89. 11. 8. 3 

0. Thomas (c & p). 5. 8. 2. 19. 

0. Thomas (o & p). 5. 8. 2. 2-5. 

E. B. Alston (p). 79. 9. 25. 10. 

C. G. Danford (o). 3. 2. 2. 3-10. 
3. 11. 8. 15. 

Budapest Museum 94. 3. 1. 22-25. 
(«)■ 

Dr. G. Ceeconi 1. 8. 2. 2-3. 
(o & p). 



8<5,2 9. 


Etupes, Doubs, 350 m. 




France. (0. Mottaz). 


3 6. 


Barcelonnette, Basses- 




Alpes. (0. Mottaz) 


4 6, 1 9. 


Strassburg, Alsace. 




(C. Mottaz.) 


1 al. 


Zermatt, Valais, Swit- 




zerland. 


3 6. 


Vitznau, Lake Lucerne. 


1 al. 


Zurich. 


9. 


Faido, Ticino. 


2.5,2 9. 


Locarno, Ticino. 


9. 


Switzerland. 


8 <J, 1 9. 


Hatszeg, Hunyad, Tran- 




sylvania, 1500-2000ft. 




Hungary. 


3. 


Csall6koz-Somorja, 




Pressburg, 400 ft. 




Hungary. 


2al. 


Vallombrosa, Florence, 




Italy. 



SOEEX AEANEUS PYRENAICUS Miller. 

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

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

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

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

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

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



SOEEX 45 

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

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

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

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

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

Porte, Pyrenees-Orientales (Spanish watershed), 9 ; l'Hospitalet, Ariege, 
12 ; Ax-les-Thermes, Ariege, 2 ; Bareges, Hautes-Pyrenees, 9. 

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

S, 9. Porte, Pyrenees-Orien- G. S. Miller (o). 8. 8. 4. 142-143. 

tales, 1600m. France. 
2 <5, 9. Ported Pyrenees-Orien- 0. Thomas (p). 8. 9. 1. 49-51. 

tales, 1600-1700 m. 
(A. Robert). 
S 4 9. L'Hospitalet, Ariege, G. S. Miller (c). 8.8.4.151-154. 

4700 ft. 301. 

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

2600 ft. 
<5 6 9. Bareges, Hautes-Pyre- G. S. Miller (c). 8. 8. 4. 144-150. 

nees, 1300-1500 m. 



SOEEX AEANEUS FRETALIS Miller. 

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

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

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

Type locality. — Trinity, Jersey, Channel Islands. 

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

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



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

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

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

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

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

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

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

?, <$. Jersey, Channel Islands. 0. Thomas (p). 8.9.2.1-2. 

(B. H. Bunting.) (8. 9. 2. 1. Type of subspecies.) 



SoBEX AEANEUS GRANARIUS Miller. 

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

Type locality. — La Granja, Segovia, Spain. 

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

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

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



SOREX 53 

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

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

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

Specimens examined. — -Two, both from the type locality. 

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

2 al. La Granja, Segovia, M.delaEsoalera(c). 6.11.4.3-4. 
Spain. (6. 11. 4. 4. Type of subspecies.) 

SOREX MINUTUS Linnaeus. 

(Synonymy under subspecies.) 

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

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

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

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



54 



INSECTIVOEA 




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

Skull. — Apart from its conspicuously smaller size (condylo- 
basal length, 14-8 to 16 mm. instead of 17 "8 
to 20 mm.) the skull diners notably from 
that of Sorex araneus and S. alpinus in the 
narrower, more elongate brain-case, the outline 
of which when viewed from above is distinctly 
oval instead of sub-circular. Antero-external 
portion of border flattened, though less notice- 
ably than in Sorex araneus. Owing to its 
different form the brain-case is less abruptly 
marked off from interorbital region than in 
Sorex i. > ;tw>. ^he °tber European species. Depth of brain-case 
Nat. size. usually less relatively to breadth than in 

S. araneus. Lachrymal foramen opening over 
posterior half of first molar. 

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distinctly smaller than anterior cusp, while from the side its 
outline is conspicuously higher and narrower than that of first 
unicuspid. Anterior lower incisor with 
first lobe on cutting edge as well defined 
from anterior point of tooth as from second 
lobe, and lengths of bases of all three lobes 
approximately equal. Upper unicuspids not 
essentially different in form from those of 
Sorex araneus, but crowns perceptibly longer 
than wide and less abruptly rounded off 
anteriorly, the resulting form less nearly Vla ' n ' 

square. In lateral view the outline is quite S °tSTin ^SSie^x *" 
as in the larger animal. The relative size 

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




SDEEX 55 

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

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

Soeex minutus minutus Linnseus. 

1766. [Sorex] minutus Linnseus, Syst. Nat., I, 12th ed., p. 73 (Siberia). 
1769. Sorex pygm&us Laxmann, Sibirische Briefe, p. 72 (Barnaul, Tomsk, 

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

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

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

Jornkoping, Sweden). 
1811. Sorex pygmseus Pallas, Zoogr. Bosso-Asiat., I, p. 134 (Ob and Yenesei 

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

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

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

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

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

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

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

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

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

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

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



56 



INSECTIVORA 



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

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

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

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

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

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

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

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

Prance : Barbizon, Seine-et-Marne, 1 ; Montrejeau, Haute-Garonne, 1 
(U.S.N.M.); Ax-les-Thermes, Ariege, 2; l'Hospitalet, Ariege, 3; Porte, 
Pyrenees-Orientales, 1. 

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

Austria-Hungary: CsaU6k8z-Somorja, Pressburg, Hungary, 1 ; Hatszeg, 
Hunyad, Transylvania, 7. 

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

Italy: Vallombrosa, near Florence, 1. 

1. Lossiemouth, Moray- G. Denson (c & p). 8. 9. 17. 1. 
shire, Scotland. 
9,1. Lhanbryde,Elgin,200ft. W. Taylor (c & p). 8.9.3.1-2. 
¥. Dunblane, Stirlingshire. W.E. Ogilvie-Grant 8.9.6.1. 

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

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

<5. Stornoway, Lewis. R. M. D. Hawker 8. 9. 18. 1. 

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

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

(c & p). 



57 



<5. Stockbriggs, Lanark- 
shire. 

1. Bridgend, Glamorgan- 

shire, Wales. 

1 al. Wales. 

9. Grimsby, Lincolnshire, 
England. 

2. Waltham, Lincolnshire. 

■2 st. Graftonbury, Hereford- 
shire. 
9. Wilbraham, Cambridge- 
shire. (S.F.Harmer.) 

1. Clifton, Gloucestershire. 

(R. I. Pocock.) 
6. Lundy Island, Devon- 
shire. 

2 <5. Chagford, Devonshire, 

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

i. Sulby Glen, Isle of Man. 

9. Clonbroch, Galway, Ire- 
land. 

1 al. Longford. 

$ st. Knock, Down. 

2 al. Dunkettle, Cork. 

2 S, 1 9. Duncannon, Wexford. 
(Bev. Dr. Martin.) 

1 al. Ireland. 

tj. Holaaker, Gudbrandsdal, 
1900 ft. Norway. 

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

2 9. Ax-les-Thermes, Ariege. 
i. 1' Hospitalet, Ariege, 

4700 m. 
(5. 1' Hospitalet, Ariege, 

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

tales, 1600 m. 

2. Querum Forest, Bruns- 

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

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

6. Marxheim, Bavaria. 

1. Csall6k8z-Somorja, Press- 
burg, 400 ft. Hungary. 

1. Hatszeg, Hunyad, Tran- 
sylvania, 2000 - 
5500 ft. 

3. Hatszeg, Transylvania, 

2000-5500 ft. 

3 6. Hatszeg, Hunyad, 1500- 

2000 ft. 



E,R. Alston (c&p). 

B.I. Pocock (c&p). 

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

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

(c & p). 
W. de Win ton 

(c & p). 
Miller Collection. 

Miller Collection. 

N. H. Joy (c&p). 

Miller Collection. 

C. H. B. Grant 

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

(c & p). 
H. F. Witherby 

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

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

child (c&p). 
MissO'Keefe(c&p). 

G. Barrett-Hamilton 

Zoological Society's 

Collection. 
B. J. Cuninghame 

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

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

0. Thomas (p). 

G. S. Miller (c). 

G. Barrett-Hamilton 

Tomes Collection. 

Lord Lilford (p). 

Lord Lilford (p). 
Budapest Museum 

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



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



79. 9. 25. 13. 

8. 9. 5. 1. 

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

11. 1. 2. 81-82. 

96. 4. 28. 20-21. 

7. 7. 7. 2850. 

7. 7. 7. 3604. 

6. 5. 16. 1. 

7. 7. 7. 4472, 
4474. 

8. 9. 7. 1-5. 

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

80. 12. 14. 6. 

I. 9. 3. 7. 

80. 11. 20. 4. 

81. 6. 29. 1. 

II. 1. 2. 83-85. 

52. 9. 13. 288. 

98. 2. 28. 5. 

8. 8. 4. 160. 

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

8. 9. 1. 45. 

8. 8. 4. 161. 

11. 1. 2. 86-87. 

7. 1. 1. 46. 

99. 1. 9. 15. 

11. 1. 1. i46. 
94. 3. 1. 25. 

3. 2. 2. 11. 

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



58 



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60 



INSECTIVORA 



1. St. Gothard, Uri, Swit- Purchased (Brandt). 46. 2. 13. 14. 
zerland. 
1 al. Vallombrosa, Florence, Dr. G. Cecconi (p.) 1. 8. 2. 4. 
Italy. 



Sorex minutus lucanius Miller. 

1909. Sorex minutus lucanius Miller, Anrj. and Mag. Nat. Hist., 8th ser., 

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

1910. Sorex minutus lucanius Trouessart, Fauna Mamm. d'Europe, p. 55. 

Type locality. — Monte Sirino, Lagonegro, Italy. 

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

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

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

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

Specimen examined. — The type. 

1. Monte Sirino, Lagonegro, O. Thomas (p). 8. 9. 1. 5. 

S. Italy. (A. Robert.) {Type of subspecies.) 



SOREX ALPINUS Schinz. 
(Synonymy under subspecies.) 

Geographical distribution. — Pyrenees, Alps, Harz Mountains, 
JRiesengebirge and Carpathians. 

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

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



SOEEX 



61 



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

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

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

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

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




62 



INSBCTIVORA 





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

"Pin 1^ 

peculiarities of form. The first, how- 
Sorex alpinus. Anterior L . . ., . , ,.~, . e ,, 

teeth, x s. ever, is strikingly dinerent from the 

corresponding tooth in S. araneus. 

Its general outline when viewed from side is irregularly 

elliptical, with longest axis parallel to that of mandible, the 

upper edge with two low, rounded cusps, the anterior of which 

is about as large as lobes on cutting edge of incisor, the posterior 

smaller. Upper cheek-teeth not essentially different from those 

of Sorex araneus, except that hypocones are less developed. 

Lower cheek-teeth as in the related species. Pigmentation of 

teeth slightly less extensive than usual in Sorex araneus. 

Sorex alpinus alpinus Schinz. 

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

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

locality, and probably not a European species). 
1857. Sorex alpinus Blasius, Saugethiere Deutschlands, p. 126. 
1870. ? Sorex intermedins Cornalia, Catal. Descrit. Mamm. Ital., p. 27 (Hills 

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

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

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

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

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

* In an adult female from Briinig, Switzerland (No. 85830, U.S.N.M.), 
the third and fourth unicuspids on right side are fused into a single 
2-cusped tooth, while those on left side are normal. 

t I have seen no Pyrenean specimens of Sorex alpinus. For record of 
its occurrence see Trutat, Bull. Soc. Hist. Nat., Toulouse, xn, p. 100, 1878 
("massif de la Maladetta"). The animal is probably less common in the 
Pyrenees than in the Alps, as I was unable to find it in several localities 
resembling those where it regularly occurs in Switzerland. 



SOEEX 63 

Diagnosis. — Skull and teeth of maximum size for the species 
(condylobasal length of skull, 19 "4 to 20 - 6; upper tooth-row, 
8 • 6 to 9 • ; lower tooth-row, 8 • to 8 • 4). 

Measurements. — Average and extremes of five specimens from 
Briinig, Switzerland: head and body, 74 • 6 (72-77); tail, 73 
(70-75) ; hind foot, 16. Average and extremes of three speci- 
mens from Vitznau, Switzerland : head and body, 72 • 3 (69-75) ; 
tail, 74-0 (72-75); hind foot, 15-5 (15-16). Adult male from 
Vitznau, Switzerland : head and body, 69 ; tail, 68 ; hind foot, 
15 • 3. Adult female from Chamonix, Haute-Savoie, France: 
head and body, 73; tail, 74; hind foot, 15 • 4. For cranial 
measurements see Table, p. 64. 

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

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

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

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

i. Vitznau, 440 m. Swit- 0. Thomas (c & p). 5. 8. 3. 15. 

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

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

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

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

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

rard). 
9. Hatszeg, Hunyad, Tran- C. G. Danford (c). 3. 2. 2. 2. 
sylvania, 2000 ft. 
Hungary. 
6, 2 ?. Hatszeg, Hunyad, Tran- 0. G. Danford (c). 3. 11. 8. 12-14. 
sylvania, 1500-2000 ft. 

SOEEX ALPINUS HBHCYNICUS Miller. 

1909. Sorex alpinus hercynicus Miller, Ann. and Mag. Nat. Hist., 8th ser., 

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

1910. Sorex alpinus liercynicus Trouessart, Eaune Mamm. d'Europe, p. 51. 

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

Geographical distribution. — Harz Mountains and Riesen- 



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

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



64 



INSBCTIVORA 



Observations. 


Teeth slightly worn. 
„ not worn. 

, slightly worn. 
„ not worn. 
»j jj 
jj »j 

>J J! 

slightly worn. 
„ not worn. 

jj jj 

jj » 
j jj 
jj jj 
jj jj 
„ slightly worn. 

j j> 
„ not worn. 
„ slightly worn. 


•(airjua) 
Avoj-'qioo; 
•iTJlTiqipirBjs; 


OTtf'iKMCKMCM-^tMOOlO O CO O O O O CO O 


cbcoascocococococooDcoco cot-cbcococot-co 




QO O O CO 00 CO O O CO CO CO CD "* cq ^ CO ^H tH ■*# o 


GO OS OS CO 00 00 OS OS CO GO CO GO CO CO CO CO CO CO CO CO 


•aiqipuB'K. 


THOOTHC0tDtHOC0THO(M OCOOOO^OO 


O rH rH O O O O jH O O tH O O OS O O O O O O 
t-It-It-It-.t-It-|i-It-Ht-Ht-Ht-1.H iH i-I i-I tH t-( t-H tH 


•(nuipatu) 9BB0 
-uyeiq jo mdaa; 


cq -^ -* cq cq no t*i •* cq co ■* cm th^imnoO(no 


•asBO-nrciq 
jo qipe'aia 


CO O O O CO O CO o o cq CO CO -^ O 00 00 CO CO 00 -^ 

©OiOOO>00)000)0)Q CSOsasOSCSCSOSOS 
r-4 i-l tH rH rH 


■qipBaiq 
IBcaiiqoBi 


OCNOCMtMCM^CqCMCNCNO OOOOOOOO 
■^H tH ^Ji "^i *Q" ^^ "^i ^1 ^1 ^" ^" ^i w* ^Ji *^ji "^ ^i ^*1 ^i ^i 


■qctpuajq 
di^buioS£z 


<OCOCOtHcOCOCOCOCDCOCO-# cqOCqcq-^CMCqcq 


UilOOWiOlOWJOlOWWlO lO lO 1C iO iO ifl io m 


•q^Snai 
fES^qot^puoo 


O -^ cq o cq o ■* co O ■* o tH cq t+i cq ^ co co cq co 
oooooooooosoos osasasasasososos 

CqcqcqcqCqcqCqCqCqTHCqrH 1 -) t H,h t Hi-(t-It-i'tH 


02 


0+0+CH-00+0+0+0+CHO+CH-0+ o+o+o+o+*o*o , <j*o 


CD 

1 

ft 


* 

iOHOia.OHK, (Ba)OHCO locoocq-^t-coos 
OioiowujtoEjS^ySS cqcqcococqcqcqcq 
*a th ^i to ow S2 23 §^ £g £2 gj| os os os os os os os as 
-* ■* t*i -* -,# -* <& ™ co <g g co cq cm cq cq cq cq cq cq 






Locality. 


CO 

.'g a 


§ ^ 3 -s Ea 

l-a° 1 ss =§ ... J *l .1=11 .. 
S-a^ ► w « Ml $ -3g 

g"S w § 

ll 1 

w g: a> 



NEOMYS 65 

(72-77); tail, 66-6 (65-68); hind foot, 15- 1 (14-8-15-8). 
Average and extremes of five specimens from the Riesengebirge : 
head and body, 73-4 (72-77); tail, 65-4 (59-68); hind foot, 
14-9 (14-8-15-0). For cranial measurements see Table, p. 64. 

Specimens examined. — Eleven (all in U.S.N.M.), from the following 
localities in Germany: Bahrenberg, Harz Mts., 2; Bodethal, Harz 
Mts., 1 ; Mauseklippe, Harz Mts., 3 ; Eulengrund, Riesengebirge, Silesia, 3 ; 
Wolfshau, Eiesengebirge, Silesia, 2. 

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

Genus NEOMYS* Kaup. 

1829. Neomys Kaup, Entw.-Gesoh. u. Natiirl. Syst. Europ. Thierwelt, i, 

p. 117 (Sorex daubentonii Erxleben). 
1829. Leucorhynchus Kaup, Entw.-Gesch. u.. Natiirl. Syst. Europ. Thier- 
welt, i, p. 117 (Sorex lineatus Geoffroy and S. leucodon Hermann ; 

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

p. 102, March, 1898). 
1829. Hydrogale Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, i, 

p. 119 (Sorex remifer Geoffroy). 
1832. Crossopus Wagler, Isis, p. 275 (Sorex fodiens Schreber). 
1835. Hydrosorex Duvernoy, Mem. Soc. du Mus. d'Hist. Nat., Strasbourg, 

ii, p. 19 (Sorex fodiens Pallas, see p. 17). 
1835. Amphisorex Duvernoy, Mem. Soc. du Mus. d'Hist. Nat., Strasbourg, 

ii, p. 23 (hermanni = fodiens skull + Sorex araneus tetragonurus 

animal). Part. 
1838. Pinalia Gray, Proc. Zool. Soc, London (1837), p. 126, June 14, 1838 

(MS. synonym of Crossopus). 
1848. Galemys Pomel, Archiv Sci. Phys. et Nat., Geneve, ix, p. 249, 

November, 1848 (part, included Brachysorex Duvernoy, Crossopus 

Wagler, and Pachyura de Selys-Longchamps). 
1854. Myosictis Pomel, Catal. Meth. Vert. Poss. Bassin de la Loire, p. 14 

(based on a fossil not distinguished specifically from Neomys 

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

Type species. — Sorex daubentonii Erxleben =S. fodiens Schreber. 

Geographical distribution. — Palsearctic region north of the 
Mediterranean from Spain and England to the Caucasus and 
Asia Minor, north to northern Scandinavia. 

Characters. — Upper unicuspid teeth 4-4, their form more pre- 
hensory than in Sorex; dental formula : i £=?, eg, pm ?=?, m |=| = 30 ; 
posterior lobe of anterior upper incisor less than half as high 
as first ; anterior lower incisor with one ill-defined lobe on cutting 
edge near middle of tooth ; third lower molar and second lower 
unicuspid as in Sorex ; points of all the teeth pigmented ; fur 
unusually soft and dense, entirely concealing ears ; feet with a 
noticeable fringe of elongated hairs at edges of soles and toes, 



66 



INSEOTIVOEA 



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

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

KEY TO THE EUROPEAN FORMS OP NEOMYS. 

Tail with median keel of stiffened hairs extending 

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

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

Peninsula) X. f. fodiens, p. 69. 

Underparts usually brownish (England and Scot- 
land) X.f- bicolor, p. 73. 

Tail without keel. 

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

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

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

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

NEOMYS fodiens Schreber. 
(Synonymy under subspecies.) 

Geographical distribution. — From Norway and England to the 
Pyrenees, northern Italy and western Siberia. Northern and 
eastern limits of range not known. 

Diagnosis. — Tail with keel always present ; feet very con- 
spicuously fringed; hind foot usually more than 17 mm. ; lachry- 
mal foramen opening over posterior half of m 1 ; anterior upper 
incisor robust. 

External characters. — Fur very dense, its depth at middle of 
back about 6 mm. in summer, 9 mm. in winter, its texture soft 
and velvety, this particularly noticeable on underparts ; a 
few longer hairs usually present on flanks and rump. Eyes 
small and inconspicuous ; ears completely hidden by the fur. 
Muzzle less slender than in Sorex araneus, the median ridge on 
upper lip with distinct wart-like outgrowth which fits between 
points of upper incisors when mouth is closed. Feet larger than 



NEOMYS 67 

in species of Sorex of approximately the same size, noticeably 
broadened ; digits proportioned as in Sorex araneus but gradua- 
tion less ; soles and palms completely bare, their surface strongly 
tuberculo-reticulate ; pads 6-6, much as in Sorex araneus but 
more widely spaced and better defined. At edge of soles and 
toes the hairs are elongated to form a dense, conspicuous fringe. 
Tail terete or slightly four-sided, its hairs minute, flattened, and 
closely appressed, nearly concealing the rather indistinct annula- 
tions, of which there are about 25 to the centimeter at middle ; 
pencil evident, usually about 3 to 5 mm. in length. Along 
median line of tail below, from extreme base to tip, the hairs 
are so crowded, elongated, and directed inward as to form a 
distinct, low keel, the presence of which is often made more 
evident by the nearly bare condition of the skin immediately 
at each side of it. Mammas : a 2-2, i 2-2 = 8. 

Colour. — Entire underparts a dark slaty grey, more bluish 
in summer, more blackish in winter, the individual hairs blackish- 
slate with seal-brown tips and a sub-terminal lighter area of 
varying distinctness. Underparts usually whitish in strong 
contrast with back, the line of demarcation everywhere evident, 
and on head passing just below base of ear and extending along 
upper lip to nostril. The underparts, while never so dark as 
back, are often strongly suffused with bufly, smoky, or wood- 
brown, and occasionally tinged with salmon-colour. A small 
whitish tuft usually springs from inner surface of ear, and a 
minute speck of the same colour is generally present behind eye. 
Feet thinly sprinkled with silvery grey hairs. Tail a dark 
indefinite brown above, slightly less dark below, the keel usually 
a silver grey, sometimes standing out in strong contrast against 
the dark surrounding parts. 

Skull. — The skull differs from that of Sorex araneus in larger 
size, relatively larger, more inflated brain- 
case (depth decidedly more than half breadth), 
and lower, more flattened rostrum and 
interorbital region. The brain-case is sub- 
circular in outline, though distinctly flattened 
antero-externally and postero-externally ; in 
front it is marked off from interorbital 
region by a conspicuous, well defined angle. 
In lateral view the profile of brain-case is 
high and rounded posteriorly, low anteriorly 
where it passes by a distinct, often conspicu- 
ously abrupt curve into the nearly flat dorsal 
outline of interorbital region and rostrum. 
Anteorbital foramen relatively larger and more l °' t j- 

widely open than in the European species of .Nat. size. 

Sorex. Lachrymal foramen over posterior 
half of m 1 . Mesopterygoid space decidedly more than half as 
wide as long ; hamular processes shorter than in Sorex araneus 

f 2 




68 



INSECTIVOEA 



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

Teeth. — Though in general not strikingly different from those 
of Sorex araneus the teeth show a tendency towards greater 
elevation of the cusps as well as a more pronounced hooking 
forward of the anterior upper incisor, peculiarities which impart 
to the entire dentition, though particularly to the teeth lying 
in front of the large premolars, an unmistakably prehensive 
character. Anterior upper incisor with main hook more projecting 
than in Sorex araneus ; posterior talon low though rather long, 
its outline when viewed from the side somewhat triangular, its 
height less than half that of main cusp and much less than that 
of first unicuspid. Lower incisor more slender than that of 
Sorex araneus, its cutting edge with only a single low, ill-defined 
lobe near middle. Upper unicuspids large and strong, their 
crowns when viewed from below decidedly longer than wide, 

truncate posteriorly, narrowed an- 
teriorly and terminating in a slight 
though evident point. In lateral 
view the cusps appear to be more 
slender and set further forward 
than in Sorex araneus, and the 
posterior part of crown is more 
produced backward, the cingulum 
often rising to a distinct postero- 
internal angle or rudimentary cusp, 

particularly in first tooth. Spaces 
Xeomys fodiens. Anterior teeth i , ,. 2 . - n L . 

in profile, x 5. between tips 01 unicuspids greater 

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




NEOMYS 69 

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

Measurements. — In external measurements this species is 
unusally variable, the head and body ranging from 72 to 96 mm., 
tail from 47 to 77 mm., and hind foot from 16 to 20 mm. These 
differences appear to be for the most part strictly individual. 
The skull is more constant : condylobasal length, 19'6 to 22-2 mm. ; 
upper tooth-row, 9 • 6 to 11 mm. Here the variation is to a 
certain extent geographic, as the skull of the British race 
averages smaller than that of the Continental form. 

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

Neomys fodiens fodiens Schreber. 

1776. Sorex aquations P. L. S. Muller, Natursyst. Suppl. u. Regist.-Band, 

p. 36 (Prance ; based on Buffon, vin, pi. 11). Not Sorex aguaticus 
Linnaeus, 1758. 

1777. Sorex fodiens Schreber, Saugthiere, in, p. 571 (Berlin, Germany). 
1777. [Sorex] daubentonii Erxleben, Syst. Regni Anim., i, p. 124 (Burgundy, 

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

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

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

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

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

synomym of Sorex eremita, perhaps from Bechstein MS.). 
1800. S[orex] f[pdiens~] albus Bechstein, Thomas Pennant's Allgem. Uebers. 

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

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

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

(Abbeville, Somme, Prance). Type in Paris Museum. 
1818. Sorex collaris Desmarest, Nouv. Diet. d'Hist. Nat., xxn, p. 65 

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

but not named by Geoffroy, Mem. Mus. d'Hist. Nat., Paris, i, 

p. 309, 1815. 



* In an adult male from Geneva, Switzerland (No. 1046 Mottaz), the 
right upper incisor and first unicuspid and right lower incisor are white 
throughout. All other teeth normal. 



70 



1NSECTIVORA 



1822. Sorex macrourus Lehmann, Observ. Zoologies in faunam Hamburg- 

ensem,i,p. 5 (Saehsenwald, near Friedrichsruh, Sohleswig-Holstein, 

Germany). 
1826. Sorex amphibius Brehm, Ornis, n, p. 44 (Renthendorf, Tbiiringen, 

Germany). 
1826. Sorex natans Brebm, Ornis, II, p. 44 (Benthendorf, Tbiiringen, 

Germany). 
1826. Sorex stagnatilis Brehm, Ornis, n, p. 47 (Renthendorf, Thiiringen, 

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

1834. S[orex] nigripes Melchior, Den Danske Stats og Norges Pattedyr, 

p. 68 (Sielland, Denmark). 

1835. Sorex fodiens Duvernoy, Mem. Soc. du Mus. d'Hist. Nat., Strasbourg, 

II, p. 17. Part : animal, not skull. 
1835. Sorex hermanni Duvernoy, Mem. Soo. du Mus. d'Hist. Nat., Stras- 
bourg, ii, p. 23 (Strassburg, Germany). Part : skull, not animal. 

1838. Amphisorex linneana Gray, Ann. Nat. Hist., n, p. 287, December, 

1838 (North Bothnia, Sweden). 

1839. [ Sorex fodiens] var. leucotis de Selys-Longohamps, Etudes de Mioro- 

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

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

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

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

nudum. 
1857. Crossopus fodiens Blasius, SiLugethiere Deutschlands, p. 120 (part) . 
1868. Sorex fimbriatus and Crossopus fimbriatus Fitzinger, Sitzungsber. 

kais. Akad. Wissensch. Wien, Math.-Naturwiss. Clasre, lvii, pt. 1, 

p. 610 (Synonyms of " Crossopus daubentonii nigripes," wrongly 

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

Wagler, 1832. 
1868. Crossopus ciliatus, griseogularis Pitzinger, Sitzungsber. kais. Akad. 

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

(Chartres, Eure-et-Loire, Prance). 
1870. Sorex intermedius Cornalia, Catal. Descrit. Mamm. Ital., p. 27 (Hills 

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

Soc. Ital. Sci. Nat. e del Mus. Civ. Stor. Nat., Milano, xxxvin, 

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

1908. 
1899. Sorex alpinus var. longobardus Sordelli, Atti Soc. Ital. Sci. Nat. e 

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

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

p. 45, April 25, 1901 (Montrejeau, Haute-Garonne, Prance). Type 

in U.S. National Museum. 
1905. Crossopus ou Sorex ignotus Patio, Arch. Sci. Phys. et Nat., Geneve, 

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

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

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

in British Museum. 



NEOMYS 71 

1906. Neomys fodiens nanus Lydekker, Zoological Record, xlu (1905), 
Mamm., p. 34, August, 1906 (Accidental renaming of naias). 

1910. Neomys fodiens, N. fodiens naias and N. fodiens minor Trouessart, 
Paune Mamm. d'Europe, pp. 56, 57. 

Type locality. — Berlin, Germany. 

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

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

Measurements. — Average and extremes of ten specimens from 
Brunswick, Germany: head and body, 86 '6 (83-92); tail, 65 ■ 5 
(58-77) ; hind foot, 18 '5 (17-19). Average of three specimens 
from Montrejeau, Haute-Garonne, France : * head and body, 83 
(82-85); tail, 52 (50-60); hind foot, 17- 1 (16-18). Average 
and extremes of seven specimens from Luchon, Haute-Garonne, 
France: head and body, 75-7 (72-80); tail, 53-1 (47-58) ; hind 
foot, 16 - 4 (16-17). Average and extremes of five specimens 
from Porte, Pyrenees-Orientales, France : head and body, 93 
(83-96) ; tail, 61 (57-65) ; hind foot, 18-2(18-19). Average and 
extremes of nine specimens from Meiringen, Bern, Switzerland : 
head and body, 82-7 (80-86); tail, 66-2 (62-71); hind foot, 
19-3 (18-20). Average and extremes of ten specimens from 
Hatszeg, Hunyad, Transylvania : f head and body, 77 ■ 7 (72-82) ; 
tail, 60 (56-66) ; hind foot, 18-5 (18-19). For cranial measure- 
ments see Table, p. 75. 

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

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

Sweden : Bothnia, 1 ; Upland, 1 ; Medstugan, Jemtland, 1 ; Upsala, 1 
(U.S.N.M.) ; no exact locality, 1. 

Belgium : Waremme, Liege, 3 (U.S.N.M.). 

France : Guinea, Pas-de-Oalais, 2 ; Abbeville, So rime, 2 (B.M. and Paris ; 
the latter type of remifer Geoffroy) ; near Paris, 1 (Paris ; type of lineatus 
Geoffroy) ; Dinan, C6tes-du-Nord, 1 ; Cadillae-sur-Garonne, Gironde, 2 
(U.S.N.M.) : Montrejeau, Haute-Garonne, 3 (U.S.N.M.) ; Porte, Pyrenees- 
Orientales, 5 ; l'Hospitalet, Ariege, 2 ; Luchon, Haute-Garonne, 7 ; Bareges, 
Hautes-Pyrenees, 3 ; Barcelonnette, Basses-Alpes, 1 ; Chamonix, Haute- 
Savoie, 1 (U.S.N.M.) ; Cranves-Sales, Haute-Savoie, 1 ; Scientrier, Haute- 
Savoie, 1 (Mottaz) ; Etupes, Doubs, 6 (Mottaz) ; no exact locality, 1. 

Germany : Brunswick, 10 (U.S.N.M.); Saxony, no exact locality, 1 
(U.S.N.M.); Bahrenberg, Harz Mts., 1 (U.S.N.M.): Frankfort, Wies- 
baden, 1 ; Niesky, Silesia, 1 ; Strassburg, 1. 

Austria-Hungary : Haida, Arva, Bohemia, 3 ; Hatszeg, Hunyad, 
Transylvania, 11. 

Switzerland : Geneva, 4 (Mottaz) ; Ohesieres, Vaud, 4 (Mottaz) ; Les 
Plans, Vaud, 3 (U.S.N.M.) ; Meiringen, Bern, 9 (U.S.N.M.) ; Grindelwald, 



* " Neomys fodiens minor " Miller. 

f "Neomys fodiens naias " Barrett-Hamilton. 



I .i INSECTIVORA 

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

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

Bemarhs. — There is much individual variation in colour as 
well as in external measurements and proportions,* though the 
size of the skull and teeth, as may be seen from the Tables, 
remains very constant. The degree of development of the fringes 
on feet and keel on tail is also variable, the depth of the keel in 
some individuals being about equal to diameter of tail, while in 
others it is too slight to be measured. This is partly, if not 
entirely, due to season, as the deepest keels are found in winter 
specimens. With regard to the variations in external measure- 
ments shown by the averages and extremes in a preceding 
paragraph, it is probable that they are much exaggerated by 
different methods in taking the measurements. This is well 
illustrated by the two series of Pyrenean specimens measured 
by A. Robert at an interval of six years. While much 
material has been examined it is not wholly satisfactory, con- 
sisting chiefly of small lots taken b}' many different collectors. 
Eventually it may be necessary to recognize certain Continental 
forms such as minor, naias, and the Scandinavian linneana, as 
distinct from true fodiens ; but for the present, in view of the 
uncertainties concerning external measurements and the striking 
similarity of skulls from the entire range of the animal, there 
seems to be no other alternative than to regard all the Con- 
tinental water-shrews with keeled tail as belonging to a single 
race. 

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

way. Waldo (c & p). 

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

2200 ft. (N. F. Tice- (p). 

hurst.) 

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

b/jem (N. F. Ticehttrst.) (p) . 

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

berg.) 

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

550 m. (G.Kolthoff.) 

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

France. 

<J, 2 9. Porte, Pyrenees-Orion- 0. Thomas (p). 8. 9. 1. 39-41. 

tales, 1600-1700 m. 

France. (A. Robert.) 



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



<5. 


L'Hospitalet, Ariege, 




1450 m. (A. Bobert.) 


?. 


L'Hospitalet, Ariege. 


<J,2. 3 9. 


Luchon.Haute-Garonne, 




600-900m. (A.Robert.) 


2<5. 


Bareges, Hautes-Pyre- 




nees, 1300-1500 m. 


9. 


Baroelonnette, Basses- 




Alpes. (C. Mottaz.) 


1. 


Cranves-Sales, Haute- 




Savoie. 


1. 


Abbeville, Somme. 


1. - 


Frankfort, Wiesbaden, 




Germany. 


9. 


Niesky, Silesia, 181 m. 




(W. Baer.) 


1. 


Strassburg, Alsace. 




(O. Mottaz.) 


2 9. 


Haida, Bohemia, 




Austria. 


1. 


Hatszeg, Hunyad, Tran- 




sylvania, 1500 ft. 




Hungary. 


6<J,49. 


Hatszeg, Transylvania, 




1500-2000 ft. 


2. 


Miirren, Bern, Switzer- 




land. 


4 9. 


Ziiberwangen, St.Gallen. 




(E. Zollikofer.) 


9. 


Faido, Ticino. 


3 6. 


Locarno, Ticino. 


]. 


Switzerland. 


(J, 2 9. 


Busalla, Liguria, Italy. 


lal. 


Vallombrosa, Florence. 



NEOMYS 

0. Thomas (p). 

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

G. S. Miller (c). 

0. Thomas (p). 

A. Robert (p). 

V. Baillon (c). 

Dr. Dieffenbach (p). 

Lord Lilford (p). 

0. Thomas (p). 

Lord Lilford (p). 

G. G. Danford (c). 

0. G. Danford (c). 

W. Gurtner (p). 

0. Thomas (p). 

0. Thomas (c & p). 
0. Thomas (c & p). 
Purchased (Parrey s) . 
0. Thomas (o & p). 
Dr. G. Cecconi (p). 



73 



8. 9. 1. 42. 



8. 8. 4. 139. 
6. 4. 1. 12. 14- 

18. 
8. 8. 4. 136- 

137. 
8. 8. 10. 13. 

5. 4. 9. 2. 

54. A. 

47. 1. 8. 46. 

99. 1. 9. 8. 

8. 8. 10. 12. 

11. 1. 1. 148- 

149. 
3. 2. 2. 1. 



3. 11. 8. 2-11. 
92. 10. 5. 1-2. 

4. 4, 5. 48-51. 

5. 8. 2. 20. 
5. 8. 2. 8-10. 
46. 6. 15. 61. 
5. 8. 3. 16-18. 
1. 8. 2. 1. 



Neomys fodiens bicoloe Shaw. 

1791. Sorex bicolor Shaw, Naturalists' Miscellany, n, pi. 55, named in 

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

England). 
1837. Amphisorexpennanti Gray, Proc. Zopl. Soc, London, p. 125 (England). 
1840. Crossopus sowerbyi iBonaparte, Iconogr. Faun. Ital., i, fasc. 29, in 

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

7th ser., xv, p. 508, May, 1905. 
1910. Neomys fodiens ciliatus Trouessart, Faune Mamm. d'Europe, p. 56. 

Type locality. — Oxford, England. 

Oeographical distribution. — Great Britain. 

Diagnosis. — Similar to Neomys fodiens fodiens, but underparts 
usually washed with wood-brown. Condylobasal length of skull 
usually less than 21 mm. 

Measurements. — External measurements of two adult females 
from Scotland (Grantown-on-Spey and Gortachy, Forfar) : head 



74 



INSEOTIVORA 



and body, 83 and 71 ; tail, 57 ■ 5 and 53 ■ 5 ; hind foot, 18 and 17. 
Adult male from Wellington, Somerset : head and body, 75 ; 
tail, 48 ; hind foot, 17. Adult male from Yalding, Kent : head 
and body, 82 ; tail, 52 ; hind fool;, 17. For cranial measurements 
see Table, p. 77. 

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

Scotland : Gordonstown, Elgin, 1 ; Dunphail, Elgin, 1 ; Grantown-on- 
Spey, Elgin, 1 (Wilson) ; Cortaohy, Elgin, 1 (Wilson) ; Cromlix, Stirling- 
shire, 1 ; Knibruok, Stirlingshire, 1 ; Aberdeen, 1 ; Penkill, Ayrshire, 1. 

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

Enqland : Mill-on-Tyne, Northumberland, 1 ; Leeds, Yorkshire, 1 ; 
Wellington, Somerset, 1 ; Great Grimsby, Lincolnshire, 2 ; Shaftesbury, 
Dorset, 1; Halesworth, Suffolk, 1; Cambridgeshire, 2; Thornhaugh, 
Northampton, 1 ; Podington, Wellingborough, Northamptonshire, 1 ; 
Henley-on-Thames, Oxfordshire, 2 ; Drinkstone Park, Bury St. Edmunds, 
Suffolk, 1 ; Woolpit, Bury St. Edmunds, Suffolk, 3 ; Halesworth, Suffolk, 1 ; 
Cambridgeshire, 2 ; Epping, Essex, 1 ; Banstead, Surrey, 3 ; Esher, Surrey, 
1; Garratt Park Lake, Surrey, 1; Godalming, Surrey, 1; Tillingbourne 
Stream, Surrey, 1 ; New Forest, Hampshire, 4 ; BasiDgstoke, Hampshire, 2 ; 
Hampshire, no exact locality, 2 ; Yalding, Kent, 1. 



6. 


Dunphail, Elgin. 


Miller Collection. 


7. 7. 7. 2848. 


lal. 9 


Aberdeen, Scotland. 


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


10. 11. 28. 1. 


1. 


Penkill, Ayrshire. 


E. R. Alston (p). 


79. 9. 25. 8. 


9. 


Fishguard, Pembroke- 
shire, Wales. 


Y. H. Mills (c & p). 


11. 1. 3. 376. 


1. 


Wales. 


S. Stokes (c & p). 


48. 9. 24. 4. 


9. 


Mill-on-Tyne, North- 


Bev. H. H. Slater 


11. 1. 3. 375. 




umberland, England. 


(c & P). 




lal. 


Wellingborough, North- 
amptonshire. 


R. R. Orlebar (c&p). 


84. 4. 21. 1. 


1. 


Halesworth, Suffolk. 


Bev. W. R. Tate 
(c & p). 


87. 6. 7. 1. 


9. 


Cambridgeshire. 


3. Baker (c&p). 
E. B. Alston (p). 


39. 9. 29. 26. 


1. 


Cambridgeshire. 


79. 9. 25. 7. 




(J. Baker.) 






1. 


Epping, Essex. 


J. Baker (p). 


40. 4. 11. 9. 


2al. 


Henley, Oxfordshire. 


W. Boyal Dawson 
(c & p). 


11. 1. 3. 377. 


lal. 


Bury St. Edmunds, 
Suffolk. » 


J. H.Powell (c&p). 


83. 8. 8. 1. 


3al. 


Bury St. Edmunds, 
Suffolk. 


Duncan Parker (p). 


84. 5. 16. 1-3. 


cSal. 


Esher, Surrey. 


E.Dalgleish(c&p). 


11. 1. 3. 378. 


9al. 


Godalming, Surrey. 


W. T. Blanford 
(c & P). 


94. 8. 5. 2. 


<5,3 9. 


New Forest, Hampshire. 


Miller Collection. 


7. 7. 7. 2845- 
2849. 


2 9. 


Basingstoke, Hampshire. 


Miller Collection. 


7. 7. 7. 4470- 
4471. 


2al. 


Hampshire. 


E. Bartlett (p). 


74. 11. 24. 1-2. 



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78 



INSECTIVOKA 



NEOMYS MILLERI Mottaz. 



1907. Neomys milleri Mottaz, Mem. Soo. Zool. de France, xx, p. 22, 

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

Type locality. — Chesieres, Alpes Vaudoises, Switzerland. 
Altitude, 1,230 m. 

Geographical distribution. — Pyrenees and Alps ; Hungary. 
Details of distribution not yet known. 

Diagnosis. — Slightly smaller than Neomys fodiens ; tail with 
keel absent, or represented by a slight lengthening of hairs on 
under side of terminal third only ; fringes on sides of feet not 
conspicuously developed; hind foot usually less than 17 mm.; 
tail usually less than 55 mm. ; lachrymal foramen opening over 
point of contact between m 1 and m 2 ; anterior upper incisor 
slender. 

External characters. — Similar to Neomys fodiens, except that 
the feet are less fringed and the tail lacks the definite keel on 
under side. When unworn the hairs on ventral surface of tail 
are slightly longer than those on upper side, and on terminal 
third this elongation is occasionally sufficient to produce a 
rudimentary keel ; but there is never any trace of a well- 
defined ridge extending to base of tail. Caudal annulations 
slightly less distinct than in Neomys fodiens. Mammse : a 1—1, 
i 4-4 = 10. 

Colour.' — The oolour is similar to that of typical specimens of 
Neomys fodiens fodiens. Among the skins examined none shows 
any noticeable wash of brown or suffusion of buff on underparts. 

Skull and Teeth. — The skull closely resembles that of Neomys 
fodiens, but may be distinguished by its smaller general size and 
relatively lower, more slender rostrum ; lachrymal foramen over 
point of contact of m? and m 2 . Teeth smaller throughout, a 
difference especially noticeable in the anterior upper incisor. 
Upper unicuspids with crowns longer and narrower than in 
Neomys fodiens, the long posterior portion especially noticeable 
in lateral view by comparison with height of cusp ; cingula 
not so well developed as in the related species, and seldom, if 
ever, forming a postero-external cusp. 

Measurements. — External measurements of type : head and 
body, 76; tail, 59; hind foot, 16; hind foot, including claws, 
17. Average and extremes of ten specimens from the type 
locality: head and body, 77 "4 (71-87); tail, 53-7 (50-59); 
hind foot, 15 '8 (15-16-2). Average and extremes of six 
specimens from Untervatz, Grisons, Switzerland : head and body, 
79-3 (75-82); tail, 46-8(45-51); hind foot, 14-16 (14-15-4). 
Two adult males from Locarno, Ticino, Switzerland : head and 
body, 86 ; tail, 50 and 53 ; hind foot, 16 and 17. Average and 
extremes of three adult females from 1' Hospitalet, Ariege, 



NEOMTS 79 

France: head and body, 78 (75-80); tail, 50-6 (47-53); hind 
foot, 15 - 7 (15-4—16). Adult male from Bareges, Hautes- 
Pyrenees : head and body, 76; tail, 56; hind foot, 16 '4. For 
cranical measurements see Table, p. 80. 

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

Switzerland : Near Geneva, 1 ; Lausanne, Vaud, 2 (U.S.N. II.) ; 
Chesieres, Alpea Vaudoises, 10 (B.M. and Mottaz) ; Meiringen, Bern, 2 
(U.S.N.M.) ; Untervatz, Grisons, 6 (U.S.N.M.) ; Zuberwangen, St. Gallen, 3 
(B.M. and U.S.N.M.) ; Sorengo, Ticino, 1 (U.S.N.M.) ; Locarno, Ticino, 2. 

Italy : Porlezza, Como, 1 (Mottaz). 

Austbia-Hungaby : Zuberec, Northern Hungary, 1. 

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

Remarks. — Though superficially resembling Neomys fodiens 
the round-tailed water-shrew is easily recognizable by the 
complete absence of any true keel on the tail. The slight 
elongation of the hairs in median region which sometimes occurs 
on terminal third of tail never results in the formation of a 
definite keel similar to that of the related species ; while no 
trace of such a structure is ever present on basal half of tail. 
The smaller hind foot is usually diagnostic, though in this 
character there is a slight overlapping between the two species. 
Though in all essential respects true water-shrews, the members 
of the round-tailed group (N. anomalus of Spain, N. milleri of 
the Pyrenees, Alps and northern Hungary, and N. teres of 
Asia Minor) are less highly specialized than Neomys fodiens. 
Their habits, however, appear to be essentially similar to those 
of the common species, as the two animals are found together in 
localities where their ranges coincide.* 

2 9. L'Hospitalet, Ariege, G. S. Miller (c). 8.8.4.138,140. 

Prance. 
9. Chesieres, Alpes Vau- C. Mottaz (p). 6. 2. 6. 4. 

doises, 1200 m. Swit- 
zerland. 
9. Ziiberwangen.St.Gallen, O. Thomas (p). 4. 4. 5. 52. 

Switzerland. 

(E. H. Zollikofer.) 
2 1. Locarno, Ticino, Swit- O. Thomas (c & p). 5. 8. 2. 6-7. 
zerland. 
1. Zuberec, Hungary. Budapest Museum 94. 3. 1. 26. 

(e). 



* At two localities in the Pyrenees (l'Hospitalet, Ariege and Bareges, 
Hautes-Pyrenees), where I found the two animals occurring together, both 
were taken alternately in the same traps set at the edge of mountain 
streams. In the original description of the species, however, Mr. Mottaz 
observes (p. 23) that according to his observations the round-tailed animal 
is the more inclined to wander away from the immediate vicinity of water. 
The habits of N. anomalus in North Central Spain appear to be as strictly 
aquatic as those of N. fodiens. 



80 



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PACHYURA 



81 



NEOMYS ANOMALUS Cabrera. 



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

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

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

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

Geographical distribution. — Northern and central Spain. 

Diagnosis. — Similar to Neomys milleri, but with, longer tail 
(usually over 55 mm.), and larger hind foot (16 '8 to 18 mm.). 
Mammse : a 1 — 1, i 4 — 4 = 10. 

Measurements. — -Type specimen, male (from Cabrera) : head 
and body, 73 ; tail, 60 ; hind foot, 17 • 5. Average and extremes 
of nine specimens from Silos, Burgos, Spain : head and body, 
83-2 (76-88); tail, 60 (56-61); hind foot, 17-4 (17-18). 
Adult male and adult female from Barracas, Castellon, Spain : 
head and body, 99 and 100; tail, 53 and 52; hind foot, 17 
For cranial measurements see Table, p. 82. 

Specimens examined. — Twenty-two, from the following localities in 
Spain : Silos, Burgos, 9 ; La Granja, Segovia, 2 ; Barracas, Castellon, 2 ; 
Lerida, 9. 

"Remarks. — The Spanish -water-shrew is well differentiated 
from Neomys milleri by its longer tail and larger hind foot. As 
yet no member of the keel-tailed group has been found in the 
Iberian Peninsula, though it is probable that N. fodiens occurs in 
the region north of the Ebro, and perhaps also in the Asturias. 

2 S, 7 ?. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 22-30. 

2 al. La Granja, Segovia. M. delaEscalera (c). 8.7.30.6-7. 

6,9. Barracas, Castellon. O. Thomas (p). 8.2.9.41-42. 
(JV. Gonzales.) 

Genus PACHYURA de Selys-Longchamps. 

1839. Pachyura de Selys-Longchamps, Etudes de Micromamm., p. 32 

(Sorex etruscus Savi). Sub-genus of Crocidura. 
1857. Pachyura Blasius, Saugethiere Deutschlands, p. 147 (Sub-genus of 

Crocidura). 
1897. Plerodus Schulze, Mamm. Europ., in Helios, Abhandl. n. Vortraga 

Gesammtb. Naturwiss., xiv, p. 90 (Crocidura suaveolens Blasius 

= Sorex etruscus Savi). 

Type species. — Sorex etruscus Savi. 

Geographical distribution. — Africa and warmer portions of 
Asia ; in Europe confined to the Mediterranean region. 

Diagnosis. — Like Crocidura (p. 86), but with upper uni- 
cuspids 4-4, the dental formula : i ^|, c j=J, pm ^, »y= 30. 

Remarks. — Although perhaps not forming a natural group, 
the 30-toothed species of Crocidura may for convenience be 

6 



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PACHYURA 83 

treated as members of a distinct genus. The only European 
species is immediately recognizable by its excessively small size. 

PACHYURA ETRUSCA Savi. 

1822. Sorex etruscus Savi, Nuovo Giorn. de' Letterati, Pisa, I, v. 60 (Pisa 

Italy). '* K 

1835. [Sorex] pachyurus Kiister, Isis, p. 77 (Cagliari, Sardinia). 
1841. Pachyura etrusca Bonaparte, Ioonogr. Faun. Ital., i, Indice distrib. 
1857. Grocidura suaveolens Blasius, Saugethiere Deutsohlands, p. 147. 
1910. Pachyura etrusca Trouessart, Faune Mamm. d'Europe, p. 43. 

Type locality. — Pisa, Italy. 

Geographical distribution. — Italy and adjoining portions of the 
Mediterranean region.* Limits of range not known. 

Diagnosis. — Much smaller than Grocidura russula (head and 
body, 35-40 mm. ; hind foot, 7 ■ 6-8 mm.), but with tail (25- 
30 mm.) relatively longer, its ratio to head and body about 
70 ; skull excessively small, its condylobasal length only about 
12-8 mm., the dorsal profile nearly straight from front of nasal 
to back of parietals, usually a little concave in interorbital 
region, the brain-case narrow but even more flattened than in 
Grocidura leucodon and C. mimula ; teeth not conspicuously 
different from those of the smaller European species of Crocidura, 
except for the presence of the fourth upper unicuspid, and their 
much smaller size (maxillary tooth-row about 6 mm.). 
r External characters. — In general external form Pachyura 
etrusca does not differ notably from Crocidura russula except in 
the relatively longer tail. The weight of a full grown individual 
is, however, probably not more than one-fourth or one-third that 
of an adult G. russula, and the head and feet are equally small 
in proportion, a character by which Pachyura etrusca mav be 
easily distinguished from young individuals of the larger animal. 
Mammse, i 3 — 3 = 6. 

Colour. — Upper parts a uniform slaty brown, perhaps best 
described as drab-grey washed with light bister, the individual 
hairs slate-grey at base. Underparts faintly contrasted, rather 
pale drab-grey with silvery reflections in certain lights. Feet 
dull light slaty grey not noticeably contrasted with back. Tail 
like back above, usually becoming somewhat darker toward tip, 
lighter and more like belly below, but with no evident contrast 
between the colours of the two surfaces. 

Skull. — Apart from the small size the skull differs from that 
of its European allies in several important characters. Most 
noticeable among these is the extreme flatness of the dorsal 
profile, which is essentially straight from nares to occipital, 
usually a little concave in interorbital region, while in the 
European forms of Crocidura it is usually a little convex 
throughout, and never evidently concave in interorbital region. 

* Spain. See footnote, p. 86.— O. T. 

G 2 



84 INSECTIVOKA 

The brain-case is excessively flattened, relatively more so than in 
Crocidura mimula, so that there is less contrast between its depth 
and that of rostrum. This flattening is not accompanied by any 
increase in breadth of brain-case, the ratio of 
a^Z^Psi which to condylobasal length is about 46, 

Jil^j^ slightly less than in Crocidura mimula. Ante- 

"* orbital foramen relatively large, its position 

normal. Lachrymal foramen over posterior 
half of m 1 . Palate relatively short and wide. 
Mesopterygoid space relatively longer than in 
the European forms of Crocidura, its anterior 
border on line with posterior edge of third 
jfio. le. molar instead of distinctly behind it, its lateral 

FaCh xT£™ USCa ' borders nearly parallel, though slightly con- 
verging posteriorly ; hamular excessively deli- 
cate, bowed outward and upward, its length relatively greater 
than in C. russula, and about equal to greatest breadth of fossa. 
Mandible not peculiar except for its small size (length about 
7 mm. or less), and very delicate structure. 

Teeth. — While in general, and aside from the presence of 
the fourth unicuspid, the teeth do not differ materially from 
those of the smaller European Crociduree except in size, they 
show several peculiarities in form. The anterior upper incisor 
projects more strongly forward, and has the anterior cusp rather 
shorter than in the species of Crocidura, while the posterior cup 
is more distinctly separated from the cingulum, these two 
characters together imparting to the tooth a form somewhat 
suggesting that assumed in Sorex. First unicuspid relatively 
larger and higher than in Crocidura russula, its cingulum less 
curved, and its width distinctly exceeding that of palate instead 
of barely equalling it. Contrast in both height and crown area 
of first and second unicuspids greater than in C. russula, the 
area of second about one-third that of first instead of evidently 
more than one-third. Third unicuspid slightly larger than second. 
Fourth about half as large as third, crowded inward from the 
tooth-row, but visible from the outside through space separating 
third unicuspid from large premolar, the width of this space 
rather more than half diameter of crown of fourth unicuspid. 
Large premolar as in C. leucodon, except that cutting blade is 
shorter and even higher, and posterior border of crown is less 
concave. Upper molars with hypocones less distinct than in 
C. russula, but otherwise not showing any tangible peculiarities. 
Mandibular teeth essentially as in Crocidura russula and C. leu- 
codon, but posterior section of third molar even more reduced. 

Measurements. — Average and extremes of six specimens from 
Florence, Italy: head and body, 38 '5 (36-42); tail, 27 '1 
(26-28); hind foot, 7-7 (7 "6-7 -8). Adult male and female 
from near Turin, Italy : head and body, 42 and 40; tail, 28 and 
27 ; hind foot, 7 ■ 8 and 7 ' 4. Adult male and female from near 



PACHYURA 



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86 



IXSECTIVORA 



1 al. 
2al. 
lal. 


Pisa, Italy. 

Tuscany. 

Italy. 


Skeleton. 
<5, 9 al. 


Italy. 
Sicily. 


lal. 


Sardinia. 


lal.* 


Lamia, Greece, 



Genoa, Italy : head and body, 41 and 42 ; tail, 26 and 28 ; hind 
foot, 7 ■ 6 and 8 ' 0. Adult female from Sassari, Sardinia : head 
and body, 41 ; tail, 29 ; hind foot, 7*8. For cranial measure- 
ments see Table, p. 85. 

Specimens examined. — Twenty- nine, from the following localities : — 

Italy : Near Turin, 2 (Genoa) ; Perti, Finalborgo, 2 (Genoa) ; near 
Genoa, 5 (U.S.N.M. and Genoa) ; Pisa, 5 (B.M. and Mottaz) ; Florence, 6 
(U.S.N.M. andilottaz) ; Tuscany, 2; Borne, 1 (Genoa) ; no exact locality, 2. 

Sicily : No exact locality, 2. 

Sakdiuta: Assuni, 1 (U.S.X.M.). No exact locality, 1. 

Greece : Lamia, 1. 

Marquis G. Doria (p). 69. 3. 4. 1. 

(No history). 

Zoological Society's 55. 12. 26. 295. 

Collection. 

Purchased. 58. 10. 21. 13. 

Purchased (Parzu- 52. 2. 26. 22-23. 

daki). 

Hon. N. C. Boths- 11. 10. 1. 1. 

child (p). 

E. MacDoneU (p). 8. 7. 22. 1. 

Genus CROCIDURA Wagler. 

1832. Crocidura Wagler, Isis, p. 275 [Sorex leucodon Hermann). 

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

1869. Leucodon Fatio, Faune Vert. Suisse, I, p. 132 (substitute for 

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

Gesammtb. Naturwiss., xiv, p. 90 (Sorex leucodon Hermann and 

S. araneus Schreber = S. russulus Hermann). 

Type species. — Sorex leucodon Hermann. 

Geographical distribution. — Africa and warmer portions of 
Europe and Asia, including the Malay Archipelago ; in Europe, 
north to northern Holland and central Germany, west to the 
Atlantic coast and the Channel Islands. Absent from Great 
Britain and Ireland. 

Characters. — Upper unicuspid teeth 3-3 (dental formula : 
i !t 3 , c J^J, pm !=1, m jj=| = 28 ; posterior lobe of upper incisor 
noticeably less than half as high as main cusp ; anterior lower 
incisor without lobes on cutting edge ; third lower molar with hypo- 
conid and entoconid coalesced, so that form of tooth is strikingly 
different from that of other molars, its crown 4-cusped instead 
of 5-cusped ; second lower unicuspid without rudimentary second 
cusp and commissure ; teeth white throughout ; skull heavier and 
more robust than in Sorex and Neomys, with conspicuously deeper 
rostrum and less contrast between width of brain-case and that 
of anterior portion ; rudimentary zygomatic process of maxillary 
obsolete ; external form rather heavy ; ear rising noticeably 

* i, 2 9. llarismas, Lower Guadal- A. Chapman (c&p). 11.12.19.1-3. 
quivir, Spain. 
Beceived December, 1911.— 0. T. 



CROGIDURA o I 

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

Remarks. — Though not highly modified in external pecu- 
liarities the genus Grocidura is the least primitive group of 
shrews occurring in Europe. This is indicated by the reduced 
number of upper unicuspids, but is more clearly shown by the 
highly modified form of the third lower molar and the completely 
unicuspid character of the lower premolar. Among the European 
members of the family Soricidee the species of Grocidura may 
at once be recognized by their moderate size, entirely white teeth, 
large ears, and by the presence of long, loosely spreading hairs 
scattered over the surface of the tail. Badly prepared specimens, 
in which the true characters are obscured, may often be detected 
among skins of Sorex by the noticeable silvery reflections on 
hairs of back. Eight species are now known to occur in western 
Europe, while the number of forms thus far described from other 
portions of the range of the genus is not far from one hundred. 

KEY TO THE EUROPEAN FORMS OP GROCIDURA. 

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

Hind foot about 14 mm. (Sicily) C. caudata, p. 110. 

Hind foot 11 to 12' 5 mm. 

Upper tooth-row about 8 - 6 mm. (Corsica) C. cyrnensis, p. 111. 

Upper tooth-row about 8 mm. (Balearic Islands)... C. balearica, p. 112. 
Tail about half as long as head and body, the ratio 
normally varying from 40 to 60. 
Condylobasal length of skull 16 to 17 • 6 mm. 
Second upper uniouspid with crown area about 

equal to that of third (Crete) C. canese, p. 109. 

Second upper unicuspid with crown area distinctly 

less than that of third C. mimuia, p. 94. 

Colour tending to be pale and greyish (Basque 

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

Colour tending to be dark and brownish. 
Brain-case not decidedly flattened (Charente, 

Prance) C. m. iculisma, p. 98. 

Brain-case decidedly flattened (entire range 

of species, western portion excepted)... C. m. mimiUa, p. 95. 
Condylobasal length of skull 17 '6 to 20 • 4 mm. 
(usually more than 18 mm.). 
Brain-case about half as high as wide or less. 
Animal conspicuously bicolor ; condylobasal 
length of skull usually more than 19 mm. 

(Central Europe, south into Italy) G. leucodm, p. 88. 

Animal obscurely bicolor ; condylobasal length 

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

Brain-oase more than half as high as wide C. russula, p. 99. 

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

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

Condylobasal length of skull usually 18 to 19 mm. 

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

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



88 



INSECTIVORA 



CROCIDURA LEUCODON Hermann. 



1780. Sorex leucodon Hermann in Zimmermann, Geogr. Gesoh., n, 

p. 382 (vicinity of Strassburg, Germany). 

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

1782. Sor[ex~] leucodon Hermann, Tabula Affinitatum Animalium, p. 79 

(footnote). 
1792. Sorex albipes Kerr, Anim. Kingd., p. 208 (based on Pennant's account 

of S. leucodon Hermann). 
1832. Croc[idura] leucodon Wagler, Isis, p. 275. 
1857. Crocidura leucodon Blasius, Saugethiere Deutschlands, p. 140. 
1869. Leucodon microurus Patio, Paune Vert. Suisse, i, p. 173 (Substitute 

for leucodon). 
1897. [Crocidura] leucodus Schulze, Mammalia Europaea, p. 18 (Substitute 

for leucodon). 
1910. Crocidura russula leucodon Trouessart, Paune Mamm. d'Europe, p. 44. 

Type locality. — Vicinity of Strassburg, Germany. 

Geographical distribution. — Central Europe, from Belgium to 
Hungary ; south into Italy. Not known from the Iberian 
Peninsula. 

Diagnosis. — Size large (among the European species) ; hind 
foot, 1 1 to 1 3 mm. ; condylobasal length of skull, 1 8 • 4 to 20 mm. ; 
upper tooth-row, 8 • 8 to 9*0 mm. ; tail short, its actual length 
28 to 38 mm., its ratio to head and body usually ranging from 
38 to 43 ; skull with brain-case noticeably depressed, its height 
less than half its width ; large upper premolar with antero- 
external cusp rather large, its height usually greater than that 
of first unicuspid ; colour of underparts whitish, strongly con- 
trasted with rather slaty brown of back, the line of demarcation 
along sides well defined. 

External characters.- — Fur shorter and more dense than in 
Sorex araneus, the length of hairs at middle of back about 
3 • 5 mm. in summer, 5 mm. in winter, its texture not specially 
modified ; a few slightly elongated hairs (8 mm.) on flanks and 
across rump. Eyes small and inconspicuous ; ears small but 
rising conspicuously above fur, the two well developed valves a 
conspicuous feature of the ear in freshly killed specimens. Feet 
not peculiar in form ; less slender than in Sorex araneus, finely 
pubescent on dorsal surface and on lateral portions of posterior 
half of sole ; fingers proportioned as in S. araneus, but graduation 
less, especially in hind foot ; pads 6-6, those on palm distinct 
though somewhat crowded, the surface of palms and soles finely 
rugose between the pads. Tail less slender than in Sorex and 
Neomys, nearly terete or with under side somewhat flattened, 
its length equal to about half that of head and body, its hairs of 
two kinds : (a) finely appressed hairs less than 1 mm. in length, 
nearly concealing the annulations and forming a very slight 
pencil ; and (6) loosely spreading hairs about 5 mm. long, rather 
thickly sprinkled among the others ; annulation ill-defined, 
about 35 to the centimeter at middle. Mammas, i 3 — 3 = 6. 



CROCIDUEA 



89 



Colour. — Upper parts varying from a slaty drab to dull 
russet, the hairs with a slight metallic gloss and with silvery 
reflections, which in certain lights produce an evident effect of 
fine speckling. Underparts and inner surface of limbs buffy 
white, dulled to a varying degree by the slaty under colour, the 
chin and throat often suffused with cream-buff. Line of 
demarcation sharply defined, extending just below ear and eye 
and alongside of muzzle to middle of pad. The two extremes of 
colour probably represent ill-defined dichromatic phases. Most 
of the specimens examined are in some degree intermediate. 
Feet dull whitish, often irregularly clouded with drab. Tail 
sharply bicolor, whitish below, concolor with back above, the 
longer hairs silvery grey. 

Skull. — The skull is slightly larger than that of Sorex araneus 
and noticeably more heavily built, particularly that portion 
lying in front of brain-case, all of which is both broadened and 
deepened, so that the general outline tapers less conspicuously 
from behind forward, whether skull is viewed from above or 
from the side. Brain-case less well marked off from interorbital 
region than in the European species of Sorex and Neomys, its 
main sutures closing early in life. Brain-case 
slightly longer than broad, its posterior out- /7 ~~~~ — — 
line rounded but broken by the slightly Lk^-i^S^p^ 
projecting points of the condyles, its antero- 
external border straight, sharply angled in 
front ; sagittal crest low but evident in adult 
skulls, meeting the complete lambdoid crests 
posteriorly. Depth of brain-case at middle 
slightly but constantly less than half greatest 
breadth (see fig. 20, page 100). Dorsal profile 
with a slight concavity at front of brain-case 
and slight convexity over middle of rostrum. v 17 

Nares squarely truncate posteriorly, the Cm idu.m lencoHon. 
lateral wall abruptly angled near middle. Nat. size. 

Anteorbital foramen relatively smaller than in 
Sorex. araneus, and region between it and edge of alveolus dis- 
tinctly wider. Lachrymal foramen over metastyle of m 1 Plate 
forming outer wall of anteorbital canal nearly three times as wide 
as lachrymal foramen. Angular shelf -like region over posterior 
molars broader and more prominent than in the European species 
of Sorex, but rudimentary zygomatic process of maxillary reduced 
to the merest trace. Mesopterygoid fossa as in Sorex araneus ; 
floor of brain- case between tympanic bones narrower and with 
distinct median ridge. 

Teeth. — Dentition noticeably heavier than in the European 
species of Sorex and Neomys, the difference in general aspect 
heightened by the absence of brown colouring matter on points 
of cusps. Anterior upper incisor with main cusp long and 
slender, abruptly hooked downward; basal lobe low and 




90 



INSECTIVOBA 




FIG. 18. 

Croeidura leucodon. Anterior 
teeth in profile. X 5. 



triangular, its height less than half that of main cusp and only 
a little more than half that of first unicuspid tooth. General 
form of first incisor not unlike that of 
Neomys fodiens, but with even more 
contrast between height of the two 
cusps. Anterior lower incisor simple, 
rather robust, the shaft slightly taper- 
ing, the point slightly bent upward, the 
cutting edge without lobes. Upper 
unicuspids robust, strongly contrasted 
in size. First about double the height 
of second and third, the points of which 
are nearly in line with that of basal 
lobe of anterior incisor. Its crown is 
somewhat longer than broad, with main 
axis slightly oblique to that of tooth-row. In lateral view it is 
nearly triangular, with anterior border about half as long as upper 
and lower. Cutting edge essentially 
as in Neomys, but better developed 
and reaching posterior border of 
tooth somewhat outside of middle. 
Crushing surface well developed, 
occupying about one-third area of 
crown. Second and third unicus- 
pids approximately equal, in both 
height and crown area, to basal 
lobe of anterior incisor, their crowns 
distinctly narrower as well as much 
shorter than that of first 
unicuspid, their cutting 
ridge and crushing sur- 
face less well differen- 
tiated than in first. 
Second smaller than 
third, its crown about 
as broad as long. Third 
separated from large pre- 
molar by a slight space, 
its crown longer than 
broad, its cusp about 
equal in height to para- 
cone of large premolar. 
Lower unicuspids essen- 
tially alike in form, the 
second differing from that 
of Sorex and Neomys in 
the complete absence of the rudimentary second cusp. First low, 
narrow and long ; second high, its crown about as broad as long. 
Upper cheek-teeth with crowns narrower than in Sorex and 




Fig. 19. 
Croeidura leucodon. Teeth X 10. 



CROCIDURA 91 

Neomya, the emargination of posterior border deeper, particularly 
in pm*, and hypocones more distinct. Large upper premolar 
with antero-external cusp (paracone) well developed, its height 
usually about equal to that of third unicuspid or somewhat 
more, the distance from its point to well-defined angle in 
cingulum over anterior root of tooth distinctly more than half 
length of anterior border of main cusp ; posterior cutting blade 
high, the angle formed between its edge and the moderately 
projecting point of main cusp ill-defined and obtuse. Third 
upper molar actually as well as relatively smaller than in Sorex 
araneus, but containing the same elements. Lower cheek-teeth 
essentially as in Sorex araneus except for the peculiarities of m 3 
characteristic of the genus. 

Measurements. — Average and extremes of nine specimens 
from Etupes, Doubs, France: head and body, 78 • 3 (70-87); 
tail, 35-3 (32-39); hind foot, 12-3 (12-13). Average and 
extremes of nine specimens from Untervatz, Grisons, Switzerland : 
head and body, 81 (77-85); tail, 31-6 (29-34); hind foot, 
12-5 (11 • 8—13). Average and extremes of four specimens from 
Lugano, Ticino, Switzerland: head and body, 80 (77-81); tail, 
32 • 2 (31-36) ; hind foot, 12-5(11- 8-13). Average and extremes 
of three specimens from Florence, Italy (in alcohol, bodies 
contracted) : head and body, 63 ■ 3 (63-64) ; tail, 34 • 3 (32-36) ; 
hind foot, 12-3 (11 -6-13). For cranial measurements see Table, 
p. 92. 

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

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

Prance: Dinan, Brittany, 1; Paris, 1 (U.S.N.M.); Manouville, 
Meurthe-et-Moselle, 1 ; Etupes, Doubs, 14 (B.M. and Mottaz). 

Germany : Brunswick, 1 (U.S.N.M.) ; Nussberg, Brunswick, 1 ; Frank- 
furt, Hessen-Nassau, 1 ; Taucha, Saxony, 1 (U.S.N.M.) ; Marxheim, 
Bavaria, 4 ; Ummerstadt, Thiiringen, 1 ; no exact locality, 1. 

Austria-Hungary : Hatszeg, Hunyad, Transylvania, 1. 

Switzerland : Geneva, 3 (Mottaz) ; Briinig, Bern, 1 (U.S N.M.) ; Mei- 
ringen, Bern, 4 (U.S.N.M.) ; Untervatz, Grisons, 15 (B.M. and U.S.N.M.) ; 
Grisons, no exact locality, 1 ; Lugano, Ticino, 4 (U.S.N.M.) ; Zuber- 
wangen, St. Gallon, 1. 

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

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



92 



INSECTIVORA 



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94 INSBCTIVORA 

2 9, 1. Esneux, Liege, Belgium Lord Lilford (p). 95. 1. 1. 1-2. 

(H. Gronvold.) 
i. Dinan, Brittany, Prance. G. Barrett-Hamilton 11. 1. 2. 158. 

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

Moselle. 
5 6, 4 9. Etupes, Doubs, 350 m. 0. Thomas (p). 8. S. 10. 33-41. 

(C. Mottaz.) 
2 9. Marxheim, Bavaria, Lord Lilford (p). 8.9.8.2-3. 

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

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

(*)■ 
1 al. Germany. Tomes Collection. 7. 1. 1. 45. 

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

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

Switzerland. 10. 8. 16. 9-13. 

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

(E. H. ZolUkofer.) 
9. Lugano, Ticino. 0. Thomas (p). 10. 8. 16. 15. 

(E. H. ZolUkofer.) 
9. Zilberwangen.St.Gallen. 0. Thomas (p). 10. 8. 16. 14. 

(E. H. ZolUkofer.) 
i. Rome. (G. Goli.) G. Barrett-Hamilton 11. 1 2. 103. 

CROCIDURA MIMULA Miller. 
(Synonymy under subspecies.) 

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

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

Colour. — Upper parts varying from a dull russet tinged with 
sepia to a dark hair-brown with a tinge of drab, the hairs with 
metallic gloss and silvery reflections ; underparts varying from 
dull ochraceous-buff to greyish cream-buff, never sharply defined 
from colour of back and sides. Feet buffy whitish, usually with 
some dark clouding. Tail g-reyish or brownish, very obscurely 
bicolor. 

Skull and teeth. — Except for their noticeably smaller size the 
skull and teeth closely resemble those of Grocidura leucodon, 
though the brain-case is less constantly flattened, sometimes 
assuming a form essentially like that in 0. russula. Plate forming 



CB0CIDT7RA 95 

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

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

Remarks. — Grocidura mimula differs from the other continental 
European members of the genus, in its small size, a character in 
which it is approached by the small races of C. russula, though 
not sufficiently to cause any confusion. In addition to the 
typical form two geographical races have been described, one from 
south-western France, the other from northern Spain, the 
status of neither of which is clearly understood. 

Ceociduea mimula mimula Miller. 

1839. ?? [Crocidura aranea] var. minor de Selys-Longchamps, Etudes de 

Micromamm., p. 35 (Silesia). 
1901. Crocidura mimula Miller, Proc. Biol. Soc, Washington, xiv, p. 95, 

June 27, 1901 (Ziiberwangen, St. Gallen, Switzerland). Type in 

U.S. National Museum. 

1901. Crocidura antipse Matschie, Sitz.-Ber. Gesellsch. Naturforsch. 

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

1902. [Crocidura'] minuta Lydekker, Zool. Record, xxxviii (1901), Mamm., 

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

Type locality. — Ziiberwangen, St. Gallen, Switzerland. 

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

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

Measurements. — External measurements of type : head and 
body, 72 ; tail, 33 ; hind foot, 11. A male and female from 
Untervatz, Orisons, Switzerland : head and body, 71 and 65 ; 
tail, 35 and 35 ; hind foot, 10 and 11. A male and female from 
Marxheim, Bavaria : head and body, 71 and 58 ; tail, 36 and 30 ; 
hind foot, 10 and 10. Average and extremes of four specimens 
from Haida, Arva, Bohemia : head and body, 59 ■ 7 (55-65) ; 
tail, 30 (28-32) ; hind foot, 11-5(11-12). A male from Gageni, 
Roumania, and female from Bustenari, Roumania : head and 
body, 64 and 66 ; tail, 31 and 34; bind foot, 11 and 11. Two 
adult males from Agay, Var, France : head and body, 64 and 66 ; 
tail, 35 and 34 ; bind foot, 11 and 11. Adult female from 
Viareggio, Italy : head and body, 70; tail, 38 ; hind foot, 11 "6. 



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98 INSECTIVORA 

Adult male and female from Corfu, Greece : head and body, 74 
and 75 ; tail, 44 and 44 ; hind foot, 1 1 ■ 8 and 11*4. For cranial 
measurements see Table, p. 96. 

Specimens examined. — Thirty-seven, from the following localities : — 
France : Abbeville, Somme, 1 ; Agay, Var, 2. 
Germany : Marxheim, Bavaria, 2. 

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

Boumania : Gageni, Prahova, 1 ; Bustenari, Prahova, 1. 
Bulgaria: Sofia, 1 (Andersen) ; Varna, 1 (Andersen). 
Switzerland : Ziiberwangen, St. Gallen, 3 (U.S.N.M. and Mottaz) ; 
Untervatz, Grisons, 2; Faido, Ticino, 1; Santa Margherita, Ticino, 3 
(Mottaz); Davesco, Ticino, 1 (U.S.N.M.); Lugano, Ticino, 2 (B.M. and 
Mottaz) ; Locarno, Ticino, 1. 

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

Greece : Corfu, 2. 

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

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

c5. Marxheim, Bavaria, Lord Lilford (p). 8.9.8.4. 

Germany. ( Wolter- 

storff.) 
<*, 9. Haida, Bohemia, Lord Lilford (p). 8.9.8.5-6. 

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

sylvania, Hungary. 
1 al. Tatra Mountains. Dr. B. Collett (p). 91. 1. 21. 2. 

9. Gageni, Prahova, Bou- Lord Lilford (p). 4. 4. 6. 11. 

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

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

land. 
<*. Locarno, Ticino. 0. Thomas (c & p). 5. 8. 2. 1. 

<5, 9. Untervatz, Grisons. 0. Thomas (p). 4.4.5.21-22. 

(E. H. Zollihofer.) 
9. Lugano, Ticino. 0. Thomas (p). 4.4.5.57. 

(E. H. Zollikofer.) 
9. Viareggio, Lucca, 5 m. 0. Thomas (c & p). 5. 8. 2. 21. 

Italy. 
6. Borne. (Goli.) G. Barrett-Hamilton 11. 1. 2. 97. 

(*)■ 
6. Corfu, 50 m. Greece. J.I. S.Whitaker (p). 8.10.1.8. 

(0. Mottaz.) 

Crocidura mimula iculisma Mottaz. 

1908. Crocidura mimula iculisma Mottaz, Bull. Soc. Zool. de Geneve, i, 

p. 119, April 30, 1908. Type in Mottaz Collection. 
1910. Crocidura mimula iculisma Trouessart, Faune Mamm. d'Europe, p. 47. 

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

Geographical distribution. — Known from the type locality 
only. 

Diagnosis. — Size as in C. mimula mimula or slightly smaller 
(hind foot, 10; condylobasal length of skull, 16); brain-case 
deep, nearly as in C. russula. 



CROCIDURA 99 

Measurements. — Type (from Mottaz) : head and body, 60 • 5 ; 
tail, 38*5; hind foot, 10 • 2. For cranial measurements see 
Table, p. 97. 

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

Remarks. — While this race appears to be distinct from true 
mimula, the material seen is insufficient to form the basis of any 
final opinion as to its status. 



Ceociduka mimula cantabra Cabrera. 

1908. Crocidura cantabra Cabrera, Bol. Eeal Soo. Espafi. Hist. Nat., vm, 

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

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

Geographical distribution. — Basque Provinces, Spain. 

Diagnosis. — Colour paler and more grey than in the other 
races ; size small. 

Measurements. — Type (from Cabrera) : head and body, 55 ; 
tail, 24 ; hind foot, 10; ear, 6*5; upper tooth-row, 7 • 2. (Cranial 
dimensions not known.) 

Remarks. — I have not seen this animal, but from the original 
description, as well as from information received from Mr. Cabrera, 
it appears to be paler than the typical form, to which it bears 
much the same relation as G. russula pulchra to true russula. 



CROCIDURA RUSSULA Hermann. 

(Synonymy under subspecies.) 

Geographical distribution. — Central and southern Europe, 
from the Mediterranean coast to Holland and central Germany. 
Not found in the British Islands. 

Diagnosis — Size rather large (among the European forms) : 
hind foot, 11 to 14 mm. ; condylobasal length of skull, 18 to 
20 • 4 mm. ; upper tooth-row, 8 • 2 to 9 mm. ; tail rather short, 
its actual length 33 to 45 mm., its ratio to head and body 
varying from 45 to 55 ; skull with brain-case not noticeably 
depressed, its height always at least half greatest width and 
usually more ; large upper premolar with antero-external cusp 
small, its height usually less than that of third unicuspid ; 
colour of underparts not strongly contrasted with that of back, 
the line of demarcation along side vaguely defined. 

External characters. — In external characters, aside from the 
relatively longer tail, Crocidura russula agrees with C. leucodon. 
Depth of fur at middle of back about 5 mm. in summer, 8 mm, 
in winter. 

h 2 



100 INSECTIVORA 

Colour. — Upper parts varying from a dark hair-brown, 
tinged with bister to a light drab with or without a shade of 
wood-brown, the darker colour more frequent in winter pelage, 
the light apparently peculiar to summer. The pelage has the 
usual metallic gloss, and the individual hairs show strong silvery 
reflections in certain lights, particularly in the long full winter 
coat. Underparts usually a dull buffy grey or ecru-drab, but 
sometimes almost whitish, rarely tinged with a bright yellowish 
brown,* never strongly constrasted with back, the line of 
demarcation along sides always vague. Feet dull buffy grey or 
light drab. Tail obscurely bicolor, like back above, like belly 
below. 

Skull and teeth. — The skull resembles that of Crocidura 
leucodon, except that the brain-case is noticeably less flattened, 
its depth at middle always exceeding one half greatest width, a 
character readily appreciable to the eye when skulls of the two 
animals are viewed from behind. Teeth essentially as in the 
related species, but large upper premolar with antero-external 






Fis. 20. Fig. 21. 

Posterior view of skull of Crocidura russula. Anterior 

Croeidura leucodon (upper teeth in profile. X 5. 

figure), and C. russula (lower 
figure), x It. 

cusp (paracone) low, its height often much less than that of 
third unicuspid, the distance from its point to ill-defined angle 
in cingulum over anterior root of tooth about half length of 
anterior border of main cusp ; posterior cutting blade not so 
high as in Crocidura leucodon, the angle formed between its 
edge and conspicuously projecting point of main cusp well 
defined and less obtuse than in the related species. 

Measurements. — In the different races the head and body 
ranges from 64 to 95 mm., tail from 33 to 46 mm., hind foot 
from 10 • 8 to 14 mm., condylobasal length of skull from 
18 to 20 "4 mm. The unusual apparent variability in length 
of head and body is probably in great part due to differences in 
method of taking the measurement and to differences in the 
condition of the specimens measured. 

* Such specimens evidently formed the basis of Savi's Sorex thoracicus 
and Dehne's S. chrysothorax. 



CROCIDUEA 101 

Orociduea russula eussula Hermann. 

1777. Sorex araneus Sohreber, Saugthiere, in, p. 573 (not of Linnaeus, 1758). 
1780. Sorex russulus Hermann in Zimmermann, Geogr. Gesch., n, p. 382 

(vicinity of Strassburg, Germany). 
1780. 11 Sorex constrictus Hermann in Zimmermann, Geogr. Gesch., n, 

p. 383 (vicinity of Strassburg, Germany). Based on young in nest. 
1792. 11 Sorex unicolor Kerr, Anim. Kingd., p. 208 (Strassburg, Germany. 

Based on Pennant's account of Sorex constrictus). 
179S. Sorex musaraneus Ouvier, Tabl. Elem. de l'Hist. Nat. des Anim., 

p. 109 (Prance). 

1800. 11 Sorex leucurus Shaw, Gen. Zool., I, pt. 2, p. 538 (Strassburg, 

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

1801. S\prex'] a[raneus] cinereus Bechstein, Gemeinn. Naturgesch. Deutsch- 

lands, I, 2nd ed., p. 867, misprinted 863 (Thiiringen, Germany). 
1801. S[orex'\ a[raneus] candidus Bechstein, Gemeinn. Naturgesch. Deutsch- 

lands, I, 2nd ed., p. 867, misprinted 863 (Thiiringen, Germany). 
1832. Sorex fimbriatus Wagler, Isis, p. 54 (Bavaria, Germany). 
1832. Croc[idura] moschata Wagler, Isis, p. 275 (Substitute for Sorex 

fimbriatus). 
1832. C[rocidura] major Wagler, Isis, p. 1218 (Bavaria, Germany). 
1832. Orocidura rufa Wagler, Isis, p. 1218 (banks of the Rhine, Germany). 
1832. Crocidura poliogaster Wagler, Isis, p. 1218 (banks of the Bhine, 

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

(near Pisa, Italy). 
1839. 1 Sorex inodorus de Selys-Longchamps, Etudes de Micromamm., 

p. 34 (Savi cited as authority, but name apparently published here 

for the first time as synonym of aranea( = russula)). 
1839. ? [Crocidura aranea] var. minor de Selys-Longchamps, Etudes de 

Micromamm., p. 35 (Silesia). 
1839. [Crocidura aranea"] var. albiventris de Selys-Longchamps, Etudes de 

Micromamm., p. 36. (No locality given.) 
1839. ? Crocidura hydruntina Costa, Fauna del Kagno di Napoli, Mamm., 

p. 6 (Otranto, Calabria, Italy). 
1855. Sorex chrysothorax Dehne, Allg. deutsche Naturhist. Zeitung, Neue 

Polge, I, p. 241 (Wilsdurf, near Dresden, Germany). 
1857. Crocidura araneus Blasius, Saugethiere, Deutschlands, p. 144. 
1895. Crocidura russula Thomas, The Zoologist, 3rd ser., xix, p. 63, 

February, 1895. 
1910. Crocidura russula Trouessart, Faune Mamm. d'Europe, p. 43. 

Type locality. — Vicinity of Strassburg, Germany. 

Geographical distribution. — Central Europe, from Holland and 
central Germany to the valley of the Garonne and the coast of 
south-eastern France (Var) ; Italy ; Sardinia ? * ; Guernsey and 
Alderney, Channel Islands. 

Diagnosis. — Size rather large (hind foot, ll - 7 to 14, condy- 
lobasal length of skull, 19 to 20-4), and colour usually dark, 
seldom, if ever, becoming a light drab except in rather worn 
summer pelage. 

Measurements. — Average and extremes of four specimens from 
Oosterbeek, Guelderland, Holland : head and body, 78 (76-81) ; 
tail. 41 (37-45); hind foot, 12-8 (12-2-13-9). Average and 

* I have seen five Sardinian specimens in the Genoa Museum 
resembling the typical form of Crocidura russula and differing widely 
from the Corsican C. cymensis (see pp. 111-112). 



102 



INSECTIVORA 



extremes of nine specimens from Esneux, Liege, Belgium : head 
and body, 77 (72-85); tail, 35-6 (33-38); hind foot, 12-4 
(11" 7-1 3). Average of ten specimens from Pas-de-Calais, France : 
head and body, 74-5 (71-80) ; tail, 40 ■ 9 (38-46) ; hind foot, 13-1 
(12 '5-13 - 5). Average and extremes of six specimens from St. 
Cergues, Vaud, Switzerland : head and body, 87 (83-95) ; tail, 
38-1 (35-41); hind foot, 12-8 (12-4-13-2). For cranial 
measurements see Table, p. 104. 

Specimens examined. — One hundred and two, from the following 
localities : — 

Holland : Oosterbeek, Guelderland, 4. 

Belgium : Esneux, Liege, 9 ; Waremme, Liege, 8 (U.S.N.M.) ; no exact 
locality, 1. 

France : Boulogne-sur-Mer, Pas-de-Calais, 8 ; Guines, Pas-de-Calais, 2 ; 
Abbeville, Somme, 2 (B.M. and Mottaz) ; Guernsey, Channel Islands, 4 ; 
Alderney, Channel Islands, 1 ; St. Briac, Brittany, 1 ; Lignieres, 
Charente, 1 (Mottaz) ; Nancy, Meurthe-et-Moselle, 1 (Merriam) ; Etupes, 
Doubs, 9 (Mottaz) ; Montauban, Haute-Savoie, 4 ; Valescure, Var, 2 ; Ax- 
les-Thermes, Ariege, 1 ; Luchon, Haute-Garonne, 2. 

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

Switzerland : Geneva, 10 (U.S.N.M . and Mottaz) ; St. Cergues, Vaud, 9 
(U.S.N.M. and Mottaz) ; Grosjoan, Vaud, 1 (Mottaz) ; Chesieres, Vaud, 1 
(Mottaz) ; Lucerne, 1 ; Vitznau, Lake of Lucerne, 3 ; Thurgau, Roggwil, 1 ; 
St. GaUen, 3 (U.S.N.M.) ; Ziiberwangen, St. Gallon, 5 (B.M. and U.S.N.M.) ; 
Degersheim, St. GaUen, 5 (B.M. and U.S.N.M.) ; Engelberg, Unterwalden, 1. 

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

3 <5, 1 ?. Oosterbeek, Guelderland, 0. Thomas (c & p). 98. 2. 1. 9-12. 



50 m. Holland. 

Oosterbeek, Guelderland. 
(0. Thomas.) 
AS, 49,1. Esneux, Liege, Belgium. 
1. Belgium. (H. Gronvold.) 
2<5,9,1. Guernsey, Channel Is- 
lands. (B.H.Bunting.) 

Alderney. 

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

Guines, Pas-de-Calais. 

Abbeville, Somme. 

St. Briac, Brittany. 

Montauban, Haute- 
Savoie. 

Montauban, Haute- 
Savoie, 900 m. 

(A. Robert.) 

Valescure, Var. 

Ax-les-Thermes, Ariege. 

Luchon, Haute-Garonne, 
600 m. (A. Robert.) 

Ingelheim, Rheinhessen, 
Germany. 

Strassburg, Alsace. 

(C. Mottaz.) 

Vitznau, Lake of 
Lucerne, 500 m. 
Switzerland. 

Engelberg, 3300 ft. 
Switzerland. 



<5,9. 



9. 
5c5,3 9. 

2(5. 
1. 
1. 
2. 

2(5. 



(5,9. 

9. 

2 9. 



9. 



3 9. 



lal. 



Miller Collection. 

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

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

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

0. Thomas (p). 



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

C. Hilgert (c). 

0. Thomas (p). 

0. Thomas (c&p). 



7. 7. 7. 3851- 

3852. 
95. 1. 1. 3-11. 

7. 1. 1. 30. 

8. 9. 2. 18-21. 

9. 3. 28. 1. 
98. 1. 9. 4-11. 

94. 6. 6. 8-9. 
56. A. 

94. 10. 3. 1. 
97. 1. 9. 2-3. 

6. 4. 2. 2-3. 



8.8.4.164-165. 
8. 3. 27. 1. 
6. 4. 1. 19-20. 

8. 11. 2. 9-10. 

8. 8. 10. 43. 

5. 8. 3. 9-11. 



Dr. J. Anderson (p). 99. 7. 17. 3 . 



CROCIDURA 103 



Ceocidura eussula pulchea Cabrera. 

1907. Crocidura russula pulchra Cabrera, Ann. and Mag. Nat. Hist., 
7th ser., xx, p. 213; September 1, 1907. Type in Cabrera col- 
lection. 

1907. Crocidura russula pulchra Cabrera, Bol. Real Soc. EspaS. Hist. 
Nat., Madrid, vu, p. 223, October, 1907. (For date see Cabrera, 
Ann. and Mag. Nat. Hist., 8th ser., I, p. 189, February, 1908.) 

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

Type locality. — Valencia, Spain. 

Geographical distribution. — Central and southern Spain ; low- 
lands of France south, of the Gironde. 

Diagnosis. — Smaller than Crocidura russula russula (hind foot, 
10 - 8 to 13, condylobasal length of skull, 18 to 19'4), and paler 
in colour, the back a light drab brown tinged with sepia or dull 
russet. 

Measurements. — External measurements of type, male (from 
Cabrera): head and body, 71; tail, 41 - 5; hind foot, 12. 
Average and extremes of seven specimens from Silos, Burgos, 
Spain: head and body, 72 -7 (67-78); tail, 36 (34-37); hind 
foot, ll - 9 (10 • 8-1 2 - 4). Average and extremes of seven 
specimens from Granada, Spain : head and body, 69 • 6 (68-74) ; 
tail, 38-2 (36-40); hind foot, 12-2 (12-12-8). Average and 
extremes of ten specimens from Cadillac-sur-Garonne, Gironde, 
France: head and body, 75 -2 (70-81); tail, 37 (33-40); hind 
foot, 11 '6 (11-12). For cranial measurements see Table, p. 106. 

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

Fbance : Cadillac-sur-Garonne, Gironde, 19 (U.S.N.M.) ; Montrejeau, 
Haute-Garonne, 3 (U.S.N.M.) ; St. Genies, near Nimes, Gard, 2.* 

Spain: Silos, Burgos, 18; Dehesa de Valencia, Valencia, 1; Alcoy, 
Alicante, 8 ; Elche, Alicante, 2 ; Venta del Baul, Granada, 2 ; Granada, 8 ; 
Barracas, Castellon, 1. 

Portugal: Sierra de Gerez, 1 (in alcohol; perhaps referable to cintree). 

9. St. Genies, Gard, 102 m. 0. Thomas (p). 8. 8. 10. 42. 

France. (C. Mottaz.) 

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

Spain. 

?. Dehesa de Valencia. O. Thomas (p). 8. 2. 9. 46. 

(N. Gonzalez.) 

2 <5, 1 9. Alcoy, Alicante. 0. Thomas (p). 8. 2. 9. 43-45. 

(N. Gonzalez.) 

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

4 6, 1 9. Granada. G. S. Miller (c). 8. 8. 4. 33-37. 

dal. Sierra de Gerez.Portugal. Dr. H. Gadow 87.3.28.1. 

(c & p). 



* Intermediate between pulchra and true russula. 



104 



INSECTIVOEA 



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106 



INSECTIVORA 



Observations. 


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)) it 

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Locality. 


C. russula russula (continued). 

Switzerland : Ziiberwangen, St. \ 
Gallen . . ./ 
Degersheim 

>> 
Lugano, Tioino . 

Italy : Ceresole d'Alba . 

Pisa .... 

C. russula pulchra. 

Spain : Silos, Burgos 

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1 08 INSECTIVORA 

Ceociduea eussula cinte^; Miller. 

1907. Crocidura russula cintrse Miller, Ann. and Mag. Nat. Hist., 7th ser., 

xx, p. 390, November, 1907. Type in British Museum. 
1910. Crocidura russula cintrse Trouessart, Faune Mamm. d'Europe, p. 45. 

Type locality. — Cintra, near Lisbon, Portugal. 

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

Diagnosis. — Size as in C. russula pulchra (hind foot, 1 1 ■ 4 to 
12-7, condylobasal length of skull, 18 to 19-2), but colour fully 
as dark as in true russula, the back between the mars-brown and 
russet of Ridgway, the hairs with a peculiar, strong, coppery 
lustre rarely indicated in the typical race. 

Measurements. — External measurements of type : head and 
body, 64 ; tail, 33 ; hind foot, 1 1 • 4. Average and extremes of 
ten specimens from the type locality : head and body, 67 ■ 6 
(64-72); tail, 37-7 (33-42); hind foot, 11-9 (11-4-12-7). For 
cranial measurements see Table, p. 107. 

Specimens examined. — Eleven, all from the type locality. 

Remarks. — In its small size the Cintra shrew agrees with the 
Spanish race, but the colour is conspicuously darker. Taken as a 
whole the series, in winter pelage, is about as dark as in French 
and Belgian russula ; but the noticeable coppery lustre is highly 
characteristic of the Portuguese form. 

6 <$, 5 ?. Cintra, Estremadura, O. Thomas (c & p). 98. 2. 2. 10-20. 
300-350 m. Portugal. (Type of subspecies 98. 2. 2. 11.) 

CROCIDURA SICULA Miller. 

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

1901. Crocidura sicula Miller, Proc. Biol. Soc, Washington, xiv, p. 41, 
April 25, 1901 (Palermo, Sicily). Type in U.S. National Museum. 

1910. Crocidura sicula Trouessart, Faune Mamm. d'Europe, p. 47. 

Type locality. — Palermo, Sicily. 

Geographical distribution. — Sicily. 

Diagnosis. — Size and colour essentially as in Crocidura russula 
pulchra (hind foot, 12 to 13 mm., condylobasal length of skull, 
17 "6 to 19 mm.); brain-case nearly as much flattened as in C. 
leucodon, its depth usually a little less than half greatest width ; 
crown area of molars slightly reduced. 

Colour. — Upper parts a light bluish drab tinged with sepia ; 
underparts faintly constrasted pale smoke-grey, with or without 
a buffy cast. Feet dull whitish grey. Tail obscurely bicolor, 
brownish above, whitish grey below. 

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



CROCIDURA 109 

the skull resembles that of the small races of Crocidura russula. 
Teeth as in C. russula, but upper molars with crown area 
somewhat reduced, a character readily appreciable on comparison, 
and large upper premolar with antero-external cusp slightly 
enlarged, its form and relative size approaching the conditions 
found in C. leucodon. 

Measurements. — External measurements of type (male) and 
a second specimen from the type locality : head and body, 68 
and 75; tail, 32 and 35; hind foot, 12 and 12. Average and 
extremes of six specimens from San Giuglielmo, Castelbuono, 
Sicily : head and body, 76-3 (72-80) ; tail, 35-3 (32-41) ; hind 
foot, 12 - 8 (12-13). For cranial measurements see Table, p. 113. 

Specimens examined. — Fourteen, all from Sioily. Exact localities : 
Palermo, 5 (B.M. and . TJ.S.N.M.) ; Marsala, 2 ; San Giuglielmo, Castel- 
buono, 6 ; Picuzza, 1. 

2 9. Palermo, Sicily. J.I. S. Whitaker (p). 98.10.6.2-2*. 

1. Marsala. (A. Robert.) 0. Thomas (p). 6. 8. 4. 26. 

3 <5, 1 9. San Giuglielmo, Castei- O. Thomas (p). 8. 9. 1. 6-9. 
buono. (A. Robert.) 

CROCIDURA CANEffi Miller. 

1909. Crocidura caneee Miller, Ann. and Mag. Nat. Hist., 8th ser., in, 

p. 418, May, 1909. Type in British Museum. 

1910. Crocidura canese Trouessart, Faune Mamin. d'Europe, p. 48. 

Type locality. — Crete. 

Geographical distribution. — Island of Crete. 

Diagnosis. — Size and general appearance as in the smaller 
forms of CrocidMra russula, and skull with similarly deep cranium ; 
but second upper premolar as large as third, and entire anterior 
portion of upper tooth-row unusually long relatively to cheek- 
teeth. 

Colour. — The colour does not differ appreciably from that of 
dark individuals of C. russula. 

Skull. — The skull is essentially similar to that of the smaller 
forms of Crocidura russula. Brain-case slightly more than half as 
high as wide. Anterior portion of palate between unicuspids 
and anterior incisors more nearly parallel-sided, and more 
elongate than in any of the related species. 

Teeth. — The teeth differ from those of all the other known 
European members of the genus in the approximately equal size 
of the two small upper unicuspids and in the longer, relatively 
narrower crown of the first unicuspid. In the related species the 
first unicuspid is so wide posteriorly that it makes an abrupt and 
noticeable break in the outline of outer side of tooth-row. In 
C. canese this tooth, though larger than usual, is not sufficiently 
wide to project beyond the general line of the outer margins of 
the unicuspid teeth. Second unicuspid fully as large as third, 
which is of normal size. The unicuspid row is thus distinctly 



110 INSECTIVORA 

increased in length, so that the distance from front of large 
premolar to front of incisor equals that from front of large 
premolar to mesostyle of second molar, while in the related 
species it equals that from front of large premolar to metastyle 
of first molar. Large premolars and molars, particularly those 
of mandible, more robust than usual though not peculiar in form. 
Measurements. — External measurements of type (male) : head 
and body, 65 ; tail, 42 ; hind foot, 11 "8. External measurements 
of adult male from Canea : head and body, 71 ; tail, 47 ; 
hind foot, 12 - 6 ; ear, 9 - 5. For cranial measurements see Table, 
p. 113. 

Specimens examined, — Two, both from Crete. 

6 al. Crete. Purchased (Linnsea, 84. 3. 14. 2. 

Frankfort). 

(Type of species.) 
i. Canea, Crete. A. Trevor Battye (p). 8. 10. 24. 1. 

(C. H. B. Grant.) 

CROCIDURA CAUDATA Miller. 

1901. Crocidura caudata Miller, Proc. Biol. Soc. Washington, xiv, p. 42, 

April 25, 1901. Type in U.S. National Museum. 
1910. Crocidura caudata Trouessart, Faune Mamm. d'Europe, p. 49. 

Type locality. — Palesmo, Sicily. 

Geographical distribution. — Sicily. 

Diagnosis. — Size about as in large specimens of G. russula 
(hind foot, 14 mm.). Tail very long, its ratio toihead and body 
about 80, and so thickened that its diameter at middle is 3 mm. 
(in other European species the diameter of tail scarely if at all 
exceeds 2 mm.). 

External characters. — Except for the unusual length of the 
tail Crocidura caudata does not differ in external characters from 
C. russula. The tail is 'so long that when laid forward over back 
it extends to between ears. It is distinctly 4-sided, broader 
below than above, its greatest diameter at middle 3 mm.* 

Colour. — After six months' immersion in alcohol the colour of 
the type specimen was essentially as in Crocidura sicula. After 
eight years more in the same fluid the back appears to have 
assumed a somewhat more brownish cast. 

Skull and teeth. — The only known skull is so injured that the 
details of its form cannot be seen. The rostral portion does not 
differ appreciably from that of C. russula. Teeth essentially as 
in C. russula, but first upper unicuspid larger, third unicuspid 
more crowded against large premolar, and cutting edge of large 
premolar higher, its antero-external cusp, however, of the same 
form as in C. russula. 

* In the type the tail is flattened laterally for about 13 mm. from tip, 
evidently as the result of an accident. 



CROCIDTJRA HI 

Measurements. — External measurements of type : head and 
body, 63 ; tail, 52 ; hind foot, 14. For cranial measurements 
see Table, p. 113. 

Specimen examined. — The type. 

Remarks. — The tail is actually as well as relatively longer in 
this species than in any other European member of the genus. 

CROCIDURA CYRNENSIS Miller. 

1907. Crocidura cyrnensis Miller, Ann. and Mag. Nat. Hist., 7th ser., xx, 

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

Type locality. — Bastia, Corsica. 

Geographical distribution. — Corsica. 

Diagnosis. — Smaller than Crocidura caudata (hind foot, 12 to 
12 '4 mm.) but with tail relatively almost as long, its ratio to 
head and body about 70. 

External characters. — Similar to C. caudata except for the 
smaller size ; tail apparently less thickened than in the Sicilian 
animal, its diameter at middle only about 2 mm. 

Colour. — Back and sides drab washed with a brown inter- 
mediate between wood-brown and raw-umber, this especially 
noticeable on posterior half of back, but scarcely extending to 
sides, which are a nearly clear drab ; underparts a light buffy 
drab-grey, inconspicuously contrasted with sides ; tail dull dark 
drab, essentially unicolor ; feet (both fore and hind) like tail on 
outer half, rather sharply contrasted pale buffy grey on inner 
half. 

Skull and teeth. — While its general size and form are essentially 
as in Crocidura russula, the skull of the Corsican shrew is 
distinguishable by its broader, more deepened rostrum. In the 
type the mandible is peculiar in the unusual depth of ramus, 
though in a second specimen this character is less marked. 
Teeth essentially as in C. russula. 

Measurements. — External measurements of type (adult male) : 
head and body, 67 ; tail, 48 ; hind foot, 12 ■ 4. Very old female 
from the type locality : head and body, 62 ; tail, 46 ; hind foot, 
12 • 4 ; ear, 8 • 2. External measurements of well made skin from 
La Foce de Vizzavona : head and body, 72 ; tail, 51 j hind foot, 
12. For cranial measurements see Table, p. 113. 

Specimens examined. — Three, all from Corsica. 

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



112 INSBCTIVORA 

Ovile Seardu and two from Zinnigas) in the Genoa museum 
appear to be strictly of the russula type, though without com- 
parison of the skulls it is impossible to say whether they are most 
nearly related to true russula or to sicula. Their average and 
extreme measurements (from spirit specimens) are as follows : 
head and body, 65-2 (60-69); tail, 36-8 (33 -6-39); hind 
foot, 12 (11 -8-12-2). 

1 La Foce de Vizzavona, Col. J". W. Yerbury (c & p). 93. 9. 15. 3. 
Corsica. 
S al. Bastia, Corsica. Mrs. Southwell (c & p). 6. 3. 14. 1. 

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

CROCIDURA BALEARICA Miller. 

1901. Grocidura russula Thomas, Proc. Zool. Soc. London, p. 39. 

1907. Grocidura balearica Miller, Ann. and Mag. Nat. Hist., 7th ser., xx, 

p. 391, November, 1907. Type in British Museum. 
1910. Crocidura balearica Trouessart, Faune Mamm. d'Europe, p. 49. 

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

Geographical distribution. — Balearic Islands, Spain. 

Diagnosis. — In general similar to Crocidura cyrnensis, but 
tooth-row distinctly shorter than in the Corsican form. 

Colour. — The three skins are more noticeably brownish above 
than in the one skin of cyrnensis at present known, though the 
actual elements of the colour are the same. Feet with the same 
colour pattern. 

Skull and teeth. — In cranial and dental characters the Balearic 
shrew agrees with Crocidura cyrnensis, except that the mandible 
is less robust (essentially as in the small races of C. russula) and 
the tooth-row is distinctly shorter. Brain-case somewhat more 
flattened than in the small races of C. russula. 

Measurements. — External measurements of type (female) : 
head and body, 62 ; tail, 45 ; hind foot, 12 • 5. External measure- 
ments of two other specimens from the type locality (male and 
female) : head and body, 71 and 72 ; tail, — and 45 ; hind foot, 
12-5 and 12. For cranial measurements see Table, p. 113. 

Specimens examined. — Three, all from the type locality. 

£, 2? San Cristobal, Minorca; O. Thomas and R. I. Pocock 0.7.1.40-42. 
Balearic Islands. (c & p). 

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



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114 INSBCTIVORA 

Family ERLTSTACEID^E. 

1821. Erinaceidse Gray, London Med. Repos., xv, p. 300, April 1, 1821. 

Geographical distribution. — Tropical and temperate Africa, 
Europe and continental Asia ; in Europe west to Ireland, north 
to central Sweden and south-eastern Norway. 

Characters. — Skull deep and heavy, not specially tapering 
anteriorly, most of the sutures persistent ; zygomatic arch com- 
plete and heavy ; floor of brain-case completely ossified ; tympanic 
bone annular, not attached to skull ; auditory process of basi- 
sphenoid large, sometimes forming an evident half-bulla ; glenoid 
surface directed downward (normal) ; a large external pterygoid 
plate ; teeth anterior to molars neither well differentiated by 
form into incisors, canines and premolars nor strictly " unicuspid," 
the anterior upper incisor higher than the others but not specially 
modified in form ; anterior lower incisor short, oblique : crowns 
of upper molars rather high, sub-quadrate in outline (except the 
reduced third), the cusps sub-equal, subterete, near margin of 
crown, the styles and commissures reduced or absent, never 
forming an important functional part of the tooth ; form short 
and heavy ; eyes and ears well developed ; snout pointed, some- 
what produced ; back normally covered with short, stiff spines. 

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

Genus ERINACEUS Linnaeus. 

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

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

1868. Herinaceus Mina-Palumbo, Ann. Agrio. Sicil., 2nd ser., xii, p. 37. 

Type species. — Erinaceus europseus Linnseus. 

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

Characters. — Skull rather short and broad, the zygomatic 
breadth distinctly more than half greatest length ; posterior 
palatal region conspicuously fenestrate ; auditory process of basi- 
sphenoid well developed, concave, sometimes forming a half-bulla ; 
external pterygoid plate rather larger than internal pterygoid 
plate, formed about equally of ectopterygoid and a broad horizontal 
outgrowth from palatine; dental formula: i jjr*, c^l, pm 3z3 
m 3 1 3 = 36 ; canines not differentiated by form from the contiguous 
teeth ; third upper molar consisting of a large protocone and 
minute paracone, all trace of crushing surface absent ; body 
short and heavy, the back covered with stiff, sharply pointed 
bristles of uniform length ; tail shorter than hind foot. 



EEINACEUS 115 

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

KEY TO THE EUROPEAN FORMS OP EEINACEUS. 

Third upper incisor definitely 2-rooted ; elevated 

portion of posterior lower premolar 2-cusped ; a 

bare area among spines at middle of forehead 

(Spain, Balearic Islands and southern Prance)... E. algirus, p. 130. 

Underpart8 clouded with brown, at least in inter- 

ramial and intercrural regions (Spain and 

southern Prance) E. a. algirus, p. 131. 

Underparts entirely whitish (Balearic Islands).... E. a. vagans, p. 133. 
Third upper incisor never definitely 2-rooted ; 
elevated portion of posterior lower premolar 
3-cusped ; no bare area among spines at middle 
of forehead. 
Greatest upper length of maxillary greater than 
or at least equal to depth of rostrum at middle ; 
anterior upper premolar sub-equal to canine, 
its posterior border with small though evident 
cusp (Eastern). 
Hind foot 40 to 43 mm. ; condylobasal length 
of skull in adult male about 58 mm. (Eastern 
Germany through Bohemia and Roumania 

to Greece) E. roumanicus, p. 127. 

Hind foot 35 to 38 mm. ; condylobasal length 

of skull in adult male about 55 mm. (Crete) E. nesiotes, p. 129. 

Greatest upper length of maxillary less than 

depth of rostrum at middle ; anterior upper 

premolar decidedly smaller than canine, its 

posterior border with cusp obsolete or absent 

(Western) E. europseus, p. 115. 

Size larger, the skull in old males exceeding 
59 mm. in condylobasal length. 
Average colour darker, the face never clear, 
pale, buffy grey, but usually with notice- 
able blackish markings (Central-western 

Europe) E. e. europseus, p. 120. 

Average colour lighter, the face usually clear, 
pale, buffy grey without noticeable black- 
ish markings (Iberian Peninsula) E. e. hispanicus, p. 122. 

Size smaller, the skull in old males not ex- 
ceeding 59 mm. in condylobasal length. 
Colour of head and shoulders lighter than 

that of contiguous spiny area (Italy) E. e. italicus, p. 123. 

Colour of head and shoulders darker than 

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

ERINACEUS EUROPSEUS Linmeus. 
(Synonymy under subspecies.) 

Geographical distribution. — Europe from the Mediterranean 
coast to Scotland and southern Scandinavia ; west to Ireland • 
eastern limits of range not known. 

Diagnosis. — Size large (head and body about 225 to 275 ; hind 
foot 40 mm. or more ; condylobasal length of skull more than 



116 INSECTIVORA 

55 mm.) ; spines coarse and heavy, extending in an unbroken 
line across forehead and decidedly overtopping ears ; skull with 
heavy, deep rostrum, the distance from posterior extremity of 
premaxillary to posterior extremity of maxillary less than rostral 
depth at middle ; auditory process of basisphenoid short, wide- 
funnel shaped, not forming a half-bulla ; third upper incisor with 
one root, this sometimes partly divided longitudinally ; elevated 
portion of posterior lower premolar with three cusps (fig. 25 a). 

External characters.— General form short, thick and clumsy, 
the legs short, the feet large, plantigrade. Legs, tail, underparts 
and head, except crown, clothed with coarse fur, the finer more 
woolly hairs of which are about 15 mm. long, the longer, straight 
hairs about 40 mm. in length. The rest of the body is covered 
by a densely-set mass of sharply pointed bristles about 25 mm. in 
length and 1 mm. in diameter. * The skin on which these bristles 
are set is loosely attached to body, and provided with a special 
system of muscles by which the edges of the spiny area can be 
drawn together ventrally over the animal's retracted head and legs, 
forming a complete protection for the entire body. Feet robust 
with short digits and well developed claws, those on hind feet 
longest. Fore foot broad and rounded with very short, thick 
fingers, the third and fourth sub-equal and longest, second slightly 
shorter, fifth extending to base of fourth, first well developed 
but not reaching base of second ; balls of all five large and pad- 
like ; palm entirely naked ; three large, semi-confluent pads 
at bases of median digits, a small tubercle (about 2 mm. in 
diameter) at base of thumb, and two large pads at back of wrist, 
the outer the larger ; skin between pads wrinkled ; hind foot 
much like fore foot but longer ; second, third and fourth digits 
sub-equal and longest, but their claws noticeably graduated from 
second to fourth, fifth digit extending to base of fourth, first not 
reaching base of second ; sole naked, the tubercles as on palm 
but more crowded, the two posterior sub-equal. Tail short and 
thick, its length much less than that of hind foot. Bar simple, 
rounded, shorter than bristles on crown, the meatus without 
valves. Muzzle moderately produced, somewhat pointed, the 
muzzle-pad well developed, naked, its surface marked with minute 
furrows, its lateral edges finely scalloped, its lower border continued 
as a pair of parallel ridges extending inward to palate. Eye 
well developed but rather small. Mammae : p 1-1, a 2-2, 
i 2-2 = 10. 

Colour. — Furred portions of body varying from dull brown to 
dirty whitish, the under fur usually a dusky hair-brown, the 
longer hairs lighter and more buffy. Belly often irregularly 
blotched and variegated with whitish and darker or lighter 
brown. Feet usually darker than sides. Cheeks and eye-ring 
often darker than rest of head. Spines buffy at base, then with 

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



BRIKACBUS 117 

a slaty area of variable width, followed by a narrow but sharply 
denned buffy annulation and an obscurely darker tip. The 
general effect is a coarse grizzle, the exact tone of which varies 
considerably in the different geographical races as well as in 
individuals of the same race. 

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




rig. 22. 

Erinaceus europseus. Nat. size. 



conspicuously wider than interorbital region, the rostrum short 
and deep (distance from anteorbital foramen to front of pre- 
maxillary less than depth through anterior root of zygoma). 
Occiput and interorbital region marked by noticeable ridges. 



118 INSEOTIVOEA 

Ventral profile straight, the dorsal profile essentially parallel to 
it from lambda nearly to front of interorbital region, then sloping 
forward at a slight angle (about 15°) ; occiput squarely or some- 
what obliquely truncate. General outline of occipital region as 
viewed from behind truncate-triangular, slightly more than half as 
high as wide, the base of the triangle formed by line joining tips 
of widely projecting mastoid processes, the apex by the narrowly 
rounded or bluntly pointed lambdal region. Paroccipital 
processes nearly as large as mastoid processes and resembling 
them in form, though more slender and directed more backward. 
Basisphenoid with deep median pit between bases of half -funnel 
formed auditory processes, the pit continuous anteriorly with 
mesopterygoid fossa. Tympanic ring open postero-externally, its 
greatest breadth (antero-internal) about 3 mm. Inner and outer 
pterygoid plates broadly triangular, approximately alike in size 
and form, each containing more of the pterygoid than palatine 
element. Hamular short, strongly curved. Mesopterygoid space 
slightly longer than broad. Palate terminating posteriorly in a 
high transverse ridge and strongly projecting median spine, the 
ridge nearly straight, its median portion well developed. In 
front of ridge the palatine bones are conspicuously and irregularly 
fenestrate. Lambdoid crest high. Sagittal crest low but 
evident, extending forward to back of interorbital region. Here 
it divides into two low, diverging ridges which pass forward 
toward lachrymal region. In some specimens they can be traced 
as far as the high, well defined ridge which occupies edge of orbit 
for a distance of about 7 mm. above lachrymal foramen. Ante- 
orbital foramen small, separated from lachrymal foramen by a 
space much greater than its own diameter, its anterior border 
over anterior root of large premolar. Upper portion of maxillary 
rather short, its length behind posterior point of premaxillary 
less than depth of rostrum at middle. Posterior termination of 
premaxillary variable in form : nearly square, broadly or narrowly 
cuneate, rounded, or rounded with supplemental inner spicule. 
Mandible short and heavy, the greatest depth of ramus about one- 
third length of alveolar line. Coronoid process high, narrow, 
sharply hooked backward at tip. Angular process about as wide 
as coronoid process, but not so long, its apex slightly bent 
inward. 

Teeth. — General aspect of teeth as compared with that in 
other European members of the order, short, heavy and blunt, 
distinctly omnivorous rather than strictly insectivorous in type. 
Anterior upper incisor about twice as high as the succeeding 
small teeth, its shaft subterete, flattened posteriorly, directed 
slightly forward and inward, the teeth separated at base by 
space about equal to height of shaft, at tip by about half this 
distance. The four succeeding teeth (two incisors, canine and 
anterior premolar) are essentially alike in form, the crown 
slightly longer than wide, its height slightly greater than length, 



ERINACEUS 



119 



the blunt point of conical cusp somewhat in front of middle of 
crown. Of these four unicuspid teeth the first is smallest, the 
second and third sub-equal and larger, the fourth intermediate. 
On posterior side of crown of each unicuspid there is a faintly 
developed ridge extending to apex of cusp, this ridge tending to 
rise posteriorly, especially in fourth unicuspid, to form a very 
rudimentary secondary cusplet. First and second incisors invari- 
ably single-rooted. Third incisor single-rooted, but root occasionally 
showing trace of longitudinal furrow on outer side. Canine 
usually single-rooted, the root with or without longitudinal 
furrow ; but in a small series of specimens every stage may be 
observed from this condition to a completely two-rooted tooth, 
each root with a distinct alveolus. First premolar single-rooted, 
the root often showing traces of longitudinal division, and perhaps 
rarely double. Anterior lower incisor essen- 
tially like the corresponding upper tooth 
but not so high. Its shaft is directed 
obliquely forward in line with symphysis, 
the teeth of opposite sides parallel, separated 
throughout by a narrow space. 
Three succeeding teeth unicuspid, 
the crowns similar in outline to 
those of upper unicuspids, but an- 
terior cusps obsolete and posterior 
cusplets relatively better developed. 
Second upper premolar scarcely 
broader than the unicuspids, but 
three-rooted and with a distinct 
protocone, metacone and postero- 
external commissure. Large upper 
premolar with well developed pro- 
tocone, hypocone, metacone and pos- 
tero-external commissure, the cusps 
much as in the molars except that 
hypocone is relatively smaller and 
metacone and its commissure larger 
and more trenchant. Large lower premolar with a high anterior 
three-cusped portion similar to first triangle of lower molars, 
except that the metaconid is reduced to a slight thickening at 
inner base of commissure of protoconid ; second triangle repre- 
sented by a mere narrow ledge or thickened cingulum. First 
upper molar sub-quadrate in outline, the crown slightly wider 
posteriorly than anteriorly. Protocone with somewhat broader 
base than the other cusps and with low ill-defined anterior 
and posterior commissures. Paracone, metacone and hypocone 
sub-equal, the metacone slightly larger than the others. All 
three are subterete with faintly indicated commissures, that 
extending outward and backward from metacone to rudimen- 
tary metastyle the most distinct. Parastyle and mesostyle 




no. 23. 

Erinaceus europseus. Teeth. 



1 20 INSBCTIVOEA 

absent.* A small but evident metaconule. Second molar like 
first but smaller, its crown area about equal to tbat of large 
premolar, its greatest diameter anterior instead of posterior. 
Cusps essentially as in first molar, except that paracone is larger 
than metacone and hypocone, and metaconule is barely indicated. 
Third molar reduced to a protocone nearly as large as in the 
other teeth, and a rudimentary paracone, the two connected by 
a cutting edge sloping obliquely outward, forward and upward. 
The tooth is single-rooted. Lower molars with the usual cusps 
and commissures, the cusps more terete and commissures less 
trenchant than in other European insectivores. Third molar 
consisting of the anterior triangle only, this somewhat smaller 
than in the other two teeth. 

Erinaceus bueopjeus europ^eus Linnseus. 

1758. [Erinaceus] europseus Linnaeus, Systema Naturse, I, 10th ed., p. 52 

(Sweden). 
1779. [Hystrix] erinaceus Blumenbach, Handbuoh d. Naturgesch., p. 72 

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

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

Nat., p. 68 (France). 

1857. Erinaceus europseus Blasius, Saugethiere Deutschlands, p. 153 (part). 

1858. E[rinaceus] caniceps Hamilton Smith, Jard. Nat. Libr., 2nd ed., xv 

(Mammalia I), p. 148 (near Brussels, Belgium). 
1897. [Erinaceus] echinus Schulze, Abh. a. Vortr. Gesammtb. Naturw. iv, 

No. 10, p. 19 (Substitute for europssus). 
1900. Erinaceus europseus Barrett-Hamilton, Ann. and Mag. Nat. Hist., 

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

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

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

Mamm. d'Europe, p. 38. 

Type locality. — Upsala, Sweden. 

Geographical distribution. — Western central Europe from 
Scotland, southern Norway, and central Sweden to the Pyrenees 
and Alps ; west to Ireland ; eastern limits of range not known. 

Diagnosis. — Size largest of the European hedgehogs (condylo- 
basal length of fully adult skulls usually 61 to 63 mm., seldom 
less than 60 mm.) ; colour dark, the sides and underparts seldom 
if ever a light buffy grey ; face with blackish area extending 
from eye to muzzle. 

Measurements. — Adult male and female from Upsala, Sweden : 
head and body, 265 and 263 ; tail, 34 and 37 ; hind foot, 44 
and 43. Adult male from Innerwick, Haddingtonshire, Scotland : 
head and body, 218; tail, 17 ; hind foot, 42. Adult male and 

"■ The cingulum usually forms a small projection resembling a rudi- 
mentary parastyle. 



ERINACBDS 121 

female from Oundle, Northampton, England : head and body, 
249 and 257 ; tail, 24 and 31 ; hind foot, 40 and 40. Adult 
male and female from Maredsous, Namur, Belgium : head and 
body, 270 and 251 ; tail, 39 and 33 ; hind foot, 44 and 44. 
Adult male from Bouconne, Gers,. France : head and body, 220 ; 
tail, 20 ; hind foot, 41. Adult male and female from St. Gallen, 
Switzerland : head and body, 279 and 297 ; tail, 41 and 43 ; hind 
foot, 44 and 47. For cranial measurements see Table, p. 124. 

Specimens examined. — Seventy-three, from the following localities : — 

Scotland : Dunphail, Elgin, 2 ; Gordonstown, Elgin, 2 ; Innerwiok, 
Haddingtonshire, 1 ; Glendoo, Inverness, 1 (Wilson) ; Lanarkshire, 1. 

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

Wales : Cardiff, 2. 

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

Norway : Asker, near Christiania, 1. 

Sweden : Upsala, 5 (U.S.N.M.) ; Upland, 1. 

Denmark: Copenhagen, 5 (Andersen). 

Holland : No exact locality, 1. 

Belgium : Maredsous, Namur, 2. 

Prance : Eoret de Bouconne, Gers, 1 ; Cranves-Sales, Haute-Savoie, 1. 

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

Switzerland : Geneva, 2 (Mottaz) ; St. Gallen, 7 (B.M. and U.S.N.M.) ; 
Uzwil, St. Gallen, 1 (U.S.N.M.) ; Heresau, St. Gallen, 1 (U.S.N.M.) ; Wolf- 
halden, Appenzell, 1 (U.S.N.M.) ; Thurgau, 2. 

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

<5, 9. Gordonstown, Elginshire, W. R. Ogilvie-Grant 11. 1. 3. 379- 

Scotland. (c & p). 380. 

<5. Innerwick, Haddington- W. Eagle Clarke (c & 0. 3. 13. 1. 

shire. p). 

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

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

i, 9. Oundle, Northampton, Hon. N. C. Roths- 11. 1. 3. 381- 

England. child (c & p). 382. 

S. Graftonbury, Hereford- W. E. de Winton (c 11. 1. 3. 383. 

shire. & p). 



* For discussion of this character see Lonnberg, Ann. and Mag. Nat. 
Hist., 7th ser., v, pp. 542-544, June, 1900, and Barrett-Hamilton, I.e., 
pp. 245-246, August, 1900. The question cannot yet be regarded as 
decided, since no adequate series of skulls has yet been brought together. 



122 



INSBCTIVORA 



2 6. 

6. 
6. 
6. 

3 juv.al. 

S, 9. 

6, 9 al. 

9. 



6 al. <5. 
2 6. 



Saffron Walden, Essex. G. Barrett-Hamilton 11, 1. 2. 91-92. 

(Wright.) 
Banstead, Surrey. 
Godalming, Surrey. 
Ockley, Surrey. 



Shrewsbury, Shropshire. 

Somerset. 

Somerset. 

Cardiff, Glamorganshire, 
Wales. 

Nenagh, Tipperary, Ire- 
land. (W. Smithwick.) 

Ennis, Glare. 

Kilmanock, Wexford. 



C.H.B.Grant(c&p). 
W.T.Blanford(o&p). 
Hon. Ella Scarlett 

(c & p). 
H. E. Forrest (c & p). 
Dr. J. Anderson (s). 
Dr. J. Anderson (p). 
E. Drane (c & p). 



11. 1. 3. 386. 
11. 1. 3. 384. 
11. 1. 3. 385. 



Asker, Christiania, Nor- 
way. 

Upland, Sweden. (G. 
Eolthoff.) 

Holland. {Seba Coll.) 

Maredsous, Namur, Bel- 
gium. 

Foret de Bouoonne, Gers, 
250 m. Prance. (A. 
Robert.) 

Cranves - Sales, Haute- 
Savoie. (A. Robert.) 

Burgheim, Bavaria, Ger- 
many. (Wolterstorff.) 

Ingelheim, Rheinhessen. 
2 <5, 9 juv. St. Gallen, 500 m. 
Switzerland. (E. H. 
Zollikofer.) 
6, 9. Thurgau, 400 m. (E. H. 
Zollikofer.) 



6 juv. 

lal. 

2 6. 



6, 9. 



6. 



0. 9. 23. 1-3. 
93. 7. 81. 1-2. 
93. 7. 31. 3-4. 
11. 1. 3. 387. 



G. Barrett-Hamilton 11. 1. 2. 95. 

(*)■ 

J. W. Scott (o & p). 93. 10. 30. 1-2. 

G. Barrett-Hamilton 11. 1. 2. 93-94. 

(o & p). 

Christiania Museum 93. 3. 1. 7. 

(e). 

LordLilford (p). 11. 1. 1. 151. 

Lidth de Jeude Coll. 67. 4. 12. 555. 

Bev. G. Pournier (c 1. 6. 2. 1-2. 

&p). 

O. Thomas (p). 6. 4. 1. 10. 



O. Thomas (p). 

Lord Lilford (p). 

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

O. Thomas (p). 



6. 4. 2. 1. 

11. 1. 1. 92, 150. 

8. 11. 2. 5. 
4. 4. 5. 27-29. 

4. 4. 5. 30-31. 



Ekinaceus euroPjEUS hispanicus Barrett-Hamilton. 

1900. Erinaceus europseus hispanicus Barrett-Hamilton, Ann. and Mag. 

Nat. Hist., 7th ser., v, p. 363, April, 1900. Type in British Museum. 
1910. Erinaceus europseus hispanicus Trouessart, Faune Mamm. d'Europe, 

p. 39. 

Type locality. — Seville, Spain. 

Geographical distribution. — Iberian Peninsula. 

Diagnosis. — Size large, essentially as in E. europseus europseus 
(condylobasal length, of skull in old individuals about 60 mm.) ; 
general colour paler than in the typical race, the fur often a 
uniform light buffy grey, seldom with any evident darker area 
between eye and muzzle. 

Colour. — There is much variation in colour, some individuals 
essentially agreeing with the palest examples of true europseus. 
In its extreme phase, however, the colour is a whitish buff, 
decidedly paler than the cream-buff of Ridgway, the muzzle and 
region about eyes washed with ecru-drab ; feet tinged with drab ; 
spines the same whitish buff, about half of them with scarcely any 



ERINACEUS 123 

dark shading, the rest with a drab sub-terminal band, the general 
effect of spiny area scarcely speckled, and nearly as pale as fur. 
In the type specimen the fur is a dull cream-buff, and the feet 
are washed with broccoli-brown ; spines drab brown with light 
tips, essentially as in true europseus. 

Measurements. — Type : hind foot (dry), 40. Adult male and 
female from Burgos, Spain : head and body, 270 and 250 ; tail, 
30 and 28 ; hind foot, 43 and 41. Adult male and female from 
Pajares, Leon : head and body, 252 and 249; tail, 22 and 21 ; 
hind foot, 44 and 42. For cranial measurements see Table, 
p. 125. 

Specimens examined. — Twenty, from the following localities in Spain : 
Arreohavaleta, Vitoria, 1 ; Pajares, Leon, 7 ; Burgos, 5 ; Palacios de la 
Sierra, Burgos, 1; Bejar, Salamanca, 2; Seville, 4. 

Remarks. — The Spanish hedgehog is a moderately well 
differentiated form. Extreme specimens are easily distinguish- 
able from typical europaeus ; but in general the difference between 
the two races must be regarded as an average one. 

6. Arreohavaleta, Vitoria, 0. Thomas (p). 8. 2. 9. 9. 

Spain. (N. Gonzalez.) 
<5, 2 9, ¥ juv. Pajares, Leon. (N. Gon- O. Thomas (p). 8. 2. 9. 2-6. 



S, 9, ? juv. Burgos. G. S. Miller (c). 8. 8. 4. 17-19. 

2 <5. Burgos. (N. Gonzalez.) O. Thomas (p). 8. 2. 9. 10-11. 

2 c5 juv. Bejar, Salamanca. (N. O. Thomas (p). 8. 2. 9. 7-8. 

Gonzalez.) 
6, 2 ?, 1. SeviUe. (Dr. A. Ruiz.) Lord Lilford (p). 95. 3. 3. 1-4. 

(Type of suhspecies 95. 3. 3. 2.) 



Erinaceus eueop^eus italicus Barrett-Hamilton. 

1857. Erinaceus europseus Blasius, Saugethiere Deutschlands, p. 153 (part). 
1900. Erinaceus europseus italicus Barrett-Hamilton, Ann. and Mag. Nat. 

Hist., 7th ser., v, p. 364, April, 1900. Type in British Museum. 
1910. Erinaceus europseus italicus Trouessart, Faune Mamm. d'Europe, 

p. 39. 

Type locality. — Siena, Italy. 

Geographical distribution. — Italian Switzerland, Italy and 
Sardinia ; Corsica 1 

Diagnosis. — Colour as in E. europseus europseus or slightly 
paler ; size less than in the typical race, the largest skulls 
probably not exceeding 59 mm. 

Colour. — The colour is about as in the paler individuals of 
typical europseus, though the speckling of the spiny area seems in 
general to be finer, and the underparts usually lack all heavy 
dark clouding. 

Measurements. — Adult female from Curoggio, Ticino, Switzer- 
land : head and body, 250; tail, 29 ; hind foot, 41. Two adult 



124 



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Locality. 


E. europseus europaeus. 

Norway : near Christiania 
Sweden: Upsala 

,, ... 

Denmark : Copenhagen 

Scotland : Haddingtonshire 

Lanarkshire 
Wales: Cardifi . 

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Belgium : Maredsous, Namur . 

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Germany: Brunswick 

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126 INSBCTIVOEA 

males from Siena, Italy : head and body, 210 and 220 ; * tail, 30 
and 28 ; * hind foot, 40 and 42.* Two adult females from the 
same locality : head and body, 200 and 208 ; tail, — and 32 ; 
hind foot, 43 and 38. For cranial measurements see Table, 
p. 125. 

Specimens examined.— Seventeen, from the following localities : — 
Switzebland: Bigorio, Ticino, 1 (U.S.N.M.); Curoggio, Tieino, 1 
(U.S.N.M.) ; Gentilino, Ticino, 1 (U.S.N.M.). 

Italy : Empoli, Florence, 1 ; Siena, 4 ; Ostia, Borne, 1 ; Borne, 4. 
Sardinia : Su Cramu, 1 ; Bare, 1 ; Marusei, 1 ; Trecorgia, 1. 

Remarks. — Though not so pale as the Spanish race the Italian 
hedgehog seems worthy of recognition as a form distinct from 
true europseus. Its status is at present unsatisfactory, owing to 
the lack of sufficient material ; but specimens from south of the 
Alps seem never to attain the large size of Central European 
adults. The Sardinian specimens that I have seen are in general 
paler than those from the mainland ; but here again the material 
is insufficient. A hedgehog is known to occur in Corsica, but 
no specimens have yet been compared with the Italian race. 

¥. Empoli, Florence, Italy. A. H. Savage Landor 97.3.7.1. 

(c & p). 
2 6, 2 ?. Siena. (S. Brogi.) Dr. E. Hamilton (p). 98. 10. 2. 5-8. 

(Type of subspecies 98. 10. 2. 5.) 
1. Ostia, Borne. Dr.L. Sambon(c&p). 1. 1. 2. 7. 

3 i. Borne. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. 3-4. 96. 

(*)■ 
£. Su Gramu, Sardinia. (W. O. Thomas (p). 0. 12. 3. 5. 

Wolterstorff.) 
t. Bare. (W. Wolterstorff.) O. Thomas (p). 0. 12. 3. 3. 



9. Marusei. (W. Wolterstorff.) O. Thomas (p). 0. 12. 3. 6. 

? juv. Trecorgia. (W. Wolterstorff.) O. Thomas (p). 0.12.3.4. 



Erinaceus europseus consolei Barrett-Hamilton. 

1900. Erinaceus europseus consolei Barrett-Hamilton, Ann. and Mag. Nat. 

Hist., 7th ser., v, p. 366, April, 1900. Type in British Museum. 
1910. Erinaceus europseus consolei Trouessart, Faune Mamm. d'Europe, 

p. 40. 

Type locality. — Palermo, Sicily. 

Geographical distribution. — Sicily. 

Diagnosis. — Size apparently as in JS. europseus italicus (only 
known specimen imperfect) ; colour differing from that of 
italicus in the uniform dusky brown head and shoulders ; quills 
unusually robust, their dark and light markings strikingly con- 
trasted. 

Colour. — Whole head and sides of neck and shoulders a 
uniform dark brown between hair-brown and sepia, sprinkled 
with buffy grey hairs. The dark brown continues back along 

* Type. 



ERINACEUS 127 

edge of spiny area to tail, but throughout this region it is overlaid 
by the uniform light cream-buff of underparts. Spines very dark 
drab with light cream-buff tips, the light area shorter on spines 
of middle of back than on those of sides, thus producing a slight 
though evident darker median dorsal area. Feet so injured that 
colour cannot be determined. 

Skull and teeth. — The imperfect skull shows no peculiarities. 
Teeth as in specimens from the mainland. 

Measurements. — Type (sex not known) : head and body, 252 ; 
tail, 50 ; hind foot, 40. For cranial measurements see Table, 
p. 125. 

Specimen examined. — The type. 

Remarks. — If not an abnormal specimen of E. europseus italicus 
the type of consolei represents a very distinct local race. 

1. Palermo, Sicily. J. I. S. Whitaker (p). 98. 10. 6. 1. 

(Type of subspecies.) 

ERINACEUS ROUMANICUS Barrett-Hamilton. 

1900. Erinaceus europseus roumanicus Barrett-Hamilton, Ann. and Mag. 

Nat. Hist., 7th ser., v, p. 365, April, 1900. (Gageni, Boumania.) 
Type in British Museum. 

1901. Erinaceus danubicus Matschie, Sitz.-Ber. Gesellsch. Naturforsch. 

Freunde, Berlin, p. 229, December, 1901. (Prundu, Boumania.) 
1910. Erinaceus europssus roumanicus and E. europseus danubicus Troues- 
sart, Faune Mamm. d'Europe, pp. 40-41. 

Type locality. — Gageni, Prahova, Roumania. 

Geographical distribution. — -From eastern Germany (Konigs- 
berg) and northern Bohemia south through Hungary, into 
Greece. 

Diagnosis. — In general like Erinaceus europseus, but colour of 
furred parts usually much darker, and chest often with a large, 
conspicuously contrasted whitish area ; skull with upper length of 
maxillary greater than depth of rostrum at middle ; teeth 
essentially as in E. europseus, but first upper premolar with 
postero-basal cusp usually more distinct. 

Colour. — Furred area dark hair-brown interspersed with buffy 
and whitish hairs, the former most numerous on sides, the latter 
forming a clear whitish pectoral area which sometimes spreads 
over entire underparts. Feet and tail sepia. Face usually with 
a seal-brown suffusion. Claws blackish horn-colour. Quills 
with colours usually less contrasted than in E. europseus, the 
general effect of the spiny area darker and less speckled. 

Skull and teeth. — The skull differs from that of E. europseus 
in the relatively longer, less deepened rostrum, a peculiarity 
which seems chiefly to involve the maxillary bone. Upper length 
of maxillary greater than depth of rostrum at middle. Posterior 



128 INSBCTIVOEA 

transverse palatal ridge usually lower and less developed, 
particularly at middle, where there is often a slight angle. 
Ridge at margin of orbit in lachrymal region not so long as in 
157. europseus, and less distinctly marked off from general contour 




Fig. 24. 
Erinaceus roumanicus. Nat. size. 

of skull, its degree of development somewhat as in E. algirus. 
Teeth not certainly distinguishable from those of Erinaeeus 
europseus, though anterior upper premolar is usually larger 
relatively to canine, and its posterior cusplet tends to be better 
developed. 

Measurements. — External measurements of adult male and 
female from vicinity of Konigsberg, Germany : head and body, 
285 and 280 ; tail, 21 and 22 ; hind foot, 42 and 41. Type 
(adult female) : head and body, 206 ; tail, 24 ; hind foot, 40 ■ 6. 
Adult female from Corfu, Greece : head and body, 263 ; tail, 
37 ; hind foot, 43 ; ear, 29. Adult male from Cephalonia, 
Greece: head and body, 260; tail, 35; hind foot, 42; ear, 31. 
For cranial measurements see Table, p. 132. 

Specimens examined. — Eleven, from the following localities : — 
Germany: Near Konigsberg, 3 (U.S.N.M.). 



ERINACEUS 129 

Austria-Hungary: Haida, Arva, Bohemia, 1; Vasoar, Eisenburg, 
Hungary, 1 (U.S.N.M.). 

Roumania : Gageni, Prahova, 2. 

Greece: Corfu, 2; Cephalonia, 1; Tatoi, near Athens, 1. 

Bemarles. — Though at first sight very similar to Erinaceus 
europseus, this species is easily recognizable by its cranial characters. 
In most specimens there is a strong contrast between the dark 
posterior portion of underparts and whitish chest, throat and 
shoulders, a pattern which appears to be rarely if ever well 
developed in the related species. 

9. Haida, Bohemia. Lord Lilford (p). 97. 8. 14. 1. 

<5, 9. Gageni, Prahova, Roumania. Lord Lilford (p). 4. 4. 6. 15-16. 

(TV. Dodson.) (Type of species 4. 4. 6. 16.) 

6. Potamos, Corfu, Greece. J. I. S. Whitaker (p). 8. 10. 1. 6. 

(C. Mottaz.) 

9. Argostoli, Cephalonia. J. I. S. Whitaker (p). 8. 10. 1. 7. 

(C. Mottaz.) 

A. Tatoi, Athens. C. Mottaz (c). 8. 11. 3. 8. 

ERINACEUS NESIOTES Bate. 

1906. Erinaceus europseus nesiotes Bate, Proc. Zool. Soc. London, 1905, n, 
p. 316, April 5, 1906. Type in British Museum. 

1910. Erinaceus europmus nesiotes Trouessart, Faune Mamm. d'Europe, 
p. 40. 

Type locality. — Near Gonia, western Crete. 

Geographical distribution. — Island of Crete. 

Diagnosis. — Similar to Erinaceus roumanicus but smaller (hind 
foot, 35 to 38 ; condylobasal length of skull in adult male, about 
55 mm.) ; spines not so coarse as in the related species ; first 
upper premolar distinctly larger than canine and of essentially 
the same height. 

Colour. — The colour is like that of E. roumanicus. Under- 
parts, sides and face dull whitish grey with faint dark clouding 
between eye and muzzle, and in one of the three skins with a 
dark wash on posterior half of underparts. 

Skull. — As in E. roumanicus, but not attaining as large size. 

Measurements. — Adult male'"' and female from the type 
locality: head and body, 208 and 204; tail, 29 and 19; hind 
foot, 40 and 38 (dry, 38 and 35). For cranial measurements see 
Table, p. 132. 

Specimens examined. — Three, all from Crete. 

Bemarles. — The Cretan hedgehog is nearly related to Erinaceus 
nnimanicus, though well differentiated by its smaller size and by 
the relatively large fourth unicuspid tooth. 

■i, ?. Gonia, Crete. Miss D. Bate (c). 5. 12. 2. 11-12. 

(5. 12. 2. 11 Type of species.) 
9. Mesoghia. Miss D. Bate (c). 5. 12. 2. 13. 

* Type. 



130 



INSECTIVORA 



ERINACEUS alcirus Duvernoy and Lereboullet. 

(Synonymy under subspecies.) 

Geographical distribution. — Northern Africa ; also in southern 
Spain, southern France, and the Balearic Islands. 

Diagnosis. — Smaller than Erinaceus europseus (hind foot less 
than 40 mm.) ; spines not so coarse ; middle of forehead with 
bare area among the spines ; skull with sagittal crest extending 
forward to middle of frontal ; a wide flattened area on each side 
of bony palate behind transverse ridge ; third upper incisor with 
two perfectly distinct roots ; elevated portion of posterior lower 
premolar with only two cusps (fig. 25 b). 

External characters. — Externally Erinaceus alyirus is dis- 
tinguishable from E. europseus by its smaller size, shorter, more 
slender and apparently more densely-set bristles, and by the bare 
area at middle of forehead. This bare area is about 7 mm. wide 
and extends back about 10 to 15 mm. from front line of spines. 
In dried skins it is sometimes partly hidden by shrinking. Fur 
more dense and less, coarse than in the larger animal. Claws on 
front feet seldom attaining a length of 8 mm. 

Colour. — The colour resembles in general that of the paler 
races of E. europseus. 

Skull. — The form of the skull is essentially as in E. europseus 
except that rostrum is less elevated posteriorly, so that the dorsal 
profile tends to become slightly concave. Sagittal crest when" 
fully developed extending forward to middle of frontal, while in 
E. europseus it is usually confined to parietals, rarely encroaching 
on posterior edge of frontal. Ridge at margin of orbit very short, 
scarcely more than a process above lachrymal foramen. Bony 
palate extending behind transverse ridge as a well-defined flat area 
divided along median suture by a longitudinal ridge representing 
the median spine of E. europseus. Basisphenoid pit narrower, 
relatively deeper, and with more overhanging edges than in 
E. europseus. 

Teeth. — In general the teeth show no departure from those 
of E. europseus. The posterior lower pre- 
molar, however, lacks all trace of the meta- 
conid, so that the resemblance of the elevated 
portion of the tooth to the first triangle lof 
)»! and m 2 is completely destroyed. In the 
upper jaw the third incisor, canine, and 
a b first premolar are two-rooted, apparently 

I'ig. 25. without exception. 

Large lower premolar of Measurements. — Head and body about 

fndT^i>«Tb) M xl 200 to 250 ; tail, 25 to 40 ; hind foot, 32 to 
37 ; condylobasal length of skull, 54 to 59 mm. 
Remarks. — This species is readily distinguishable from Erinaceus 
europseus by the bare spot among spines of forehead, the perfectly 
two-cusped large lower premolar, and the two-rooted third upper 




ERINACEUS 



131 



incisor. .Although occurring wild in southern Spain, on the 
Balearic Islands, and in south-eastern France, it seems not 
improbable that the animal owes its presence in Europe to the 
agency of man. 

Erinaceus algirus algirus Duvernoy and Lereboullet. 

1840. Erinaceus algirus Duvernoy and Lereboullet, Mem. Soc. Mus. 

d'Hist. Nat. Strasbourg, in, fasc. 2, p. 4. 
1898. Erinaceus algirus de Winton, Proo. Zool. Soc. London, 1897, p. 955. 

Type locality. — Oran, Algeria. 

Geographical distribution. — Northern Africa ; also in southern 
Spain and south-eastern Prance. 

Diagnosis. — Condylobasal length of skull in individuals with 
distinctly worn teeth 57 to 59 mm. ; underparts dusky through- 
out or with at least an evident dark wash in interramial and 
intercrural regions. 

Colour. — Furred area bufly white to base of hairs, except on 
muzzle, cheeks, interramial region, a narrow band along sides 
bordering quills and spreading posteriorly to cover tail, hind 
legs and intercrural region, all of which are a dark brown, very 
nearly the bister of Ridgway. Feet a lighter shade of the same 
brown. Occasionally the brown suffusion extends over most of 
ventral surface. Quills dull horn-colour, each with a whitish 
sub-terminal area about 7 mm. in length, the extreme tip usually 
dark. Throughout the spiny area the whitish strongly pre- 
dominates, especially when animal is viewed from in front. 
Claws light yellowish horn-colour. 

Measurements. — Adult male from Schaf-el-Kab, Morocco 
(teeth much worn) : head and body, 206 ; tail, 26 ; hind foot, 32. 
For cranial measurements see Table, p. 132. 

Specimens examined. — Numerous specimens from Northern Africa ; also- 
an adult from "Andalucia," Spain; a young, less than half grown, from 
Elche, Alicante, Spain ; and a still younger specimen from Lecques, Var, 
France. 

Remarks. — Owing to the unsatisfactory nature of the Spanish 
and French material the status of the Continental European 
hedgehogs of the Erinaceus algirus group is at present doubtful. 
Should they prove to be identical with the North African form 
it would seem probable that they have been introduced within 
historic times. The specimen from Elche, though undoubtedly a. 
wild-bred animal, is too young to be positively determined as to 
geographical race. The same is even more true of that from Var. 
The adult from " Andalucia " lacks detailed history. Externally 
it resembles the African form as compared with E. algirus vagans, 
but the skull is small,* essentially as in the Balearic race. 

* Condylobasal length, 53 - 8; zygomatic breadth, 32 - 0; least inter- 
orbital breadth, 14 ■ 8 ; mandible, 41 • 2 ; upper tooth-row, 27 • ; lower tooth- 
row, 22-0. 

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ERINACEUS 



133 



1. Andaluoia, Spain. Lord Lilford (p). 94. 6. 11. 5. 

S juv. Blohe, Alicante. G. S. Miller (c). 8. 8. 4. 16. 

juv. al. Lecques, Var, France. Dr. P. Siepi (p). 98. 8. 25. 1. 

Eeinacbus algirus vagans Thomas. 

1901. Erinuceus algirus vagans Thomas, Proe. Zool. Soc. London, p. 38. 

Type in British Museum. 
1910. Erinaceus algirus vagans Trouessart, Paune Mamm. d'Europe, p. 39. 

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

Geographical distribution. — Balearic Islands. 

Diagnosis. — Condylobasal length of skull in individuals with 
worn teeth about 54 mm. ; furred area of body whitish through- 
out, with no evident dark wash on face, in interramial or 
intercrural regions, or on feet. 

Measurements. — Type (adult male) : head and body, 250 ; 
tail, 40 ; hind foot, 37. Adult male from Inca, Majorca . head 
and body, 218; tail, 31 ; hind foot, 35. For cranial measure- 
ments see Table, p. 132. 

Specimens examined. — One from Majorca and five from Minorca, 
Balearic Islands. 

Remarks. — The Balearic form of Erinaceus algirus is dis- 
tinguishable from the African race by its paler colour and smaller 
size. Its relationship to the animal occurring on the mainland 
of southern Spain is not at present clear. 

6. Inca, Majorca ; 300 m. O. Thomas & R. I. Pocock 0. 7. 1. 6. 
Balearic Islands. (c & p). 

4 <5, 9. San Cristobal, Minorca. O. Thomas & B. I. Pocock 0. 7. 1. 35-39. 

(c & p). 

(0. 7. 1. 36. Type of subspecies.) 



134 



CHIROPTERA 



Order CHIROPTERA. 

1779. Ghiroptera Blumenbach, Handbuch der Naturgeschichte, p. 74. 

Geographical distribution. — Practically cosmopolitan ; only 
absent from the treeless arctic and antarctic regions, and from 
the most remote islands of the Pacific and South Atlantic 
oceans. 

Characters. — Terrestial placental mammals with the anterior 
limbs modified for true flight, the fingers greatly elongated (third 
usually at least as long as head and body) and joined together 
by a membrane which extends to sides of body and legs ; 
shoulder girdle much more developed than pelvis, the sternum 
usually keeled ; knee directed backward. 

Remarks. — The order Ghiroptera, containing the only living 
vertebrates, except birds, capable of true flight, is the most 
sharply circumscribed of the main groups of mammals. Not 
only are its living members invariably distinct from all other 
recent forms, but the fossils also are, so far as known, equally 
well differentiated. Therefore no intermediate stage has yet 
been found connecting the bats with any other order. That 
they are, however, not distantly related to the Insectivora, is 
shown by numerous peculiarities of structure, among others the 
relatively simple character of the brain. Two sub-orders are 
recognized among the recent members of the order, the 
Megachiroptera, not represented in Europe, with less highly 
modified skeleton of fore limb, and more modified teeth, and the 
Microchiroptera with more highly developed wing and in most 
instances more primitive teeth. 

Sub-Order MICROCHIROPTERA. 

1821. Insectivora Gray, London Medical Repository, xv, p. 299, April 1, 

1821. 
1872. Animalivora Gill, Arrangement of the Families of Mammals, p. 16, 

November, 1872. 
1875. Microchiroptera Dobson, Ann. and Mag. Nat. Hist., 4th ser., xvi, 

p. 346, November, 1872. 
1878. Microchiroptera Dobson, Catal. Chiropt. Brit. Mus., p. 2. 
1907. Microchiroptera Miller, Families and Genera of Bats, p. 78, June 29, 

1907. 

Geographical distribution. — The same as that of the order. 
In Europe north to the limits of tree growth, west to Ireland 
and the Azores. 



MICROCHIEOPTEEA 



135 



Characters. — Anterior limb very highly modified, the second 
finger scarcely if at all independent of third, its ungual phalanx 
never present, the humerus with trochiter and trochin large, the 
former usually articulating with scapula ; mandible with angular 
process well developed, long and narrow ; teeth usually not 
modified for frugivorism (never in European species), the cheek- 
teeth of upper and lower jaws very different from each other 
(except when excessively reduced, as in the South American 
Desmodontidse) ; margin of ear not forming a ring ; tragus usually 
present. Mammse in all European genera, p 1-1 = 2. 

Bemarlcs. — The sub-order Microchiroptera is essentially cos- 
mopolitan in distribution. Though more highly modified than 
the Megachiroptera in wing structure, the members of this group 
for the most part retain the primitive tuberculo-sectorial type of 
molar tooth, though certain South American frugivorous forms 
show the stages through which the Megachiropterine molars 
have probably passed. At present 17 families and nearly 150 
genera are recognized ; the species are too imperfectly known to 
permit any approximate estimate of their number. Three 
families and ten genera are found in Europe. 

KEY TO THE EUROPEAN FAMILIES AND SUB-FAMILIES 
OF MICBOCHIBOPTEBA. 

Tragus absent ; muzzle with leaf -like outgrowths ; 
premaxillaries represented by palatal branch 
only, not fused with surrounding parts (often 

lost in prepared specimens) Bhinolophidai, p. 136. 

Tragus present ; muzzle without leaf-like out- 
growths ; premaxillaries represented princi- 
pally (entirely in European genera) by nasal 
branch, very early and completely fused with 
surrounding parts. 
Fibula robust, its diameter about half that of 
tibia ; tail projecting conspicuously beyond 
hinder edge of narrow interfemoral mem- 
brane Molossidas, p. 276. 

Fibula very slender, its diameter much less than 
half that of tibia ; tail scarcely or not pro- 
jecting beyond hinder edge of broad inter- 
femoral membrane Vespertilionidss, p. 165. 

Presternum with median lobe much smaller 
than body of bone ; coracoid curved out- 
ward ; second phalanx of third ringer not 

specially elongated Vespertilioninse, p. 165. 

Presternum with median lobe larger than 
body of bone ; coracoid straight, directed 
inward; second phalanx of third finger 
nearly three times as long as first Miniopterinx, p. 26S. 



136 CHIEOPTERA 

KEY TO THE GENERA OP EUROPEAN BATS. 

(A wholly artificial key based primarily on external characters.) 

Muzzle with leaf-like outgrowths Rhinolophus, p. 137. 

Muzzle without leaf-like outgrowths. 

Tail projecting conspicuously beyond membrane ... Nyctinomus, p. 276. 
Tail not projecting conspicuously beyond membrane. 
Ears joined. 

Ear longer than head Plecotus, p. 256. 

Ear shorter than head Barbastella, p. 263. 

Ears separate. 

Second phalanx of third finger nearly three 

times as long as first Miniopicrus, p. 268. 

Second phalanx of third finger less than twice 
as long as first. 
Fifth finger about as long as metacarpal of 

fourth or third Nyctalus, p. 242. 

Fifth finger much longer than metacarpal of 
fourth or third. 
Ear wider than high, its lower margin 
forming a small pocket near angle of 

mouth Vespcrtilio, p. 238. 

Ear higher than wide, its lower margin 
not forming pocket near angle of 
mouth. 

Upper cheek-teeth 6-6 Myotis, p. 166. 

Upper cheek-teeth less than 6-6. 

Upper cheek-teeth 5-5 Pipistrellus, p. 202. 

Upper cheek-teeth 4-4 Epteshus, p. 224. 

Family RHINOLOPHIDiE. 

1827. Rhinolophina Lesson, Man. de Mammalogie, p. 81 (part). 
1857. Phyllostomata Blasius, Saugethiere Deutschlands, p. 26. 
1866. RMnolophidai Gray, Proc. Zool. Soc. London, p. 81 (part). 
1878. Rhinolophidai Dobson, Catal. Ghiropt. Brit. Mus., p. 100 (part). 
1907. Rhinolophidse Miller, Families and Genera of Bats, p. 106, June 29, 
1907. 

Geographical distribution. — Tropical and temperate portions of 
the Old World from Ireland east to the Philippine Islands, 
Solomon Islands and north-eastern Australia. In Europe north 
to northern England and the Baltic coast of Germany. 

Characters. — Ear without tragus ; muzzle with conspicuous 
leaf-like cutaneous outgrowths (fig. 26) consisting of a horizontal 
anterior horseshoe, a perpendicular median sella, and a posterior 
erect lancet; skull with premaxillaries represented by palatal 
branches only, the two bones partly cartilaginous and not fused 
with surrounding parts (often lost in prepared specimens) ; 
shoulder girdle highly abnormal, the seventh cervical and first 
dorsal vertebrae, first and second ribs, and presternum fused into a 
continuous ring ; secondary articulation of humerus with scapula 
small but distinct ; fibula thread-like ; foot normal, the hallux 
with two phalanges, the other toes with three. 

Remarks. — The ffliinolophidse are the most widely distributed 



EHINOLOPHUS 



"137 



of the Old World leaf-nosed bats, and the only family known 
to occur in Europe. Notwithstanding its extensive distribution 
and its large number of species the group is represented by a 
single genus. 

Genus RHINOLOPHUS Lacepede. 

1799. Bhinolophus Lacepede, Tabl. des div. sousdiv. ordres et genres des 

Mammiferes, p. 15 (femcm-equinum). 
1836. Rhinocrepis Gervais, Diet. Pittoresque d'Hist. Nat. IV, pt. 2, p. 617 

(attributed to Geoflroy and Ouvier, Mag. Encyclopedique, 1795, 

tut the name does not occur in the paper alluded to). 
1847. Aquias Gray, Proc. Zool. Soc. London, p. 15 (luctus and trifoliatus) . 
1857. Bhinolophus Blasius, Saugethiere Deutschlands, p. 26. 
1866. Phyllotis Gray, Proc. Zool. Soc. London, p. 81 (philippcnsis) not 

Phyllotis Waterhouse, 1837. 
1866. Caelophyllus Gray, Proc. Zool. Soc. London, p. 427 (ccelophyllus). 
1878. Bhinolophus Dobson, Gatal. Chiropt. Brit. Mus., p. 100. 
1901. Euryalus Matschie, Sitz.-Ber. Gesellsch. naturforsch. Preunde, 

Berlin, p. 225 (mehelyi). 
1904. Euryalus Matschie and Andersen, Sitz.-Ber. Gesellsch. naturforsch. 

Freunde, Berlin, p. 71 (euryala group). 
1907. Bhinolophus Miller, Families and Genera of Bats, p. 108, June 29, 

1907. 

Type species. — Vespertilio ferrum-equinum Schreber. 
Geographical distribution. — Same as that of family (p. 136). 
Characters. — Dental formula: i^ ^, c^J, pi» |~? , m |^f = 32. 

2, 2, XI-* 1 oo o <j 

Upper incisor very small, but usually well formed and with 
distinct rounded crown with slight cusp on inner side. Lower 
incisors trifid, the outer larger than inner, the four teeth 
forming a continuous row between canines. Upper canine 
heavy, but without secondary cusps or conspicuous cingulum. 
Lower canine rather weak. Anterior upper premolar (pm 3 ) and 
middle lower premolar (pm 3 ) small, functionless, usually crowded 
quite out of tooth-row. Other teeth showing no special peculiari- 
ties ; m 1 and m 2 without hypocone, m 3 with five cusps and 
three commissures (in many species a rudimentary fourth com- 
missure), the crown area much more than half that of m 1 or 
m 2 . Skull with large brain-case and much shortened, globular ly 
inflated rostrum, beyond which the maxillaries, bearing the large 
canines, conspicuously project ; palate so deeply emarginated 
both anteriorly and posteriorly that its median length is less 
than least distance between tooth-rows. Tail well developed, 
extending to edge of wide interfemoral membrane. Calcar 
slender. Ears large, separate, without tragus. Muzzle with 
conspicuous leaf-like cutaneous outgrowths, consisting of a hori- 
zontal anterior horseshoe, a perpendicular median sella and an 
erect posterior lancet (fig. 26). 

Remarks. — Among European bats the members of the genus 
Bhinolophus are at once recognizable by the presence of the nose- 



138 



CHIEOPTERA 



leaf and absence of tragus. The skull differs from that of all 
other members of the fauna in the short, globularly inflated 
rostrum and long, projecting maxillaries, between which lie the 










FlS. 26. 

Noseleaf of Rhiaolophvx ferrum-equinum (a), R. hipposideros (b), R. euryale (c), 
and R. blasii (d). Nat. size. 

horizontal free premaxillaries (often lost in prepared specimens), 
About 100 forms have been described,* eight of which occur in 
Europe. 

KEY TO THE EUROPEAN FOBMS OP BHINOLOPHUS. 

Noseleaf with connecting process broadly 
rounded above ; skull with nasal swellings 
long, rising gradually above line of fore- 
head. 
Forearm over 50 mm. ; condylobasal length 
of skull over 20 mm. ; sella pandurate ; 
large upper premolar in contact with 

canine (Greater Horseshoe) R. ferrum-eqitimmi, p. 139. 

Wing relatively long ; forearm 54-58 mm., 
longest finger 84 to 92 mm. (Southern 

and central Continental Europe) R. f. ferrum-egvimim, p. 142. 

Wing relatively shorter ; forearm 52 to 
55 mm., longest finger 83 to 88 mm. 
(England) R. /• imidanus, p. 147. 



* See Andersen, Ann. and Mag. Nat. Hist., 7th ser., svi, pp. 648-662. 



RHIN0L0PHUS 139 

Forearm under 43 mm. ; oondylobasal length 
of skull under 16 mm. ; sella ouneate ; 
large upper premolar not in contact 

with canine (Lesser Horseshoe) B. hipposideros, p. 147. 

Greatest length of skull 14-5 to 15-5 mm. 

(Mediterranean region) r. ;,. minimus, p. 151. 

Greatest length of skull more than 
15 '5 mm. 
Forearm 36-3 to 39 mm. (England and 

Ireland) r. u. minutus, p. 154. 

Forearm 39 to 41 -7mm. (Central Europe) r. /,. hipposideros, p. 149. 
Noseleaf with connecting process acutely 
pointed above ; skull with nasal swellings 
short, rising abruptly above level of fore- 
head. 
First phalanx of fourth finger more than 
half as long as second; sella bluntly 
cuneate ; no marked contrast between 
crown areas of anterior and posterior 
lower premolars (Eastern Mediterranean 

region) R. blasii, p. 162. 

First phalanx of fourth finger less than half 
as long as second; sella parallel-sided, 
broadly rounded above ; a marked con- 
trast between crown areas of anterior 
and posterior lower premolars. 
Size smaller, forearm 44-6 to 49 mm., 
upper tooth-row 6-2 to 6 ■ 6 mm. ; 
gradation between phalanges of fourth 
finger abrupt (ratio of first to second 
about 38) ; point of lancet gradually 

narrowed, never linear R. euryale, p. 155. 

Size larger, forearm 48-6 to 51-4, upper 
tooth-row about 7 mm. ; gradation 
between phalanges of fourth finger 
less abrupt (ratio of first to second 
about 44) ; point of lancet linear R. mehelyi, p. 159. 

RHINOLOPHUS FERRUM-EQUINUM Schreber. 

(Synonymy under subspecies.) 

Geographical distribution. — From southern Japan and China, 
through the Himalayas, the Mediterranean sub-region (exclusive 
of Egypt), and central Europe to southern England (Andersen). 

Diagnosis. — Size largest of the European species (forearm 
more than 50 mm., condylobasal length of skull about 21 mm., 
mandible, 15 to 16 mm.); noseleaf with horseshoe less than 
10 mm. wide, the sella pandurate, the connecting process low, 
abruptly rounded ; fourth finger with first phalanx considerably 
more than half as long as second ; large upper premolar broadly 
in contact with canine, the small premolar minute (sometimes 
absent), completely external to tooth-row. 

External characters. — Size large and form rather heavy 
(among the European members of the genus). General outline 
of noseleaf a rather elongate ovate-pyriform, the width of horse- 
shoe slightly greater than distance from flat area at base of sella 



140 



CHIROPTERA 



to tip of lancet. Sella broadly rounded at tip, noticeably con- 
stricted somewhat above middle, the resulting outline pandurate. 
Connecting process rising slightly but evidently above level of 
sella, its upper border straight anteriorly, its tip rather abruptly 
rounded off. Ear large, extending when laid forward slightly 
beyond extremity of muzzle, abruptly narrowed to a rather acute 
recurved tip ; antitragal lobe less than half as high as conch, 
its width about equal to its height, its upper border nearly 
horizontal. Wings broad, the membrane attached to ankle. 
First phalanx of fourth linger noticeably more than half as long- 
as second. Foot slender, nearly half as long as tibia. 

Colour. — General effect a light greyish or drabby brown 
produced by varying combinations of the pale ecru-drab under 
colour and the darker hair tips, the region between ears, across 
shoulders and at sides of lumbar region usually paler than back. 
The colour of the tips is usually either a clear, rather light sepia, 
or a buffy brown resembling the wood-brown of Ridgway. While 
intermediate shades occur the extremes are more often met with, 
evidently representing two dichromatic phases. Underparts 
usually rather lighter than back but never approaching whitish. 
SJcull. — General outline of skull long and narrow, the breadth 
of brain-case much less than twice that across canines, and con- 
tained about 2 }j times in greatest length ; zygomata projecting 
slightly beyond general outline ; interorbital region greatly con- 
stricted and conspicuously hour-glass shaped ; occipital portion of 
brain-case noticeably overhanging foramen 
magnum and marked off from main portion 
by a slight transverse depression corre- 
sponding to suture between parietals and 
unusually large interparietal ; sagittal 
crest well developed, extending forward to 
narrowest portion of interorbital region : 
lambdoid crest low but evident ; auditory 
bullte small, covering less than half surface 
of large cochleae, between which the floor 
of brain-case is reduced to a longitudinal 
bridge less than 1 mm. in diameter ; 
mesopterygoid fossa slightly wider 
anteriorly than posteriorly, about 1£ 
times as long as wide, its rounded 
anterior margin at level of postero- 
internal angle of to 2 ; palate with an 
evident emargination on each side ex- 
tending between m 3 and mesopterygoid space ; anterior palatal 
emargination extending back to line joining protocones of 
anterior molars ; premaxillary ligulate, somewhat wider pos- 
teriorly than anteriorly, the inner border entire, the outer border 
with a deep almost circular emargination posteriorly, the two 
bones closely applied to each other along inner margin and to 




y^&sg&A 




Rhinolophusferrum-equinum . 
Nat. size. 



RHINOLOPHUS 



141 



bottom of palatal emargination posteriorly, but otherwise free; 
rostral inflation evident but low, its posterior border running 
gradually into that of interorbital region, its anterior margin 
over middle of anterior molar ; maxillary triangular in outline 
when viewed from the side, the heavy canine projecting forward 
and downward from its anterior apex ; anteorbital foramen small, 
over middle of second molar and directly beneath minute 
lachrymal foramen, the plate forming outer wall of canal 
thread-like, occasionally absent. 

Teeth. — Except for the minute premolars and upper incisors the 
teeth are robust and heavy relatively to size of skull. Upper 
incisor minute, low, the crown subterete, wider than root, lower 
externally than internally, each tooth placed near middle of 
oblique anterior border of premaxillary, the space between the 
two nearly double diameter of crown. Lower incisors strongly 
imbricated, forming a short, very convex row between canines, 
their crowns longer than high, compressed (the outer tooth less 
than the inner), deeply and equally trifid. Upper canine very 
large, and noticeably the highest tooth in the maxillary series, 
the root oblique, the shaft abruptly bent downward at level of 
well developed cingulum, its length along cingulum about three- 
fourths height ; cross section of shaft triangular, the inner surface 
flattened, slightly concave near cingulum, the posterior edge 
trenchant, the anterior edge narrowly sub-trenchant, the outer 
surface with well developed median longitudinal ridge and 
noticeable posterior concavity. Lower canine not so large as 
upper, the flattened surface of its shaft directed posteriorly to 
oppose front of upper canine in mastication. Anterior upper 
premolar minute, resembling upper incisor in both size and form, 
entirely external to tooth-row, occasionally absent. Posterior 
upper premolar large, closely crowded against canine, its crown 
area about equal to that of second molar, without secondary 
cusps, its main cusp intermediate in height between canine and 
metacone of first molar ; posterior border of crown slightly but 
evidently emarginate. Anterior lower premolar less than half as 
high as posterior premolar and with barely half its crown area, 
the two teeth crowded closely together between canine and first 
molar ; shaft of anterior tooth with slightly developed concave 
area on inner side, that of posterior tooth with better defined 
posterior concavity, its inner side convex ; middle lower premolar 
resembling anterior upper premolar, its position equally external 
to tooth-row. Upper molar with large though not unusually high 
protocone, behind which in m 1 and m 2 there is a low-lying 
heel (best developed in m 1 ), but no indication of a true hypocone ; 
paracone and metacone well developed, the latter slightly the 
higher in m 1 and m 2 , this reversed in m 3 ; styles and commis- 
sures in m 1 and m 2 well developed, forming a normal VV-pattern ; 
in m 3 the metastyle and fourth commissure are absent, and the 
third commissure is much reduced in length ; crown area of m 3 



142 



CHIEOPTBEA 



about two-thirds that of wr ; lower molars with protoconid 
noticeably higher than the other cusps, the posterior triangle 
slightly the wider of the two in m 1 and ro 2 , a little the narrower 
in m 3 ; behind entoconid the cingulum forms a minute though 
evident accessory cusp. 

Remarks. — Among the leaf-nosed bats of Europe this species 
is easily recognizable by its large size taken in connection with 
the low, rounded upper margin of the connecting process of sella. 
In It. mehelyi, which approaches it in size, the connecting process 
is sharply pointed above. Two geographical races are known, one 
occupying the Continental range of the species, the other confined 
to Great Britain. 



Rhinolophus ferrum-equinum ferrum-equinum Schreber. 

1774. Vespertilio ferrum-equinum Schreber, Saugthiere, i, pi. lxii, upper 

figures ; description, i, p. 174 under name : Die Hufeisennase 

(part). Prance ; based primarily on Daubenton. 
1776. Vespertilio equinus P. L. S. Miiller. Natursyst. Suppl. u. Regist.- 

Band, p. 20 (part), Prance. 
1779. [Vespertilio] perspicillatus ~Blumenba,ch, Handb. d. Naturgesch., p. 75 

(part : included the leaf -nosed bats of Europe and South America). 
1785. [Vespertilio'] ungula Boddaert, Elenchus Animalium, i, p. 71 

(Burgundy). 
1792. Vesp[ertilio] fer[rum]-equ[i'ium] major Kerr, Anim. Kingd., p. 99 

(not V. molossus major Kerr, I.e., p. 97), Prance. 
1798. Vespertilio hippocrepis Schrank, Pauna Boica, I, p. 64 (Renaming 

of ferrum-equinum, Schreber) . ' 
1803. Bhinolophus major Geoflroy, Catal. Mamm. Mus. Nat. d'Hist. Nat., 

Paris, p. 56 (Burgundy). 
1813. Bhinolophus unihastatus Geoffrey, Ann. Mus. d'Hist. Nat., Paris, 

xx, p. 257 (Prance). 
1829. ? Bhinolophus unifer Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. 

Thierwelt, I, p. 104 (nomen nudum). 
1857. Bhinolophus ferrum-equinum Blasius, Saugethiere Deutschlands 

p. 31. 
1863. [Rhinolophus ferrum-equinum] a var. germanicus Koch, Jahrb. des 

Vereins fur Naturkunde im Herzogthum Nassau, xvin, p. 522 

(Wiesbaden, Hessen-Nassau, Germany). 
1863. [Bhinolophus ferrum-equinum] B var. italicus Koch, Jahrb. des 

Vereins fur Naturkunde im Herzogthum Nassau, xvni, p. 523 

(Italy). 
1878. Bhinolophus ferrum-equinum Dobson, Catal. Ohiropt. Brit. Mus., 

p. 119 (part). 

1885. Bhinolophus unihastatus, var. homorodalmasiensis Daday, Orvos- 

Termeszettudomanyi ErtesetiS, Kolozsvar, x, p. 274 (Homorod- 
Almas cave, Hungary). 

1886. Bhinolophus unihastatus var. homorodalmasiensis Daday, Verhandl. 

u. Mittheilungen des Siebenbiirgischen Vereins f iir Naturwissensch. 
in Hermannstadt, xxxvi, p. 79. 

1887. Bhinolophus ferrum-equinum var. homorodensis Daday, Brtekezesek a 

Termeszettudomanyok Korebi'il, Budapest, xvi, pt. 7, p. 13 
(Renaming of homorodalmasiensis) . 



RHINOLOPHUS 143 

1904. Rhinolophus ferrum-equinum obscurus Cabrera, Mem. Soc. Espafi. 

Hist. Nat. Madrid, n, p. 257 (Valencia, Spain). 

1905. Rhinolophus ferrum-equinum typicus Andersen, Proo. Zool. Soc. 

London, 1905, n, p. 113, October 17, 1905. 

1905. Rhinolophus ferrum-equinum obscurus Andersen, Proc. Zool. Soc. 
London, 1905, n, p. 116, October 17, 1905. 

1910. Rhinolophus ferrum-equinum, R. ferrum-equinum obscurus and R. 
ferrum-equinum homorodensis Trouessart, Paune Mamm. d'Europe, 
pp. 4-5. 

Type locality. — Burgundy, France. 

Geographical distribution. — Central and southern Continental 
Europe. 

Diagnosis. — Wing relatively long ; forearm, 54 to 58 mm. ; 
third finger, 84 to 92 mm. 

Measurements. — Average and extremes of six males from 
Silos, Burgos, Spain : head and body, 62 ■ 1 (58-64 ■ 4) ; tail, 
39-9 (37 ■ 6-43) ; tibia, 23-2 (21-8-24); foot, 11-7 (11-13); 
forearm, 55-1 (54-8-55-4); third finger, 89-3 (88-92); fifth 
finger, 72-5 (72-73); ear from meatus, 24-9 (24-26). Average 
and extremes of five females from Silos, Burgos, Spain : head 
and body, 61-8 (60-66); tail, 38-8 (37-42); tibia, 23-1 
(22-6-24) ; foot, 11-7 (11-13) ; forearm, 56-2 (56-57) ; third 
finger, 89-6 (88-91); fifth finger, 73-4 (71-76); ear from 
meatus, 25 -1 (24-8-26). Two males from Granada, Spain 
forearm, 54 and 54. Two females from the same locality 
forearm, 56 and 58. Two males from Elche, Alicante, Spain 
forearm, 54 and 55. Average and extremes of six adults 
(3 males and 3 females) from St. Genies, Gard, Prance : tibia, 
24-1 (23-4-24-6); foot, 12-] (11-8-12-8); forearm, 554 
(54-65) ; third finger, 87-3 (84-91) ; fifth finger, 71-5 (69-74). 
Female from Marseilles, France : forearm, 57. Male and female 
from near Genoa, Italy : forearm, 57 and 57. Two females from 
Rimini, Italy : forearm, 53 and 57. Female from Siena, Italy : 
forearm, 54. Male from Rome, Italy : forearm, 56. Male from 
the Parnassus region, Greece : forearm, 54. Adult female from 
Tubingen, Wurtemberg, Germany : head and body, 63 ; tail, 37 ; 
tibia, 24-4; foot, 11; forearm, 56-6; third finger, 88; fifth 
finger, 72; ear from meatus, 24; width of ear, 16. Adult 
female from Ofener Mts., Hungary : head and body, 61 ; tail, 40 ; 
tibia, 24; foot, 11-6; forearm, 56; third finger, 87; fifth 
finger, 73; ear from meatus, 23'4; width of ear, 16. Two 
males from Herkulesbad, Hungary : forearm, 56 • 6 and 58. Four 
females from Herkulesbad, Hungary: forearm, 56, 56-4, 56-6 
and 57 - 4. For cranial measurements see Table, p. 144. 

Specimens examined. — One hundred and forty-one, from the following 
localities : — 

Portugal : Cintra, 1. 

Spain : Silos, Burgos, 22 ; Granada, 4 ; Elche, Alicante, 3 ; San 
Cristobal, Minorca, Balearic Islands, 4. 



144 



CHIEOPTEEA 





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146 



CHIEOPTEEA 



France : Troubate, Hautes-Pyrenees, 6 ; St. Genies, Gard, 43 (Mottaz) ; 
Marseilles, 1 (U.S.N.M.) ; Meounes, Var, 1. 

Italy: Turin, 1 ; near Genoa, 8 (B.M., U.S.N.M. and Genoa); Grotta 
de Isoverde, Liguria, 7 (B.M. and Genoa) ; Rimini, 2 (U.S.N.M.) ; Siena, 1 
(U.S.N.M.); near Rome, 5 (U.S.N.M. and Genoa); no exact locality, 1 ; 
Sicily, 2. 

Corsica : Commune di Barbaggio, 1. 

Sardinia : Sassari, 1. 

Greece : Parnassus region, 1 (U.S.N.M.) ; Island of Syra, Cyclades, 

1 (U.S.N.M.) ; Mt. Goria Monastery, Crete, 3. 

Germany : Tubingen, Wiirtemberg, 1. 

Switzerland : Geneva, 6 (U.S.N.M. and Mottaz) ; Boudry, Neuchatel, 

2 (Mottaz) ; Tremona, Tioino, 1 ; Mendrisio, Ticino, 2 ; Lugano, Ticino, 3. 

Austria-Hungary : Herkulesbad, Hungary, 6 ; Ofener Mts., Hungary, 1. 

Remarks. — Spanish specimens of JRhinoloplius ferrum-equinwu 
have been regarded by both Cabrera and Andersen at represent- 
ing a peculiar race, B. f. obscurus, distinguished by small size. 
The material which I have examined, however, indicates that 
the Iberian animal cannot be treated as distinct. 



6. 


Cintra, 500 mm. Portugal. 


0. Thomas (c&p). 


98. 2. 2. 1. 


6al. 


Silos, Burgos, Spain. 




N. & S. Gonzalez 

(c). 
G. S. Miller (c). 
G. S. MiUer (c). 
G. S. MiUer (a). 


8. 7. 7. 38-43. 


2 6, 2 9. 


Silos, Burgos. 




8. 8. 4. 1-4. 


6,9. 


Granada, 2250 ft. 




8. 8. 4. 8-9. 


2 6, 9. 


Elche, Alicante, 20 m. 




8. 8. 4. 5-7. 


3,2 9. 


San Cristobal, Minorca, 
learic Islands. 


Ba- 


0. Thomas and 
R. I. Pocock 
(c & P). 


0. 7. 1. 24-26. 


<5al. 


San Cristobal, Minorca. 




0. Thomas and 
R. I. Pocock 
(c & P). 


0. 7. 1. 68. 


6 6. 


Troubate, Hautes-Pyrenees, 


0. Thomas (p). 


6. 4. 1. 1-6. 




France. (A. Robert.) 








lal. 


Meounes, Var. 




Dr. K. Jordan 
(c & p). 


8. 3. 15. 1. 


9al. 


Turin, Italy. 




Prof. BoneUi (p). 






?al. 


Isoverde, Genoa. 




O.Thomas (c&p). 


88. 12. 7. 1. 


Skeleton. 


Italy. (Prince Bonapan 


ie.) 


Tomes CoUection. 


7. 1. 1. 727. 


2<Jal. 


Sicily. 












cJal. 


Barbaggio, Corsica. 




Dr. C. I. Forsyth 


6. 4. 14. 1. 



6 al. Sassari, Sardinia. 

<5, 2 ?. Mt. Goria Monastery, Crete. 
9 al. Tiibingen, Wiirtemberg, Ger- 
many. 
9. Tremona, Ticino, Switzerland. 

(E. H. Zollihofer.) 
2 9. Mendrisio, Ticino. (E. H. 

Zollihofer.) 
3. Lugano, Ticino. (E.H. Zolli- 
hofer.) 
2 ci, 4 9. Herkulesbad, Hungary. 

9 al. Ofener Mts., Budapest. 



Major (c&p). 
Marquis G. Doria 6. 12. 1. 11. 

Miss D. Bate (c). 5. 12. 2. 1-3. 
Dr. A. Giinther 66. 2. 1. 1. 

(*)■ 
0. Thomas (p). 



0. Thomas (p). 
0. Thomas (p). 



2. 8. 4. 1. 
2. 8. 4. 2-3. 
4. 4. 5. 1-3. 
7. 9. 16. 1-6. 



Hon. N. C. Roths 

child (p). 
Budapest Museum 94. 7. 18. 5. 

(■)• 



RHINOLOPHUS 



147 



Rhixolophus pbeeum-equinum insulands Barrett-Hamilton. 

1910. Rhinolophus ferrum-equimminsulanus Barrett-Hamilton, Ann. and 
Mag. Nat. Hist., 8th ser., v, p. 292, March, 1910. Type in British 
Museum. 

1910. Rhinolophus ferrum-equinum insulanus Trouessart, Faune Mamm. 
d'Europe, p. 273. 

Type locality. — Cheddar, Somersetshire, England. 

Geographical distribution. — Central and southern England. 

Diagnosis. — Wing relatively short ; forearm, 52 to 55 ; third 
finger, 83 to 88. 

• . Measurements. — Type (adult male) : head and body, 67 ; tail, 
37 ; tibia, 23-4 ; foot, 11 ; forearm, 54 ; third finger, 86 ; fifth 
finger, 70; ear from meatus, 23 • 6. Average and extremes of 
seven males from the type locality : head and body, 65 (63-67) ; 
tail, 36-9 (35-40); tibia, 22-8 (22-24); foot, 11-2 (11-12); 
forearm, 53-5 (52-54) ; third finger, 85-5 (83-87) ; fifth finger, 
69-9 (69-71) ; ear from meatus, 24-2 (23-25). Three females 
from the type locality : head and body, 64, 66 and 64 ; tail, 35, 
34 and 36; tibia, 23-4, 23 and 23; foot, 10-6, 12 and 12; 
forearm, 55, 54 • 6 and 54; third finger, 87, 86 and 88 ; fifth 
finger, 72, 72 and 72 ; ear from meatus, 25, 24 and 24. For 
cranial measurements see Table, p. 144. 

Specimens examined. — Twenty-four, from the following localities in 
England : — Cheddar, Somersetshire, 10 ; Wells, Somersetshire, 10 ; Dorset- 
shire, 3 ; Bonchurch, Isle of Wight, 1. 

6, 9. Cheddar, Somerset, J. A. Coward (c & p). 7. 1. 10. 1-2. 

England. 

6 al. Cheddar. J. A. Coward (o & p). 7. 1. 10. 3-10. 

(J, 2 Sal. Wells. Hon. N.C.Rothschild 5.1.23.1-3. 

(*)■ 

3 6, 6 juv., 9 st. Wells. Hon.N. C.Rothschild /l. 9. 3. 1-4. 

(p). \2. 9. 6. 1. 

6,9. Wells. Hon. N.C.Rothschild 11.1.3.3-4. 

(*)■ 
■1 9. Dorset. W. M. Hardy (c & p). 11. 1. 3. 1-2. 

1. Bonchurch, Isle of Rev.C. A.Bury (c&p). 11.1.3.388. 

Wight. 

RHINOLOPHUS hipposideros Bechstein. 

(Synonymy under subspecies.) 

Geographical distribution. — -From Gilgit through the Mediter- 
ranean sub-region and central Europe to Ireland ; north in 
•continental Europe to the Baltic, and in Great Britain to about 
the southern border of Scotland. 

Diagnosis. — Size small, the forearm less than 43 mm. in 
length, condylobasal length of skull about 14 to 15 mm., mandible 
about 10 mm.; noseleaf (fig. 26 6) with bluntly cuneate sella 
And low, broadly rounded connecting process ; fourth finger with 

l 2 



148 



CHIROPTBEA 



first phalanx slightly more than half as long as second ; large 
upper premolar separated from canine by a noticeable space 
occupied by the well developed small premolar, which lies per- 
fectly in the tooth-row. 

External characters. — A much smaller, more delicately formed 
animal than Bhinolophus ferrum-equinum. General outline of 
noseleaf narrower than in B. ferrum-equinum, the width of horse- 
shoe less than distance from flat area at base of sella to tip of 
lancet ; sella narrowly rounded at tip, the sides straight, slightly 
convergent above, the resulting outline bluntly cuneate ; connect- 
ing process essentially as in the larger species but relatively 
wider and lower, its upper extremity about on level with that of 
sella ; lancet slender, scarcely or not contracted at middle, the 
tip cuneate. Ear when laid forward extending about 5 mm. 
beyond extremity of muzzle, the narrow tip abruptly curved 
backward ; antitragal lobe more than half as high as conch, its 
width less than its height, its upper margin noticeably oblique. 
Wings and feet essentially as in B. ferrum-equinum. 

Colour. — The colour does not differ appreciably from that of 
Bhinoloplius ferrum-equinum. 

Skull. — The skull differs from that of Bhinoloplius ferrum- 
equinum chiefly in its conspicuously smaller size (greatest length 
about 16 mm. instead of about 24 mm.). There are also some 
slight peculiarities in form, the principal of which is the greater 
contrast between width of brain-case and anterior maxillary 
region, the breadth of former being about twice that of latter. 
As the breadth of brain-case is contained distinctly less than 
2|- times in greatest length it is probable that the difference 
between the two animals is due rather to enlarged brain-case in 
the smaller than to widened palate in the larger. Sagittal crest 
slightly developed ; lambdoid crest obsolete. Nasal region 
relatively more inflated than in B. ferrum-equinum, particularly 
at side of nares, but dorsal outline not rising abruptly above 
level of interorbital region. Mesopterygoid fossa so conspicu- 
ously widened anteriorly that it occupies almost entire extent of 
palate between posterior molars, leaving no space for lateral 
palatal emarginations, a peculiarity by which the skull may be 
distinguished from that of all other European members of the 
genus. 

Teeth. — Though in genera] resembling those of Bhinoloplius 
ferrum-equinum, apart from their much smaller size, the teeth of 
B. hipposideros show several notable peculiarities. Upper canine 
relatively small, its apex in line with large cusps of molars, and 
slightly below that of posterior premolar ; lower canine corre- 
spondingly short ; anterior upper premolar a well developed 
functional tooth lying perfectly in tooth-row, with crown area 
equal to nearly half that of canine, its shaft subterete though 
flattened posteriorly, and only a little less than half as high as 
main cusp of large premolar ; anterior and posterior lower pre- 



RHINOLOPHUS 



149 



molars less contrasted in size than in the larger animal, and 

separated from each other ,by a slight space in which lies 

the minute middle premolar, less 

crowded outward than in S. ferrum- 

equinum. Large upper premolar 

and upper molars essentially as in 

B. ferrum-equinum except that in 

tn 3 there is a distinct meta- 

style and fourth commissure, 

and area of tooth is nearly 

equal to that of m 2 . Lower 

molars as in Bhinolophus 

ferrum-equinum, except that 

in m 3 , almost exactly resem- 
bles the other teeth, the second 

triangle having undergone 

practically no reduction. 

Measurements. — Head and 
body about 40 mm., tail about 
30 mm., forearm, 34-5 to 
41 • 7 mm., condylobasal length 
of skull, 13 • 8 to 15 mm. 
(greatest length 14-5tol6'2). 
Details under subspecies. 

Bemarhs. — Three imperfectly differentiated forms of Bhino- 
lophus hipposideros occur in Europe : a larger central race, a 
smaller Mediterranean race, and an intermediate form peculiar 
to Great Britain and Ireland. Without examination of much 
more complete material than that now available it is impossible 
to reach any wholly satisfactory conclusion as to the status and 
interrelationships of these forms. The characters here given are 
those published by Andersen in 1905.* 





Fig. 28. 
Ithinolophus hipposideros. Teeth X 10_ 



Rhinolophus hipposideros hipposideros Bechstein 

1789. VespertiUo ferrum-equinum B. , Die kleine Huf eisennase, Bechstein. 

Gemeinn. Naturgesch. Deutsehlands, i, 1st ed., p. 186. 
1792. Vcsp[ertilio~] fer\rum\-eqit,\inum\ minor Kerr, Anim. Kingd., p. 99 

(not V. molossus minor Kerr, I.e., p. 97) (France). 
1800. VespertiUo hipposideros Bechstein, Thomas Pennant's Allgemeine 

Uebersicht der vierfiissigen Thiere, n, p. 629. 
1803. Bhinolophus minor Geoffroy, Oatal. Mamm. Mus. Nat. d'Hist. Nat., 

Paris, p. 57 (Neighbourhood of Paris). 
1813. Bhinolophus bihastatus Geoffroy, Ann. Mus. d'Hist. Nat., Paris, xx, 

p. 259 (Neighbourhood of Paris). 
1816. ? Phyllorhina minuta Leach, Syst. Catal. Spec. Indig. Mamm. and 

Birds Brit. Mus., p. 5 (nomen nudum : " Small Leafnose"). 



* For further discussion of the subject see Mottaz, Mem. Soc. Zool. de 
France, Paris, xx, pp. 21-22, September, 1907 ; Andersen, Ann. and Mag. 
Nat. Hist., 7th ser., xx, pp. 384-389, November, 1907. 



1 50 CHIEOPTBEA 

1829. ? Bhinolophus bifer Kaup, Entw.-Gesch. u. Natiirl. Syst. Burop. 

Thierwelt, I, p. 104 (nomen nudum). 
1857. ? Bhinolophus hipposideros Blasius.'Saugethiere Deutschlands, p. 29. 
1863. [Bhinolophus hipposideros'] a var. typus Koch, Jahrb. des Vereins fur 

Naturkunde im Herzogthum Nassau, xvm, p. 530 (Wiesbaden). 
1863. [Bhinolophus hipposideros] var. alpinus Koch, Jahrb. des Vereins 

fiir Naturkunde im Herzogthum Nassau, xvm, p. 530 (Alps). 
1870. Bhinolophus eggentioffner Pitzinger, Sitzungsber. kais. Akad. 

Wissensch. Wien, Math.-Naturwiss. Classe, lxi, Abth. i, p. 151 

(MS. synonym of bihastatus). 
1878. Bhinolophus hipposideros Dobson, Oatal. Chiropt. Brit. Mus., p. 117. 

1885. Bhinolophus bihastatus var. kisnyiresiensis Daday, Orvos-Termesz- 

ettudomanyi Ertesito, KojLozsvar, x, p. 274 (Kis-Nyires, Szolnok- 
Dobaka, Hungary). 

1886. Bhinolophus bihastatus var. kisnyiresiensis Daday, Verhandl. u. 

Mittheilungen des Siebenburgischen Vereins fiir Naturwissensch. 
in Hermannstadt, xxxvi, p. 80. 

1887. Bhinolophus hipposideros var. troglophilus Daday, Ertekezesek a 

Termeszettudomanyok Korebol, Budapest, xvi, pt. 7, p. 8 (Re- 
naming of kisnyiresiensis). 

1904. Bhinolophus euryale helvetica Bretscher, Vierteljahrsschrift der 

Naturforsch. Gesellsch. in Zurich, xlix, p. 256. See Mottaz, 
Bull. Soc. Zool., Geneve, ±., p. 172, 1908 (Baar, Zug, Switzerland). 

1905. Bhinolophus hipposiderus typicus Andersen, Proc. Zool. Soc. London, 

1905, n, p. 141, October 17, 1905. 
1910. Bhinolophus hiijposiderus Trouessart, Faune Mamm. d'Europe, p. 9. 

Type locality. — France. 

Geographical distribution. — Central Europe, north of the Alps, 
east through Armenia and north-west Persia to the Himalayas. 

Diagnosis. — Forearm* usually 39 to 41 • 7 mm.; greatest 
length of skullf about 16 mm. 

Measurements. — Two males from Strass near Burgheim, 
Bavaria, Germany: tibia, 17 and 18-4; foot, 7 - 8 and 7*4; 
forearm, 38-8 and 39 • 2 ; third finger, 61 and 60 ; fifth finger, 
54 and 53. Female from the same locality : tibia, 17 • 4 ; foot, 
7-4; forearm, 38-8; third finger, 60; fifth finger, 53. The 
six following extremes of forearms of European specimens are 
given by Andersen (I.e. p. 142) : 1ST. Bulgaria (1), 39 ; Roumania 
(13), 39 to 41-2 ; Transsylvania (2), 40 to 41 ; S. Carpathians 
(1), 39-3; Schlangenbad, Nassau, Germany (2), 40 to 40" 1; 
Strassburg, Germany (3), 39 to 40*1. Forty-nine males from 
the vicinity of Geneva, Switzerland (Mottaz collection),! 37 ' 3 
to 39. Thirty females from the same locality (Mottaz collection), J 
38-5 to 40-6. Four males from the same locality (U.S.N.M.), 
37 to 38-4 Six females from the same locality (U.S.N.M.), 
37 -4 to 40. Adult female from Dions, Gard, France (Mottaz 
collection), 40. For cranial measurements see Table, p. 152. 

* Bases of metacarpals included, 
t Prom back of occiput to front of canine. 

X Measured by Chas. Mottaz (base of metacarpals included) and verified 
by Gerrit S. Miller. 



RHINOLOPHUS 151 

Specimens examined. — One hundred and seventeen, from the following 
localities: — 

France : Dions, Gard, 1 (Mottaz) ; St. Gergues, Haute-Savoie, 1. 

Germany : Strass, near Burgheim, Bavaria, 3 ; Mainz, 1 (Strassburg) ; 
Bitsoh, Alsace, 1 (Strassburg). 

Austria-Hungary : Hatszeg, Hunyad, Hungary, 1. 

Switzerland: Near Geneva, 94 (B.M., U.S.N.M. and Mottaz); St. 
Moritz, 1 ; Thayngen, Schaffhausen, 2 (U.S.N.M.) ; Roggwil, Thurgau, 
7 (B.M. and U.S.N.M.) ; Canton Thurgau, 5 (B.M. and U.S.N.M.). 

£. St. Cergues, Haute-Savoie, A.Robert (c &p). 5.4.9.1. 

France. 

2 9. Burgheim, Bavaria, 375 m. Lord Lilford (p). 11. 1. 1. 117- 

Germany. 118. 

<J. Hatszeg, Hunyad, Tran- C.G.Danford(c). 8.11.8.1. 

sylvania. 

3 6. Grand Pre, Geneva, Swit- C. Mottaz (c & i>). 6. 2. 6. 1-3. 

zerland. 
1. St. Moritz, Grisons. Leon 0. Galliard 75. 9. 20. 3. 

(o & p). 
3 6, 2 9, imm. Roggwil, Thurgau. (E. H. 0. Thomas (p). 2. 8. 4. 4-9. 
ZolUkofer.) 
?al. Thurgau. (E.H. ZolUkofer.) 0. Thomas (p). 2.8.4.55. 

Rhinolophus hipposideros minimus Heuglin. 

1861. Rhinolophus minimus Heuglin, Nov. Act. Acad. Caes. Leop.-Carol., 
xxix, Abhandl. vin, p. 6 (articles separately paged) (Keren, 
Abyssinia). 

1863. [Rhinolophus hipposideros'] y var. pallidus Koch, Jahrb. des Vereins 
fiir Naturkunde im Herzogthum Nassau, xviii, p. 531 (Mediter- 
ranean region). 

1904. Rhinolophus phasma Cabrera, Mem. Soc. Espan. Hist. Nat., Madrid, 
ii, p. 252 (Madrid, Spain). 

1904. Rh[inolophus] h[ipposiderus] minimus Andersen, Ann. and Mag. Nat. 

Hist., 7th ser., xiv, p. 456, December, 1904. 

1905. Rhinolophus hipposiderus minimus Andersen, Proc. Zool. Soc. 

London, 1905, n, p. 140, October 17, 1905. 
1910. Rhinolophus hipposiderus minimus Trouessart, Faune Mamm. 
d'Europe, p. 10. 

Type locality. — Keren, Abyssinia. 

Geographical distribution. — Mediterranean region. 

Diagnosis. — Forearm usually 34 ■ 7 to 38 mm. ; greatest 
length of skull, 14-5 to 15 ■ 5 mm. 

Measurements. — The following measurements of forearms are 
given by Andersen (I.e. p. 141) : Keren (type), 36 • 3 ; Cyprus (6), 
34-7 to 37-7; Malta (8), 36 to 37; Ostia, Italy (2), 35*7' to 
36-8; Corsica (1), 37-7; Balearic Islands (7), 36-2 to 37-6; 
Seville, Spain (1), 37-7; Cintra, Portugal (1), 36-2. Specimens 
in U.S. National Museum : Ficuzza, Sicily ( $ ), 35 • 2 ; Rome ( <J ), 
36-4; Siena (3 $ ), 36 '8 to 38; Marseilles, France (?), 38; 
Dions, Gard, France ( <J ), 34 ■ 8, ( ° ), 36 • 6 Two females from 
Ax-les-Thermes, Ariege, France, 36 ■ 6 and 38 '4. Two males 
from Silos, Burgos, Spain, 36 and 37. Female from the same 
locality, 38. Five males from Ticino, Switzerland, 37 - 
(36 '2-38). Mottaz collection: St. Genies, Gard, France (15, 



152 



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both sexes), 36 • 4 to 38 ■ 2) ; Sardinia (2 ? ), 37 ■ 4 to 37 ■ 8. For 
cranial measurements see Table, p. 153. 

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

Spain: Inoa, Majorca, Balearic Islands, 1; San Cristobal, Minorca, 
Balearic Islands, 3 ; Elche, Alicante, 2 ; Silos, Burgos, 5 ; Seville, 1. 

Portugal : Cintra, 1. 

France: Ax-les-Thermes, Ariege, 5; St. Genies, Gard, 17 (U.S.N.M. 
and Mottaz) ; Marseilles, 1 (U.S.N.M.). 

Switzerland : Locarno, Ticino, 4 (U.S.N.M.) ; Gordola, Ticino, 2 
(U.S.N.M.) ; Minusio, Ticino, 6 (U.S.N.M.). . 

Italy: Liguria, 3 (U.S.N.M.); Siena, 3 (U.S.N.M.); Paenza, 1 
(U.S.N.M.) ; Borne, 3 ; Ostia, Rome, 2 ; Picuzza, Sicily, 6. 

Sardinia : No exact locality, 2 (Mottaz) ; Zimmigas, Siliqua, 2. 

Corsica : Dintomi di Patrimonio, 3 ; no exact locality, 1. 

Malta : 7. 

9. Inca, Majorca, Balearic 0. Thomas and R. I. 0. 7. 1. 1-2. 
Islands. Pocock (c & p). 

6, 9. San Cristobal, Minorca. 0. Thomas and R. I. 0. 7. 1. 27-28. 

Pocock (c & p). 

1 al. San Cristobal, Minorca. 0. Thomas and R. I. 0. 7. 1. 69. 

Pocock (c & pi 

9. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 11. 

6. Elche, Alicante, Spain. G. S. Miller (c). 8. 8. 4. 12. 

£ al. Cintra, 500 m. Portugal. O. Thomas (c & p). 98. 2. 2. 57. 

6, 9. Ax-les-Thermes, Ariege, G. S. Miller (o). 8.8.4.124- 

2400 ft. Prance. 126. 

S. Borne, 37 m. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. 42. 

if)- 
2. Ostia, Rome. Dr. L. Sambon (c & p). 1. 1. 2. 5-6. 

6 9. Picuzza, Sicily. (A.Robert.) 0. Thomas (p). jg ®" * - ]^ 16 - 

(5, 9 al. Zimmigas, Siliqua, Sardinia. Marquis G. Doria (p). 6.12.1.20-21. 

(R. Meloni.) 
3 <$, 2 9. Malta. (J. Micalleff.) Lord Lilford (p). 11. 1. 1. 119- 

123 
2. Malta. Lord Lilford (p). 95. 3.' 2. 1-2. 

Rhinolophus hipposideeos minutus Montagu. 

1808. Vespertilio minutus Montagu, Trans. Linn. Soc. London, ix, p. 163. 
1905. Rhinolophus hipposiderus minutus Andersen, Proc. Zool. Soc. 

London, 1905, u, p. 142, October 17, 1905. 
1910. Rhinolophus hipposiderus minutus Trouessart, Faune Mamm. 

d'Europe, p. 10. 

Type locality. — Wiltshire, England. 

Geographical distribution. — England and Ireland. 

Diagnosis. — Forearm, 36 '3 to 39 mm . • greatest length of 
skull about 16 mm. 

Measurements. — Forearm in 30 English and Irish specimens 
measured by Andersen (I.e. p. 142), 37 '6 (36-3 to 39). For 
cranial measurements see Table, p. 152. 

Specimens examined. — Sixteen, from the following localities in England : 
— Great Grimsby, Lincolnshire, 1; Bowdon, Cheshire, 1; St. Asaph, 
Denbighshire, 3 ; Conway, Carnarvonshire, 1 ; Hope End, Herefordshire, 1 ; 
Wells, Somersetshire, 1 ; Devizes, Wiltshire, 1 ; Zeals, Wiltshire, 2 ; Devon- 
shire, 2 ; Ragley House, Warwickshire, 3. 



5. 


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1. 1. 289- 




291. 



RHIN0L0PHUS 155 

3 i. St. Asaph, Denbighshire, Charles Oldham (c & p). 11. 1. 3. 5-7. 

Wales. 

1. Conway, Carnarvonshire. Sir W. Jardine (c & p). 60. 9. 17. 1. 

?. Great Grimsby, Lincoln- G. Barrett-Hamilton (p). 11. 1. 2. 98. 
shire, England. (Caton 
Haigh.) 

6. Bowdon, Cheshire. (T.A. G. Barrett-Hamilton (p). 11.1.2.99. 
Coward.) 

i al. Hope End, Herefordshire. N. C. Hewitt (c & p). 

6 al. Wells, Somerset. S. Lewis (o & p). 

1. Devizes, Wiltshire. J. E. Harting (c & p). 

6, 9 al. Zeals, Wiltshire. F. Norgate (c & p). 

2 ?. Devonshire. Oxley Grabham (c & p) 

3. Ragley House, Warwick- Tomes Collection, 
shire. 

RHINOLOPHUS EURYALE Basius. 

1853. Bhinolophus euryale Blasius, Wiegmann's Arehiv fur Naturgesch., 

1853, i, p. 49 (Milan, Italy). 
1857. Bhinolophus euryale Blasius, Saugethiere Deutschlands, p. 35. 
1878. Bhinolophus euryale Dobson, Catal. Chiropt. Brit. Mus., p. 116. 
1904. E[uryalus] toscanus Andersen and Matschie, Sitz.-Ber. Gesellsch. 

Naturforsch. Freunde, Berlin, p. 77 (Caverna di Parignano, Mt. 

Pisani, Italy). 
1904. E[uryalus'] atlanticus Andersen and Matschie, Sitz.-Ber. Gesellsch. 

Naturforsch. Freunde, Berlin, p. 77 (St. Paterne, Indre-et-Loire, 

France). 
1904. E\uryalus~\ cabrerse Andersen and Matschie, Sitz.-Ber. Gesellsch. 

Naturforsch. Freunde, Berlin, p. 78 (Alcala de Henares, Madrid, 

Spain). 
1910. Bhinolophus euryale, B. euryale atlanticus, B. euryale toscanus, 

B. euryale cabrerai Trouessart, Faune Mamm. d'Europe, pp. 5-7. 

Type locality. — Milan, Italy. 

Geographical distribution. — Southern Europe from Portugal 
to 1 Greece, north to Hungary and central France. 

Diagnosis. — Size medium, forearm, 44 ■ 6 to 49, condylobasal 
length of skull, 16-4 to 18, mandible, 12 to 13, upper tooth-row 
6 • 2 to 6 • 6 ; noseleaf with parallel-sided, bluntly rounded sella 
and high, sharply pointed connecting process, the lancet gradually 
narrowing to a bluntly cuneate tip ; fourth finger with first 
phalanx slightly more than one-third as long as second (ratio 
about 38) ; large upper premolar separated from canine by a 
narrow space occupied by the much reduced small premolar. 

" External characters. — Size intermediate between that of the 
Greater and Lesser Horseshoes. General outline of noseleaf 
(fig. 26 c) about as in Bhinolophus ferrum-equinum ; sella parallel 
sided, rounded off above, connecting process sharply linear- 
pointed, rising conspicuously above sella ; lancet with slight con- 
striction above middle, beyond which the tip narrows gradually to 
a bluntly cuneate point. Ear when laid forward extending about 
■5 mm. beyond extremity of muzzle, its tip less attenuate and less 
noticeably curved backward than in Bhinolophus ferrum-equinum 



156 



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and B. hipposideros ; antitragal lobe about balf as bigb as conch, 
its width about equal to its height. Wing peculiar in the 
shortening of the first phalanx of fourth finger to a little more 
than one-third that of second (average of 10 specimens from Gard, 
France: first phalanx, 6 - 6 mm.; second phalanx, 17 '4 mm.; 
ratio of first to second, 38 +)• Foot averaging slightly more 
than half as long as tibia. 

Colour. — While essentially as in BMnolophus ferrum-equinum 
and B. hipposideros, the colour usually differs slightly in the more 
evident contrast of the light area between ears and the more 
drabby general effect of underparts. Median region below 
occasionally rather paler than usual, sometimes nearly as in 
B. mehelyi. 

Skull. — In both size (greatest length about 19 mm.) and form 
the skull is somewhat intermediate between that of BMnolophus 
ferrum-equinum and B. hipposideros. It resembles or surpasses 
the latter in the breadth of brain-case relatively to narrow 
maxillary region, but more nearly agrees with the former in the, 
nearly parallel-sided mesopterygoid space, the anterior border of 
which is separated from posterior molars by well defined palatal 
emarginations. The mesopterygoid space is, however, shorter in 
proportion to its width than in B. ferrum-equinum. Floor of 
brain-case between cochleee less narrowed than in the preceding 
species. Nasal region slightly less inflated than in B. hipposideros, 
and inflated area relatively shorter, its posterior border rising 
abruptly above interorbital level. 

Teeth. — In all respects the teeth closely resemble those of 
BMnolophus ferrum-equinum (apart from their smaller size), except 
that the upper canine is relatively less robust, the anterior upper 
premolar is less reduced (its crown area about double that of 
upper incisor), and anterior lower premolar is less crowded 
between canine and posterior premolar. 

Measurements. — For cranial and external measurements see 
Tables, pp. 156 and 160. 

Specimens examined. — About 130, from the following localities : — 

Portugal : Cintra, 6. 

Spain: Villalba, Lugo, 1; Madrid, 2; Silos, Burgos, 1. 

France : St. Paterae, Indre-et-Loire, 3 (B.M. and U.S.N.M.) ; St. Genies, 
Gard, about 50 (Mottaz) ; Gapeau Eiver, Var, 12. 

Italy: Near Genoa, 33 (B.M., U.S.N.M., Genoa, and Mottaz); Monte 
Pisanino, 2 ; Siena, 3 (U.S.N.M.) ; Rome, 2 ; VeUetri, Borne, 5 (U.S.N.M.) ; 
Nieotera, Calabria, 1 ; Marsala, Sicily, 2. 

Sardinia : Mount Gennargentu, 3 (U.S.N.M.). 

Austria-Hungary : Ofener Mountains, 2 ; Orsova, 1. 

Dalmatia : Zara, 1. 

Greece : Missolungi, Acarnania, 2 (U.S.N.M.). 

Bemarks. — BMnolophus euryale is so readily distinguishable 
from all the other European members of the genus, except 
B. mehelyi, as to require no special comparisons. From B. mehelyi 
it is most easily distinguished by the form of the lancet and 



EHINOLOPHUS 



159 



antitragus, together with its rather smaller size and usually 
darker colour. With the material at hand I am unable to 
recognize the local forms of this species described by Andersen 
and Matschie, as the alleged differences appear to be within the 
range of normal individual variation. 

2 9. Cintra,500m. Portugal. O. Thomas (c & p). 98. 2. 2. 2-3. 

2 <?, 2 ? al. Cintra, 500 m. 0. Thomas (c & p). 98. 2. 2. 53-56. 

9 al. Villalba, Lugo, N.W. Dr. V. L. Seoane (p). 94. 1. 1. 1. 
Spain. 

i, 9 al. Madrid. A. Cabrera (p). 5. 2. 3. 1-2. 

1 al. St. Paterne, Indre-et- Boyal Army Medical 9. 1. 4. 9. 

Loire, France. College (p). 

9 al. St. Paterne, Indre-et- G. E. Dobson (p). 80. 12. 14. 3. 

Loire. 

9 al. Gapeau Eiver, Var. Dr. K. Jordan (c & p). 8. 3. 15. 2-10. 

3 <5, 9 al. Finalborgo, Liguria, Marquis G. Doria (p). 6.12.1.14-17. 

Italy. (A. Gagero.) 

2 al. Monte Pisanino, Liguria. Lord Lilford (p). 73.1.8.6. 

■2 6. Eome. (C. Coli.) G. Barrett-Hamilton 11. 1. 2. 40-41. 

(*)■ 
6 al. Nicotera, Calabria. Florence Museum (e). 85.7.6.1. 

2 6. Marsala, Sicily. (A. 0. Thomas (p). 6. 8. 4. 10-11. 

Robert.) 
S, 9 al. Ofener Mts., Budapest. Budapest Museum (e). 94. 7. 18. 2-3. 
9. Orsova, Hungary. Hon. W. Bothschild 7. 9. 16. 7. 

(*)■ 
9. Zara, Dalmatia, 50 m. Lord Lilford (p). 11. 1. 1. 128. 

[Kolombatovic.) 
1 al. S. Europe. Purchased (Parreys). 47. 5. 27. 44. 

RHINOLOPHUS MEHELYI Matschie. 

1901. Bhinolophus mehelyi Matschie, Sitz.-Ber. Gesellsch. Naturforsch. 

Freunde, Berlin, p. 225 (Bucharest, Boumania). 
1904. Bhinolophus carpetanus Cabrera, Mem. Soc. Espafi. Hist. Nat., n, 

p. 254 (Madrid, Spain). 
1910. Bhinolophus euryale mehelyi and B. carpetanus Trouessart, Faune 

Mamm. d'Europe, pp. 7-8. 

Type locality. — Bucharest, Roumania. 

Geographical distribution. — Roumania, southern France 
(Gard), Sardinia, central Spain. Details of distribution not 
known. 

Diagnosis. — Like Bhinolophus euryale but larger (forearm, 
48 • 6 to 51 • 4 ; upper tooth-row about 7 mm.); noseleaf with 
lancet abruptly narrowed to a linear tip ; ear with antitragal 
lobe relatively broad and low ; fourth finger with first phalanx 
decidedly more than one-third as long as second (ratio about 44) ; 
colour usually paler than in the related animal. 
<j External characters. — Slightly larger and more robust than 
Bhinolophus euryale, a difference especially noticeable in freshly 
killed individuals. Noseleaf as in B. euryale, except that the 
lancet is very abruptly narrowed above middle to a distinctly 
linear tip. Ear as in B. euryale but broader, the antitragal lobe 



160 



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162 CHIROPTEEA 

scarcely half as high as conch, its width slightly greater than 
height. Wing differing from that of the related animal in the 
less degree of shortening of the first phalanx of fourth finger as 
compared with second (average of ten specimens from Gard, 
Prance, first phalanx, 8'1 ; second phalanx, 18 - 0; ratio of first 
to second, 44 +)• Foot as in B. euryale. 

Colour. — Though not invariably distinguishable the colour is 
usually paler than that of Rhinolophus euryale, a difference 
especially noticeable in the region between ears, on sides of face 
and neck, on chin and throat, and along median portion of chest 
and belly, all of which are frequently a very pale almost whitish 
drab-grey.* 

Skull and teeth. — Except for its slightly greater average size 
(greatest length about 20 mm.) the skull agrees with that of 
Rhinolophus euryale. Teeth more robust than those of the 
related animal, but not peculiar in form. 

Measurements. — For external and cranial measurements see 
Tables, pp. 161, 163. 

Specimens examined. — About fifty-five, from the following localities : — 

Spain: Near Madrid, 1 (paratype of carpetanus). 

Pbance : Near St. Gerries, Gard, about 50, skins and in flesh (Mottaz) . 

Sardinia: Sassari, 3 (B.M. and U.S.N.M.). 

Boumania : Bucharest, 1 (U.S.N.M.) ; Dobrudscha, 1 (Mottaz). 

Remarks. — At first sight this species appears very similar to 
Rhinolophus euryale, together with which it occurs ; but its 
characters when once understood are readily appreciable. The 
ranges of the two animals will probably be found to be essentially 
coincident, though Rhinolophus mehelyi may prove to be more 
strictly confined to the Mediterranean region than the smaller 
form. 

9. Madrid. A. Cabrera (p.) 5. 2. 3. 1. 

(Paratype of R. carpetanus Cabrera.) 
2 <5 al. Sassari, Sardinia. Marquis G. Doria(p). 6.12.1.18-19. 

rhinolophus blasii Peters. 

1857. Rhinolophus clivosus Blasius, Siiugethiere Deutschlands, p. 33. Not 
of Riippell, 1824 (Italy, Sicily, Istria and Dalmatia). 

1866. Rhinolophus blasii Peters, Monatsber. k. Akad. Wissensch. Berlin, 
p. 17 (Renaming of clivosus Blasius). 

1878. Rhinolophus blasii Dobson, Catal. Chiropt. Brit. Mus., p. 117. 

1910. Rhinolophus blasiusi Trouessart, Paune Mamm. d'Europe, p. 9. 

Type locality. — South-eastern Europe. 

Geographical distribution. — Eastern portion of the Mediter- 
ranean region : Cyprus, Greece, Italy? 

Diagnosis. — Size essentially as in Rhinolophus euryale ; nose- 

* When seen by candle-light flying in caverns these bats are said to 
appear entirely white. 



KHINOLOPHUS 



163 



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R. mehelyi. 

Roumania: Dobrudscha . 

Bucharest 
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France: near Nimes, Gard 

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Spain : near Madrid 

R. blasii. 
Cyprus .... 

Greece: Nauplia 



M 2 



164 CHIROPTERA 

leaf with cuneate sella and very high, sharply pointed connecting 
process ; fourth finger with first phalanx more than half as long 
as second ; no marked contrast between crown areas of anterior 
and posterior lower premolars, a character unique among the 
European members of the genus! 

Colour. — The only skin of this species which I have examined 
is in bad condition. It indicates that the colour is not essentially 
different from that of Rhinolophus euryale. 

Skull. — In general the skull resembles that of Rhinolophus 
euryale, with which it agrees in size and in the form of the nasal 
swellings as well as in that of mesopterygoid fossa and posterior 
portion of palate. Constriction at front of interparietal more 
pronounced than in any of the other European species, noticeably 
marking off the occipital region from rest of brain-case. 

Teeth. — Incisors, canines and molars as in Rhinolophus euryale. 
Small upper premolar slightly less reduced, perfectly in the 
tooth-row, but showing no tendency to develop a cusp. Large 
upper premolar with anterior and posterior margins of crown 
essentially parallel, the posterior border nearly straight. Lower 
premolars differing from those of all the other European members 
of the genus in the approximately equal crown areas of the two 
larger teeth, the anterior subterete, the posterior with trapezi- 
form section ; shaft of posterior tooth with diameter in axis of 
tooth-row much less than transverse diameter, the cusp when 
viewed from the side appearing to rise from middle of crown with 
noticeable flat area before and behind it. 

Measurements. — Average and extremes of four females from 
Cyprus: head and body, 48-4 (44-51); tail, 24-7 (24-25); 
tibia, 19-3 (19-20); foot, 9-7 (9-4-10); forearm, 45-3 (44-6- 
47) ; thumb, 7 '5 (7-8) ; third finger, 70-3 (69-72) ; fifth finger, 
57'7 (56-60); ear from meatus, 19 -7 (19-20) ; ear from c'rown, 
15-5 (15-16-4); width of ear, 14-7 (14-15). Adult from 
Nauplia, Greece : tibia, 18 • 8 ; foot, 9 ■ 6 ; forearm, 44 • 6 ; thumb, 
8 ; third finger, 69 ; fifth finger, 57. For cranial measurements 
see Table, p. 163. 

Specimens examined. — Five, from the following localities : — 
Cyprus : No exact locality, 4. 
Gbeecb : Nauplia, 1 (U.S.N.M.). 

lieimirkt. — This species is so readily distinguished from the 
other European members of the genus by the peculiarities of its 
noseleaf and lower premolar as to require no special comparisons. 
Its range appears to be strictly confined to the eastern portion 
of the Mediterranean region, not extending west of Italy. 



VESPEKTILIONINJE 165 

Family VESPERTILIONIDSE. 

1821. Vespertilionidm Gray, London lied. Repos., xv, p. 299, April 1, 1821 
(part). 

1857. Vespertiliones Blasius, Saugethiere Deutschlands, p. 37. 

1878. Vespertilionidm Dobson, Catal. Chiropt. Brit. Mus., p. 167 (except 

the genera NataVus and Thyroptera). 
1907. Vespertilionidse Miller, Families and Genera of Bats, p. 195, June 29, 

1907. 

Geographical distribution. — Eastern and western hemispheres 
to the limits of tree growth ; in the Atlantic to the Azores, and 
in the Pacific to the Galapagos and Hawaiian Islands from 
America, and to Australia, New Zealand and Samoa from Asia. 

Characters. — Ear with tragus ; muzzle without distinct leaf- 
like outgrowths ; skull with premaxillaries represented by nasal 
branches only, the two bones very early fused with surrounding 
parts ; median length of palate greater than least distance between 
tooth-rows ; auditory bulla not emarginated on inner side; shoulder 
girdle normal, without fusion of its elements ; secondary 
articulation of humerus with scapula better developed than in 
the Bhinolophidse ; fibula very slender, not adding appreciably to 
strength of leg ; foot normal, the toes slender ; tail not project- 
ing conspicuously beyond membrane. 

Remarks. — This family is the most widely distributed group 
of bats as well as one of the richest in genera and species. Forty- 
one genera are at present known, eight of which occur in 
Europe.* 

Sub-Family VESPERTILIONIJST^E. 

1878. Vespertiliones Dobson, Catal. Chiropt. Brit. Mus., p. 168 (except 

genera Kerivoula and Harpiocephalus). 
1907. Vespertilioninse Miller, Families and Genera of Bats, p. 197, June 29, 

1907. 

Geographical distribution. — Same as that of the family 
Vespertilionidse. 

Characters. — Sternum slender, its entire length considerably 
more than twice greatest width of presternum ; median lobe 
very much smaller than body of presternum ; six ribs connected 
with sternum ; seventh cervical vertebra not fused with first 

* The American Nycteris cinerea has been recorded (under the name 
Vespertilio pruinosus) from South Bonaldshay, Orkney Islands, but the 
occurrence seems open to question (see Wolley, The Zoologist, vn, p. 2343, 
1849; viii, pp. 2695-96, 2813-14, 1850; Barrett-Hamilton, Hist. Brit, 
llamm., i, pp. 222-224, March, 1911). 

A specimen of another North American member of this genus 
IN. borealis), bearing the label: "Villevgque (Maine & Loire), 8. 1. 89," is 
figured by Trouessart in Bull. Soc. Zool. de France, xxx, p. 152, 1905. 
This is copied, with change of locality from Maryland to France, from a 
figure published in Bull. U.S. Nat. Mus., No. 39, pt. N, 1899 and 1901. 
Though intended merely as a guide in preparing specimens, Dr. Trouessart's 
publication might be misinterpreted as a French record of the species. 



166 CHIEOPTEEA 

dorsal ; scapula with coracoid curved outward ; nostrils simple ; 
lower incisors in all known genera, 3-3. 

Remarks. — The sub-family Vespertilioninse contains all but 
eight of the known genera of Vespertilionidse, and all but one, 
Miniopterus, of the eight found in Europe. It is the central, 
least specialized portion of the family. 

KEY TO THE EUROPEAN GENERA OP VESPEBTILION1D&]. 

Cheek teeth jtjj Myotis, p. 166. 

Cheek teeth less than ||. 
Upper premolars 1-1. 
Rostrum noticeahly concave on each side of middle 
line ; nares extending about halfway to inter- 
orbital constriction ; palatal emargination 

broader than deep Vespertilio, p. 238. 

Rostrum evenly convex laterally; nares not ex- 
tending halfway to interorbital constriction ; 

palatal emargination deeper than broad Eptesicus, p. 224. 

Upper premolars 2-2. 

Lower premolars 3-3. 

Auditory bulla large, its greatest diameter 

more than twice width of basioccipital ; 

ear much longer than head; second 

phalanx of third finger shorter than first Plecotus, p. 256. 

Auditory bulla small, its greatest diameter 

about equal to width of basioccipital ; ear 

, shorter than head; second phalanx of 

third finger nearly three times as long as 

first Miniopterus, p. 268. 

Lower premolars 2-2. 

Fifth finger shortened, its length only a little 
more than that of metacarpal of fourth 

or third Nyctalus, p. 242. 

Fifth finger normal, its length greater than 
that of metacarpal and first phalanx of 
fourth or third. 
Upper surface of rostrum convex; ears 

separate Pipistrellus, p. 202. 

Upper surface of rostrum concave ; ears 

joined Barbastella, p. 263. 

Genus MYOTIS Kaup. 

1829. Myotis Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, I, 
p. 106 (myotis). 

1829. Nystactes Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, I, 

p. 108 (bechsteimi). 

1830. Leuconoe Boie, Isis, p. 256 (daubentonii). 

1839. Vespertilio Keyserling and Blasius, Wiegmann's Archiv fiir Natur- 
gesch., 1839, I, p. 306 (Not Vespertilio Linnaeus, 1758). 

1841. Selysius Bonaparte, Iconogr. Faun. Ital. i., Introd. alia Classe 
Mamm., p. 3 (mystacinus). 

1841. Capaccinius Bonaparte, Iconogr., Fauna Ital., i, Indice Distrib., 

p. 1 (capaccinii). 

1842. Trilatitus Gray, Ann. and Mag. Nat. Hist., x, p. 258, December, 

1842 (hasseltU, macellus = adversus and blepotis). 



MYOTIS 167 

1849. Tralatitus Gervais, Diet. Univ. d'Hist. Nat., xiii, p. 213 (Modifica- 
tion of Trilatitus). 

1856. Brachyotus Kolenati, Allgem. deutsch. Naturhist. Zeitung, Dresden, 
neue Folge, n, p. 131 (mystacinus, daubentonii, and dasyeneme). 
Not Brachyotus Gould, 1837. 

1856. Isotus Kolenati, Allgem. deutsch. Naturhist. Zeitung, Dresden, neue 

Folge, ii, p. 131 (nattereri and emarginatus). 

1857. Vespertilio Blasius, Saugethiere Deutschlands, p. 78. Not Vespertilio 

Linnaeus, 1758. 

1866. Tralatitius Gray, Ann. and Mag. Nat. Hist., 3rd ser., xvn, p. 90, 

February 1866 (Modification of Trilatitus). 

1867. Ptemopterus Peters, Monatsber. k. preuss. Akad. Wissensch. Berlin, 

p. 706 (sub-genus of Vespertilio = Myotis, type lobipes = muricota) . 

1870. Exochurus Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien, 
Math.-Naturwiss. Classe, lxii, p. 75 (macrodactylus, horsfleldii = 
adversus and macrotarsus). 

1870. Aeorestes Pitzinger, Sitzungsber. kais. Akad. Wissensch. Wien, 
Math.-Naturwiss. Classe, lxii, p. 427 (villosissimus, albescens, and 
nigricans). 

1870. Comastes Pitzinger, Sitzungsber. kais. Akad. Wissensch. Wien, Math.- 
Naturwiss. Classe, lxii, p. 565 (capaccinii, megapodius, dasyeneme, 
and limnophilus). 

1878. Vespertilio Dobson, Catal. Chiropt. Brit. Mus., p. 284. Not Vespertilio 
Linnaeus, 1758. 

1897. Myotis Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 382, 
October, 1897. 

1899. Euvespertilio Acloque, Paune de Prance, Mammiferes, p. 38 (emargin- 
atus, mystacinus, murinus = myotis, nattereri, and bechsteinii). 

1907. Myotis Miller, Families and Genera of Bats, p. 201, June 29, 1907. 

Type species. — Vespertilio myotis Borkhausen. 

Geographical distribution. — Entire mainland of Eastern and 
Western hemispheres to limits of tree growth ; also the Malay 
Archipelago, New Guinea, Australia and Samoa, and in America 
the Lesser Antilles. 

Characters. — Dental formula : i g, c J--1, pm jg, m g = 38. 
General form slender and delicate, even in such large species as 
M. myotis, the skull slender and lightly built ; muzzle narrow ; 
ear narrow and rather long, without special peculiarities of form, 
the tragus at least half as high as conch, straight or slightly 
curved, tapering gradually to a narrow or acute point. 

Remarks. — The genus Myotis is the most widely distributed 
of the genera of bats. It is also probably the richest in species, 
though these are at present so imperfectly known that no estimate 
of their number can be made. Nine occur in Europe. These 
present considerable differences in size, ranging from nearly the 
smallest to nearly the largest members of the group ; they also 
differ considerably among themselves in certain details of 
structure, notably in the relative size of the hind foot ; but all 
are recognizable, apart from their dental formula, by a certain 
slenderness and delicacy of form, especially noticeable in the 
muzzle, ear, tragus and skull. 



168 CHIHOPTEKA 

KEY TO THE EUBOPEAN SPECIES OP MYOTIS. 

Size large (forearm 53 to 64 mm., oondylobasal 
length of skull 18 • 6 to 23 ■ 6 mm., upper tooth- 
row 8 • 2 to 10 • 6 mm.) ; middle upper premolar 
normally crowded inward from axis of tooth- 
row; third lower molar with second triangle 
much smaller than first and noticeably different 
from it in form. 
Oondylobasal length of skull 22 to 23 '6 mm.; 
mandible 17 '8 to 19 mm. ; maxillary tooth- 
row 9-8 to 10"6 mm. (Central and southern 

Europe) M. myotis, p. 192. 

Oondylobasal length of skull 18 • 6 to 21-4 mm. ; 
mandible 15 • 2 to 17 ■ 2 mm. ; maxillary tooth - 

row 8-2 to 9'4 mm. (Mediterranean region) M. oxygnathus, p. 199. 
Size small or medium (forearm 34 to 47 mm., oondy- 
lobasal length of skull 12 ■ 4 to 16 ■ 8 mm. , upper 
tooth -row 5 to 7 mm.) ; middle upper premolar 
not crowded inward from axis of tooth-row ; 
third lower molar with second triangle nearly 
as large as first and essentially like it in 
form. 
Foot relatively large, obviously more than half 
as long as tibia ; calcar about twice as long 
as free border of interfemoral membrane ; 
skull broad, the width of brain-oase more 
than half greatest length; crown area of 
molars relatively small ; upper molars with 
evident protoconule. 
Forearm about 47 mm. ; oondylobasal length 
of skull about 16 mm. (Central and 

southern Europe) M. dasycneme, p. 189. 

Forearm less than 45 mm. ; oondylobasal length 
of skull never more than 15 mm. 
Tibia and adjacent membrane densely furred; 
forearm about 42 mm. ; oondylobasal 
length of skull 14-0 to 14-8 mm. (Medi- 
terranean region) 71/. capaccinii, p. 187. 

Tibia and adjacent membrane not furred ; 
forearm about 35 mm. ; oondylobasal 
length of skull 13-2 to 13 -8 mm. 

(Distribution general) M. daubentonii, p. 184. 

Foot relatively small, about half as long as tibia ; 
calcar about as long as free border of inter- 
femoral membrane ; skull narrow, the width 
of brain-case less than half greatest length ; 
crown area of molars relatively large ; upper 
molars without protoconule. 
Ear not specially elongated, extending slightly 
beyond nostril when laid forward. 
Forearm about 40 mm. ; oondylobasal length 
of skull about 15 mm. ; posterior border 
of ear conch with deep, almost angular 

emargination slightly above middle M. emarginatus, p. 177. 

Forearm about 34 mm. ; oondylobasal length 
of skull 12-6 to 13 • 2 mm.; posterior 
border of ear conch with shallow, in- 
conspicuous emargination M. mystacinus, p. 169. 

Ear elongated, extending conspicuously beyond 
nostril when laid forward. 



MYOTIS 169 

Combined length of tibia and foot less than 

25 mm. ; oondylobasal length of skull 

14 • to 14 • 6 mm. ; ear narrow, its width 

about 10 mm. ; tragus conspicuously 

more than half as high as conch 21. nattercri, p. 174. 

Combined length of tibia and foot about 

30 mm. ; oondylobasal length of skull 

16 to 17 mm. ; ear broad, its width about 

14 mm. ; tragus scarcely half as high as 

conch 31. bechsteinii, p. 179. 

MYOTIS MYSTACINUS Kuhl. 

1819. Vespertilio mystacinus Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv 

(= Neue Ann., i), pt. 2, p. 202. 
1821. Vespertilio collaris Schinz, Das Thierreich von Cuvier, i, p. 177 

(Mt. Blanc, Haute-Savoie, Prance). 
1834. Vespertilio humeralis Baillon, Mem. Soc. Royale d'Emulation 

d'Abbeville, 1833, p. 50 (Abbeville, Somme, Prance). 
1837. Vespertilio schinsii Brehm, Ornis, Heft in, p. 27 (Renthendorf, 

Thiiringen, Germany). 
1843. V[espertilio] schranhii Wagner, Wiegmann's Archiv fur Naturgesch., 

tx, Bd. n, p. 25 (Munich, Germany ? See Pitzinger, Sitzungsber. 

kais, Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1, 

p. 219, 1871). 
1857. Vespertilio mystacinus Blasius, Saugethiere Deutschlands, p. 96. 
1863. [Brachyotus mystacinus] var. nigricans Koch, Jahrb. des Vereins fur 

Naturkunde im Herzogthum Nassau, xvin, p. 444 (Wiesbaden, 

Nassau, Germany). 
1863. [Brachyotus mystacinus] var. rufofuscus Koch, Jahrb. des Vereins 

fur Naturkunde im Herzogthum Nassau, xvin, p. 444 (Wiesbaden, 

Nassau, Germany). 
1863. [Brachyotus mystacinus] var. aureus Koch, Jahrb. des Vereins fur 

Naturkunde im Herzogthum Nassau, xvin, p. 445 (Breisgau, 

Germany). 
1869. [Vespertilio mystacinus] var. nigricans Patio, Faune Vert. Suisse, 

i, p. 92 (Switzerland). Not of Koch, 1863. 
1869. [Vespertilio] lugubris Patio, Paune Vert. Suisse, i, p. 93 (Alternative 

for nigricans Patio). 
1871. Vespertilio mystacinus, nigro-fuscus Pitzinger, Sitzungsber. kais. 

Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1, 

p. 217 (Renaming of V. schinzii Brehm). 
1878. Vespertilio mystacinus Dobson, Catal. Chiropt. Brit. Mus., p. 314. 
1900. Myotis mystacinus Mehely, Monogr. Chiropt. Hungariae, p. 200. 
1910. Myotis mystacinus Trouessart, Paune Mamm. d'Europe, p. 33. 

Geographical distribution. — Entire Continent of Europe north 
to about the limits of tree growth ; west to Ireland ; east into 
Asia. 

Diagnosis. — Smallest species of European Myotis (forearm 
about 34 mm., longest finger about 60 mm., oondylobasal length 
of skull 12-6 to 13 "2 mm.) ; ear moderately long, extending 1 to 
2 mm. beyond tip of muzzle when laid forward, its posterior 
border with shallow inconspicuous emargination ; foot about half 
as long as tibia ; wing membrane extending to base of outer 
toe ; last caudal vertebra free. 



170 



CHIROPTERA 



External characters. — General form slender and delicate, the 
legs and tail rather long, the membranes thin and semi-trans- 
parent. Muzzle with rather noticeable glandular swellings. 
Ear extending slightly beyond nostril when laid forward, its 
general form rather slender, the tip narrowly rounded off, the 
posterior border with shallow inconspicuous concavity extending 
from just below tip to near middle of conch ; inner surface of 
conch without well defined transverse striations. Antitragus 
small (length about 2 mm.) but well defined Tragus a little 
more than half as high as conch, its width slightly above level 
of anterior base contained about 2J times in length of anterior 
border, the anterior border straight, the posterior border convex 
below ; from widest region it narrows rather rapidly upward to 
a rather blunt point ; posterior basal lobe well defined. Wing 
rather narrow, with no special peculiarity of form, the third, 
fourth and fifth metacarpals sub-equal, their distal extremities 
falling short of elbow by about 3 mm. ; membrane inserted at base 
of outer toe. Foot about half as long as tibia ; calcar slender, 
with barely indicated rudiment of keel and terminal lobe, its 
length slightly greater than that of free border of interfemoral 
membrane. Tail about as long as head and body, and twice as 
long as tibia, its terminal vertebra free from membrane. 

Fur and colour. — The fur is soft and loose, the hairs on middle 
of back about 10 mm. in length, those of underparts a little more 
than half as long ; it is closely confined to body, extending on 
wings to extreme base of membrane only, slightly farther below 
than above, and on interfemoral membrane over basal fifth or 
fourth of both surfaces ; free border of ufopatagium not fringed. 
Colour of upper parts a clear light brown resembling the wood- 
brown of Ridgway, but usually more yellow and always with a 
distinct metallic gloss, the basal portion of the hairs blackish- 
slate, this colour sometimes appearing at the surface and 
producing a general darkening effect ; underparts paler and more 
buffy, usually not forming any decided contrast with back, but 
occasionally almost whitish, especially on 
6 "-^— ^ chest ; muzzle and cheeks dusky ; ears and 

V, W s Sirf« membrane blackish. 

£5Ud<oa^ Skull. — The skull is slender and lightly 

built, the breadth of brain-case conspicuously 
greater than that of rostrum and slightly 
but appreciably less than half greatest length. 
Dorsal profile rising abruptly above low ros- 
trum in interorbital region and forming a 
strong convexity over anterior portion of 
Fig. 29. brain-case ; occipital region distinctly pro- 

mote gw duced backward and rising slightly but evi- 
dently above level of anterior portion of 
brain-case, from which it is marked off by a shallow but notice- 
able transverse constriction following anterior margin of inter- 




171 



parietal. Ventral profile rising slightly but evidently in region 
of floor of brain-case. Greatest depth of brain-case about three- 
quarters mastoid width ; sagittal and lambdoid crests slightly 
indicated in fully adult individuals. Interorbital region obscurely 
short hour-glass shaped. Anteorbital foramen small, its posterior 
border over posterior root of large premolar. Posterior palatal 
region rather narrow, its width immediately behind molars about 
equal to its greatest length, the median spine broad and short ; 
mesopterygoid fossa slightly wider than long, the hamulars turned 
inward. Auditory bulla moderate, its greatest diameter nearly 
equal to distance between inner margins of bullse. 

Teeth. — Upper incisors sub-equal, about half as high as canine, 
each pair in contact or nearly so at base but diverging at tips, the 
cingulum of inner tooth horizontal, that of outer tooth oblique ; 
crown of outer incisor squarish in cross section, that of inner 
somewhat elongated in axis of tooth-row ; near point of contact 
each shaft bears a secondary cusp, this usually though not always 
better developed in outer than in inner tooth ; distance between 
canine and outer incisor about equal to diameter of incisor, that 
between pairs a little more than 
twice as great. Lower incisors 
slightly but evidently imbricated, 
forming a continuous, broadly 
V-shaped row between 
canines ; crown of i 3 sub- 
terete, its area more than 
half that of canine and 
about equal to that of pm v 
its blunt main cusp, the 
highest in the incisor series, 
situated at outer side, the 
three smaller cusps (third 
obsolete) on inner margin ; 
i x and i 2 sub-equal, slightly 
lower than i 3 , their crowns 
compressed, longer than 
high, trifid, that of second 
wider posteriorly than an- 
teriorly, and usually with 
a minute postero-internal 
cusp. Upper canine rela- 
tively large, slightly higher than main cusp of large premolar, 
its shaft somewhat triangular in cross section, with broad 
postero-internal concavity and anterior and postero-external 
groove ; posterior cutting edge well developed ; cingulum com- 
plete but not forming evident secondary cusps. Lower canine 
lower and less acutely pointed than upper, its tip about on level 
with highest cusps of molars ; cingulum usually forming a slight 
secondary cusp anteriorly. First and second upper premolars 





FIS. 80. 
Myotis mystacinus. Teeth X 10. 



CHIROPTERA 



alike in form, crowded between canine and large premolar but 
perfectly in the tooth-row, the first about the same size as the 
upper incisors, the second with slightly more than half the 
height and crown area of first, both with fully developed 
cingulum and subterete, conical cusp. Large upper premolar 
with crown area only a little less than that of first molar, the 
main cusp large and with strongly trenchant posterior cutting 
edge, its height about equal to that of largest molar cusps ; 
anterior and posterior borders of crown slightly concave, inner 
border narrowly rounded, usually with a distinct though small 
cusp anteriorly, sometimes with another barely indicated pos- 
teriorly ; a slight though evident concave crushing surface 
between cingulum and inner base of main cusp. First and 
second lower premolars essentially similar to the corresponding 
upper teeth but slightly less reduced in size and with somewhat 
higher cusps ; third lower premolar similar to first and second 
but with rectangular crown nearly twice as large as that of second 
tooth, and main cusp as high as protoconid of first molar ; a 
small cingulum cusp usually present at antero-internal angle. 
Upper molars rather large relatively to size of skull, the crown 
area of second tooth slightly greater than that of first ; anterior 
and posterior borders slightly concave, inner border narrowly 
rounded, especially in m 2 , the antero-internal and postero-internal 
outlines often flattened or even a little concave ; protocone large, 
its base occupying entire inner border of tooth, its cusp a little 
in front of middle, its anterior commissure simple, extending 
uninterruptedly outward to parastyle, its posterior commissure 
terminating in a thickened rib-like rudiment of a hypocone, 
between the outer base of which and inner base of metacone lies 
a deep furrow ; metacone larger than paracone ; styles and outer 
commissures well developed ; third upper molar with crown area 
equal to about three-quarters that of m l , the protocone as in the 
other teeth but smaller, paracone larger than metacone, the first 
outer commissure longer than in m 1 and m 2 , the second and third 
about as in the other teeth but set at a different angle, the fourth, 
together with metastyle absent. First and second lower molars 
with second V slightly larger than first in cross section, this 
condition reversed in m 3 ; protoconid decidedly higher than 
hypoconid in all three teeth ; inner cusps of about equal height 
throughout (slightly more than half as high as protoconid) ; a 
distinct cingulum cusp behind entoconid. 

Measurements. — Adult female from Skane, Sweden : tibia, 
1 5 ; foot, 7 ' 6 ; forearm, 32 ; thumb, 6 • 2 ; third finger, 49 ; fifth 
finger, 38; ear from meatus, 12; width of ear, 8. Adult male from 
Madrano, Tyrol : head and body, 38; tail, 38; tibia, 16 ; foot, 
8 ; forearm, 35 ; thumb, 6 • 2 ; third finger, 56 ; fifth finger, 47 ; 
ear from meatus, 13 ; width of ear, 9. Adult female from the 
Carpathian Mountains : head and body, 44 ; tail, 40; tibia, 15' 4; 
foot, 8 ; forearm, 34 ; thumb, 7 ; third finger, 54 ; fifth finger, 



MYOTIS 



173 



45 ; ear from meatus, 14 • 2 ; width of ear, 9. Forearm, in other' 
specimens : Waremme, Liege, Belgium, 33 and 33 ; Strass, near 



Burgheim, Bavaria, 31 • 6 
Switzerland, 33 and 34*4. 
p. 182. 



Dresden, 33 • 2 ; Grotte de Vallorbe, 
For cranial measurements see Table, 



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

England : Newby Bridge, Lake Windermere, Cumberland, 1 ; Aberia 
Merionethshire, 2; Colwyn, Denbighshire, 1; Cheadle, Staffordshire, 1 
Manchester, Lancashire, 1 ; Ragley House, 1 ; Welford, Herefordshire, 2 
Macclesfield, Cheshire, 1 ; Pewsey, Wiltshire, 1 ; Dover, Kent, 2 ; Hastings, 
Sussex, 1; Bath, Somerset, 1. 

Sweden : Skullno, 1 ; Skane, 1 (U.S.N.M.). 

Belgium : Waremme, Liege, 2 (U.S.N.M.). 

Germany : Moritzburg, Saxony, 1 (U.S.N.M.) ; Dresden, 1 ; Strass, 
near Burgheim, Bavaria, 1 ; Bavaria, no exact locality, 2 (U.S.N.M.). 

Austria-Hungary : Haida, Arva, Bohemia, 1 ; Carpathian Mts., 1 
(U.S.N.M.) ; Csall6k6z-Somorja, Pressburg, 2 ; Tatra Mts., Hungary, 1. 

Switzerland : Geneva, 1 (Mottaz) ; Grotte de Vallorbe, Vaud, 2 
(Mottaz) ; Stein, Appenzell, 1 (U.S.N.M.) ; St. Gallen, 1 (U.S.N.M.) ; no 
exact locality, 1 (Geneva: type of lugubris Patio). 



Bemarkts. — Myotis mystacinus, the smallest European member 
of the genus, is recognizable by its small size and relatively short 
foot in combination with the insertion of wing membrane at base 
of outer toe. From the small species of Pipistrellus it is at once 
distinguished by the narrow muzzle and slender tragus. 



2 ?. Aberia, Merionethshire, 

Wales. 
9 st. Colwyn, Denbighshire. 

1 al. Newby Bridge, Cumber- 

land, England. 

9. Cheadle, Staffordshire. 

6. Manchester, Lancashire. 

<J. Ragley House, Warwick- 
shire. 

2 9. Welford, Herefordshire. 
6. P e w s e v, Wiltshire. 

(P. S.'Hembly.) 
9. Dover, Kent. (B. Hesse.) 

1. Hastings, Sussex. 
6 st. Bath, Somerset. 
1. Skullno, Sweden. 

9. Haida, Bohemia. (Wol- 

terstorff.) 
1. Csall6kbz - Somorja, 

Pressburg, Hungary. 
1 al. Tatra Mountains, 

Hungary. 



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

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

T. Paul (c & p). 

E. Blagg (c & p). 
C. Oldham (p). 
Tomes Collection. 

Tomes Collection. 
C. H. B. Grant (p). 

C. H. B. Grant (p). 
Miss I. Roods (c & p). 
G. Dalgleish (c & p). 
Stockholm Museum 

(»■) 
Lord Lilford (p). 

Budapest Museum 

(e) 
Dr. R. Collett (p). 



11. 1. 3. 13-14. 

6. 2. 4. 1. 

94. 9. 3. 1. 

11. 1. 3. 15. 
11. 1. 3. 10. 

7. 1. 1. 498. 

7.1.1.496-497. 
11. 1. 3. 11. 

11. 1. 3. 12. 
49. 1. 16. 1. 
4. 10. 13. 3. 
46. 1. 2. 22. 

11. 1. 1. 6. 

94. 3. 1. 20-21. 

91. 1. 21. 1. 



174 CHIROPTERA 

MYOTIS NATTERERI Kuhl. 

1818. Vespertilio nattereri Kuhl, Ann. Wetterau. Gesellsch. Naturk., it 

(= Neue Ann., i), pt. 1, p. 33. 
1857. Vespertilio nattereri Blasius, Saugethiere Deutschlands, p. 88. 
1863. [Isotus nattereri] var. typus Koch, Jahrb. des Vereins fiir Natur- 

kunde im Herzogthum Nassau, xviii, p. 430 (Wiesbaden, Nassau , 

Germany). 
1863. [Isotus nattereri] var. spelseus Koch, Jahrb. des Vereins fur Natur- 

kunde im Herzogthum Nassau, xviii, p. 430 (Erdbach, Nassau, 

Germany). 
1878. Vespertilio nattereri Dobson, Cat'al. Chiropt. Brit. Mus., p. 88. 
1900. Myotis nattereri MShely, Monogr. Chiropt. Hungariae, p. 179. 
1904. Myotis escalerai Cabrera, Mem. Soo. Espafi. Hist. Nat., n, p. 279 

(Bellver, Lerida, Spain). 
1910. Myotis nattereri and M. escalerai Trouessart, Paune Mamrc. d'Europe, 

pp. 29-30. 

Type locality. — Hanau, Hessen, Germany. 

Geographical distribution. — Central and southern Europe, 
west to Ireland, north to southern Sweden. 

Diagnosis. — Size medium among the European species (forearm 
about 38 mm., longest finger about 70 mm., condylobasal length 
of skull, 14" to 14 - 6 mm.); ear elongated, extending about 
5 mm. beyond tip of muzzle when laid forward, the conch narrow 
(about 10 mm.), obscurely emarginate on upper half of posterior 
border, the tragus relatively longer than in any other European 
species, its height distinctly more than half that of conch ; foot 
about half as long as tibia ; wing membrane extending to base of 
outer toe ; edge of interfemoral membrane fringed. 

External characters. — General form essentially as in Myotis 
mystacinus, but differing in the following particulars : ear much 
longer and relatively narrower, extending conspicuously beyond 
nostril when laid forward, its extremity more broadly rounded 
off (owing to more uniform convexity of anterior border), and 
concavity on posterior border even less evident ; antitragus 
about as large as in M. mystacinus but less well defined ; tragus 
relatively longer and more attenuate than in any other European 
bat, its height conspicuously more than half that of conch, its 
greatest width contained about 3£ times in length of anterior 
border, its terminal third almost linear, sometimes faintly 
recurved, the basal lobe usually small and ill defined ; insertion 
of wing membrane as in M. mystacinus, but membrane at first 
very narrow, so that in some specimens, particularly those that 
have been hardened in strong alcohol, the point of insertion 
appears to be on side of metatarsus ; * tail rather shorter than 
head and body, only the cartilaginous extreme tip free ; calcar 
better defined than in M. mystacinus, about as long as the 
distinctly fringed free border of interfemoral membrane. 

* Such specimens seem to have formed the basis of the Myotis escalerai 
of Cabrera. 



MYOTIS 175 

Fur and colour. — Quality and distribution of fur essentially as 
in M. mystacinus, but free border of interfemoral membrane 
distinctly though not very densely fringed with hairs about 

I mm. long. Colour of upper parts a lighter and less yellowish 
brown than in M. mystacinus, the exact shade intermediate 
between the wood-brown and broccoli-brown of Ridgway, the 
longer hairs with faintly darker tips visible in certain lights ; 
underparts rather sharply contrasted whitish buffy grey ; a well 
defined line of demarcation extending from shoulder to base of 
ear ; basal portion of hairs clove-brown ; muzzle and cheeks dusky, 
but not so dark as in M. mystacinus ; ears and membranes dark 
brown. 

Skull. — In all its dimensions the skull is appreciably larger 
than that of Myotis mystacinus. In general form it is slightly 
less slender. Forehead rising more abruptly but occipital region 
not higher than main portion of brain-case, so that dorsal profile 
shows a stronger concavity in interorbital region than that of 
M. mystacinus, while over greater extent of brain-case it is 
essentially flat, or slightly falling away posteriorly, instead of 
rising by two well defined curves to lambdal region. Posterior 
palatal region about as in M. mystacinus, except that mesopterygoid 
space extends further forward, and median spine is better 
developed. Auditory bulla relatively smaller than in M. mysta- 
cinus, its greatest diameter decidedly less than distance between 
bullae. 

Teeth. — In general the teeth resemble those of Myotis 
mystacinus apart from their greater size. They differ, however, 
in certain details of form : crown area of outer upper incisor 
appreciably greater than that of inner tooth ; angle at front of 
lower incisor row wider, though evident ; upper canine relatively 
weaker, its posterior cutting edge less developed, the cross section 
of its shaft half-terete owing to the obsolescence of postero- 
external and anterior longitudinal furrow ; lower canine with 
length of base much greater in proportion to height of shaft, and 
cingulum more oblique ; second upper premolar nearly equal to 
first in cross section ; crown area of large premolar not so great 
relatively to that of first molar, the cusps on inner border 
obsolete or absent ; lower premolars with less slender crowns, 
the cingulum of the third forming a very low, sometimes obsolete 
antero-internal cusp ; molars, especially m 3 , with narrower 
crowns. 

Measurements. — External measurements of adult male from 
Colpin, Brandenburg, Germany, and adult male from Magdeburg, 
Germany: head and body, 50, 44; tail, 41, 40; tibia, 17, 16; 
foot, 8- -2, 9-0; forearm, 39-2, 40-2; thumb, 7-0, 6-2; third 
finger, 71,71; fifth finger, 56, 56 ; ear from meatus, 18, 18 ; tragus, 

I I • 0, 1 1 • 2. Two adult females from Spain (No. 94. 1 . 1. 8, Seville, 
and No. 8. 7. 23. 4 from Bellver, Lerida, paratype of M. escalerai) : 
head and body, 46, 46 ; tail, 41, 43 ; tibia, 15, 17 ; foot, 7 ■ 6, 9 • 2 ; 



176 



CHIROPTERA 



forearm, 36-4, 40-6; thumb, 7 -2, 8-4; third finger, 69, 72; 
fifth finger, 52, ,16; ear from meatus, 16, 16-6; tragus, 10' 2, 
10±. For cranial measurements see Table, pp. 182-183. 

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

Scotland : Inverary, Argyllshire, 1. 

England : Harlech, Merionethshire, 3 ; Arrow Church, 6 ; Alcester, 
Warwickshire, 8 ; Lilford Hall, Northamptonshire, 5 ; Henley-on-Thames, 
Oxfordshire, 3 (B.M. and U.S.N.M.) ; Bradfield, Berkshire, 1 ; Queen 
Camel, Somersetshire, 1 ; Devonshire, 1 ; no exact locality, 3. 

Ireland : Co. Longford, 1 ; Woodpark, Co. Galway, 1. 

Germany : Oberlausitz, Silesia, 1 ; Colpin, Brandenburg, 1 ; Magdeburg, 
Saxony, 2; Moritzburg, Saxony, 3 (U.S.N.M.). 

Austria-Hungary : Haida, Arva, Bohemia, 1. 

Switzerland: Canton Thurgau, 3 (U.S.N.M.); St. Gallon, 9 (B.M. 
and U.S.N.M.). 

Italy: Arezzo, 4 (B.M. and U.S.N.M.) ; Valesia, 1 (U.S.N.M.) ; Siena, 1 
(Mottaz) ; no exact locality, 4 (U.S.N.M.). 

Spain: Bellver, Lerida, 2 (B.M. and Genoa; paratypes of escalerai 
Cabrera) ; Seville, 1. 

Remarks. — This species is easily recognized by its rather small 
size, large ear, and very long, attenuate tragus. The exact point 
of insertion of wing membrane on side of foot is probably in all 
specimens the base of outer toe, but owing to a peculiar narrow- 
ing of the membrane along edge of metatarsal, the point of 
insertion sometimes appears to be at middle of side of foot. This 
effect is often increased by the action of strong alcohol. 



9. Inverary, Argyllshire, 

Scotland. 
1 al. Longford, Ireland. 

9. Woodpark, Galway. 

2 9, 9 juv. Harlech, Merioneth- 
shire, Wales. 

6. Arrow Church, Warwick- 

shire, England. 

8 al. Alcester, Warwickshire. 
5 9 al. Lilford Hall, Northamp- 
tonshire. 

9. Bradfield, Berkshire. 

9 st. Henley, Oxfordshire. 
<5 st. Henley, Oxfordshire. 

6 al. Queen Camel, Somerset. 
1 al. England. 
9. Oberlausitz, Silesia, 

340 m. Germany. 

(W. Baer.) 
i al. Colpin, Brandenburg, 

Prussia. 
<$ al. Magdeburg, Saxony. 

i. Magdeburg, Saxony. 

(Wolterstcrff.) 
9. Haida, Bohemia. (Wol- 

terstorff.) 
3<5,9. St. Gallen,600m. Swit- 
zerland. (E. H. Zolli- 
kofer.) 



Duke of Argyll (p). 58. 8. 16. 1. 

Dr. G. E. Dobson 76. 11. 3. 2. 

(c & p). 

R. F. Hibbert (p). 11. 1. 3. 20. 

J. Backhouse (p). 11. 1. 3. 16-18. 



Tomes Collection. 

Tomes Collection. 
Lord Lilford (o & p). 

N. H. Joy (c & p). 
J. G. Millais (c & p). 
Heatley Noble (c & p) . 
R. H. Read (p). 
Dr. J. E. Gray (p). 
Lord Lilford (p). 



7.1.1.488-493. 

7.1.1.736-743. 
72. 8. 21. 3-5. 
72. 11. 12. 13. 
11. 1. 3. 19. 
1. 11. 2. 1. 
0. 3. 23. 1. 



11. 
51. 
99. 



3. 21. 
29. 12. 
9.6. 



Dr. H. Gadow(c&p). 82.7.31.2. 
Dr. W. Wolterstorff 92. 12. 1. 1. 
Lord Lilford (p). 



Lord Lilford (p). 
O. Thomas (p). 



11. 1. 1. 8-9. 
11. 1. 1. 7. 
4. 4. 5. 4-8. 



MYOTIS 177 

2 al. Arezzo, Italy. Florence Museum (e). 85.7.6.4-5. 

9 al. Bellver, Lerida, Spain. A. Cabrera (p). 8. 7. 23. 4. 

(Paratype of M. escalerai, Cabr.) 
1 al. Seville. Dr. V. L. Seoane 94. 1. 1. 8. 

(o & p). 

MYOTIS EMARGINATUS Geoffroy. 

1806. Vesp[ertilid] cmarginatus Geoffroy, Ann. Mus. d'Hist. Nat., Paris, 
viii, p. 198 (Charlemont, Givet, Ardennes, Prance). 

1844. Vesp[ertilio] rufescens Crespon, Faithe Meridionale, i, p. 20 (near 
Nimes, Gard, Prance). Type in Nimes Museum. 

1853. V[espertilio] ciliatus Blasius, Wiegmann's Archiv fur Naturgesch., 
1853, i, p. 287 (near Cologne, Germany). 

1856. V[espertilio] schrankii Kolenati, Allgem. deutsche Naturhist. Zeitung, 
Dresden, neue Polge, n, p. 178. A nomen nudum (ex Koch and 
Giebel) cited as synonym of cmarginatus. Not of Wagner, 1843. 

1858. Vespertilio ciliatus Blasius, Saugethiere Deutschlands, p. 91. 

1878. Vespertilio emarginatus Dobson, Catal. Chiropt. Brit. Mus., p. 303. 

1880. Myotis ciliata var. budapestiensis Marg6, " Magyar orv. es termes- 
zetvizsg. xx,nagygydl. munk, p. 255" (Budapest, Austria-Hungary). 
See Mehely, Monogr. Chiropt. Hungariae, p. 170. 

1890. Vespertilio neglectus Patio, Arch. Sci. Phys. et Nat., Geneve, 3rd ser., 
xxiv, p. 512, November 15, 1890 (Valavran, near Geneva, Swit- 
zerland). Type in Geneva Museum. 

1900. Myotis emarginatus Mehely, Monogr. Chiropt. Hungariae, p. 170. 

1910. Myotis emarginatus Trouessart, Paune Mamm. d'Europe, p. 28. 

Type locality. — Charlemont, Givet, Ardennes, France. 

Geographical distribution. — Central and southern Continental 
Europe. 

Diagnosis. — Size essentially as in Myotis nattereri (forearm 
about 40 mm., condylobasal length of skull about 15 mm.) ; ear 
moderately long, extending 2 to 3 mm. beyond tip of muzzle 
when laid forward, its posterior border with a deep almost 
angular emargination slightly above middle ; foot about half as 
long as tibia ; wing membrane extending to base of outer toe ; free 
margin of interfemora'l membrane sometimes fringed ; fur some- 
what woolly in texture, the hairs of back tricolored, drab at 
base, yellowish at middle and dark brown at extreme tip. 

External characters. — Whole animal larger and more robust 
than Myotis mystacinus, though of essentially the same form and 
proportions. Ears and membranes relatively thick and leathery, 
not semi-transparent as in the related small species. Ear 
moderately long, extending about 2 mm. beyond nostril 
when laid forward, its size and general form much as in M. 
mystacinus except that emargination of posterior border is deep 
and conspicuous, in most specimens forming an evident angle 
below. Inner surface of conch marked by seven or eight short 
but well developed cross ridges. Antitragus small and ill-defined. 
Tragus slightly more than half as high as conch, its greatest 
width contained nearly three times in length of' anterior border, 
both its margins essentially straight from level of anterior base 



178 



CHIEOPTERA 



to rather acute tip. Wing as in the related species ; metacarpals 
falling short of elbow by about 3 mm. ; membrane inserted at 
base of outer toe. Calcar slender, its termination usually marked 
by a distinct lobe, its length about equal to that of free border 
of interfemoral membrane. Tail relatively shorter than in M. 
mystacinus, extending to between ears when laid forward, only 
the minute cartilaginous tip free from membrane. 

Fur and colour. — Fur shorter and more dense than in M. 
mystacinus and M. nattereri, and of a slightly woolly texture 
unique among the European species, the hairs at middle of back 
about 8 mm. in length. Distribution of fur not peculiar ; free 
border of uropatagium sometimes with evident fringe. General 
colour buff, light and clear on underparts, dulled and irregularly 
clouded by darker brownish hair tips throughout upper parts ; 
basal half of hairs drab ; muzzle and cheeks dusky ; ears and 
membranes an indefinite rather light brown. 

Skull. — In form the skull does not differ appreciably from 
that of M. nattereri, except that the rostral portion and palate are 
relatively more elongate and interorbital concavity is less strongly 
pronounced ; occipital region similarly low as compared with that 
of 31. mystacinus. Mandible slightly more robust than in M. 
nattereri, but of similar form. 

Teeth. — The teeth are larger than those of Myotis mystacinus, 
in this respect agreeing with those of M. nattereri. Upper 
incisors higher and more slender than in the preceding species, 
their crowns sub-equal in cross section ; lower incisors very 
slightly imbricated, forming a broadly and evenly convex 
(U-shaped) row without anterior angle, the crowns of i 1 and «, 
alike in form and distinctly 4-cusped, that of i x not thickened 
posteriorly and with no trace of postero-internal cusp. Canines 
about equal to those of M. nattereri in size, but shaft of upper 
tooth with evident postero-external longitudinal groove. First 
and second upper premolars even more strongly contrasted in 
size than those of M. mystacinus, the cusp of second only a httle 
exceeding cingulum of first ; the two teeth less crowded between 
canine and large premolar than in M. mystacinus and M. nattereri ; 
lower premolars essentially as in 31. nattereri, but less closely 
crowded, and second relatively larger. Upper molars as in 
M. nattereri, but crowns less narrowed ; a rudimentary commissure 
extending outward from base of hypocone and another extending 
inward from base of paracone ; lower molars not peculiar. 

3Ieasurements. — Two adult females from Florence, Italy 
head and body, 46' 6 and 50 ; tail, 40 and 42 ; tibia, 19 and 19 
foot, 8 - 4 and 86; forearm, 40 and 41 ; thumb, 8 and 7 - 8 
third finger, 67 and 70 ; fifth finger, 56 and 57 ; ear from meatus, 
16 - 6 and 17 ; width of ear, 11 "4 and 12. For cranial measure- 
ments see Table, p. 183. 

Specimens examined. — Six, from the following localities : — 
Holland : Maastricht, 1 (U.S.N.M.). 



MYOTIS 179 

France: Near Ntmes, Gard, 1 (Nimes : type of rufescens Crespon). 
Austria-Hungary : Herkulesbad, 2. 

Switzerland : Valavran, near Geneva, 1 (Geneva : type of neglectus 
Patio). 

Italy : Florence, 2 (U.S.N.M.) ; no exact locality, 1. 

Bemarhs. — The peculiar form of the ear, the short, somewhat 
woolly fur, and the yellowish colour are highly characteristic of 
this well-defined species. 

2 9. Herkulesbad, Hungary. Hon. N. C. Roths- 7. 9. 16. 9-10 

child (p). 
skeleton Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 733. 

without parte.) 
skull. 

MYOTIS BECHSTEINII Kuhl. 

1818. Vespertilio bechsteinii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv 

(= Neue Ann., i), pt. 1, p. 30 (Hanau, Hessen, Germany). 
1857. Vespertilio bechsteinii Blasius, Saugethiere Deutschlands, p. 85. 
1878. Vespertilio bechsteinii Dobson, Oatal. Chiropt. Brit. Mus., p. 308. 
1900. Myotis bechsteinii Mehely, Monogr. Chiropt. Hungarise, p. 184. 

1905. Vesp\ertilw\ bechst[emii] ghidinii vel Vesplertilio] ghidinii Patio, 

Arch. Sci. Phys. et Nat. ; Geneve, 4th ser., xix, p. 511, May 15, 
1905 (Lugano, Ticino, Switzerland). Type in Geneva Museum. 

1906. Myotis bechsteinii favonicus Thomas, Ann. and Mag. Nat. Hist., 

7th ser., xviii, p. 220, September, 1906 (La Granja, Segovia, 
Spain). Type in British Museum. 
1910. Myotis bechsteini and M. bechsteini favonicus Trouessart, Faune 
Mamm. d'Europe, pp. 30-31. 

Type locality. — Hanau, Hessen-Nassau, Germany. 

Geographical distribution. — Central and southern Europe, 
west to England, north to southern Sweden. 

Diagnosis. — Size slightly greater than that of M. nattereri 
(forearm about 40, condylobasal length of skull, 16 to 17) ; ear 
elongated, extending about 8 mm. beyond tip of muzzle when laid 
forward, the conch broad (about 15 mm.), its posterior margin 
obscurely emarginate above, the tragus scarcely half as high 
as conch ; foot about half as long as tibia ; wing membrane 
extending to base of outer toe. 

External characters. — In general like Myotis nattereri, but with 
broader ears and relatively as well as actually larger legs and 
feet (combined length of tibia and foot about 30 mm. instead of 
less than 25). Muzzle with moderately developed glandular 
swellings (these less evident than in M. mystacinus). Ear extend- 
ing considerably beyond nostril when laid forward, its length thus 
about as in M. nattereri, but its breadth so much greater (about 
17 mm. instead of about 10 mm.) that the ear is relatively larger 
than in any other European bat except Plecotus auritus ; form 
of ear essentially as in M. mystacinus, the upper half of posterior 
border faintly concave ; inner surface of conch with about eight 
rather ill-defined transverse ridges near posterior border ; anti- 
tragus about 2-5 mm. in length along base, rather abruptly 

n 2 



180 



CHIROPTERA 



rounded off above, its inner margin not continuous with posterior 
border of conch. Tragus about half as high as conch, its form 
essentially as in M. mystacinus though a little more slender 
(greatest width contained nearly three times in height of 
anterior border) ; basal lobe small but well developed. Wing 
essentially as in M. mystacinus; third, fourth and fifth meta- 
carpals sub-equal, falling short of elbow by about 5 mm. ; 
membrane inserted at base of outer toe. Calcar and free border 
of interfemoral membrane as in M. mystacinus. Tail shorter 
than head and body (laid forward it extends to middle of crown), 
the terminal vertebra free. 

Fur and colour. — Quality and distribution of fur as in Myotis 
mystacinus and M. nattereri. Upper parts uniform wood-brown 
with a slight tinge of umber, the exact shade almost impossible 
to describe exactly ; underparts a buffy grey in strong contrast, 
irregularly clouded by the slaty brown of underfur ; ears and 
membranes an indefinite dark brown. 

Skull. — In general aspect the skull resembles that of Myotis 
myotis, due allowance being made for its much smaller size, since 
it is much more slender than in any of the other small species ; 
breadth of bi - ain-case equal to about one-half distance from 
lambda to posterior border of narial emargination. Dorsal 
profile rising gradually above rostrum and forming a strong 
convexity over anterior portion of brain-case, behind which it is 
essentially flat to lambda ; ventral profile nearly flat, slightly 
elevated posteriorly ; occipital region slightly projecting, just 
sufficiently to conceal condyles when viewed from above. Brain- 
case ovate, noticeably wider than rostrum ; sagittal crest low but 
evident ; lambdoid crest moderately developed at sides, obsolete 
at middle ; greatest depth of brain-case about two-thirds mastoid 
breadth ; floor of brain-case with obscurely defined lateral grooves. 
Interorbital region not evidently hour-glass shaped, owing to the 
slight widening at lachrymal level ; anterior upper border of 
orbit slightly ridged. Rostrum slender, widening a little at 
anterior extremity, the dorsal surface smoothly rounded off at 
sides, though with indication of a slightly developed median 
groove ; narial emargination narrow but deep, extending back 
about half way to level of anterior rim of orbit ; rostral depth 
at front of orbit about equal to distance from orbit to middle of 
canine ; palate long and narrow as compared with that of the 
other small species, both transverse and lateral concavities evident, 
especially just behind middle ; anterior emargination rather large, 
sub-circular in general outline, extending back about to level of 
space between canine and first premolar ; posterior extension of 
palate about as broad as long ; interpterygoid space wider than 
long, encroached on by blunt median palatal spine. Mandible 
essentially as in Myotis myotis, the coronoid process higher and 
with more oblique posterior border than in the other small species. 

Teeth. — The teeth are rather small relatively to size of skulh 



MYOTIS 181 

Upper incisors essentially as in M. mystacinus but more slender. 
Lower incisors slightly imbricated, the row as a whole (J -shaped 
or very broadly V-shaped, the form of the individual teeth not 
peculiar. Canines with no special peculiarities. Small upper 
premolars completely in tooth-row, not crowded, the crown area of 
first equal to about half that of canine, that of second to about 
two-thirds that of first, the height of first a little less than half 
that of canine, that of second about two-thirds that of first ; 
cingulum complete but not forming secondary cusps. Large 
upper premolar with crown area nearly three-quarters that of 
first molar, its antero-internal cusp well developed. Lower 
premolars essentially as in M. nattereri. Upper molars with no 
special peculiarities except that m 3 is more reduced than in the 
other small species, its tranverse diameter through metacone 
noticeably less than half length of anterior border. Lower 
molars normal, but second triangle of m 3 a little more reduced 
than in the other small species, though much less so than in 
M. myotis and M. oxygnathus. 

Measurements. — Adult female from Sweden : head and body, 
46 ; tail, 38 ; tibia, 19 • 6 ; foot, 9 • 8 ; forearm, 42 ; thumb, 9 • 6 ; 
third finger, 67 ; fifth finger, 56 ; ear from meatus, 22 ; ear 
from crown, 19; width of ear, 13 - 6; tragus, 10. Two adults 
from the New Forest, Hampshire, England : tibia, 18-8 and 20 ; 
foot, 9 • 8 and 10 • 6 ; forearm, 39 and 42 ■ 4 ; thumb, 8 • 4 and 10 ; 
third finger, 64 and 68 ; fifth finger, 53 and 57. Adult male 
from Lugano, Switzerland (Geneva : type of ghidinii Fatio) : 
tibia, 19-4; foot, 10; forearm, 41 - 2; thumb, 10- 4; third 
finger, 65; fifth finger, 53; ear from meatus, 21 ±; ear from 
crown, 19; width of ear, 13; tragus, 11±. Adult female from 
Ste. Baume, Var, France : head and body, 53 ; tail, 37 • 4 ; tibia, 
20; foot, 10-4; forearm, 42- 6; thumb, 9; third finger, 70; 
fifth finger, 57 ; ear from meatus, 25' 6 ; ear from crown, 24; 
width of ear, 16' 2; tragus, 11" 8. Adult male from Cadillac, 
Gironde, France : head and body, 45-)- ; tail, 42 ; tibia, 20 ; 
foot, 9 ; forearm, 41 ; thumb, 8 ■ 8 ; third finger, 67 ; fifth 
finger, 56; ear from meatus, 26; width of ear, 15 • 6. Adult 
male from La Granja, Segovia, Spain (type of favonicus Thomas) : 
head and body, 48 ; tail, 38 • tibia, 20 ; foot, 8 • 4 ; forearm, 40 ; 
thumb, 8 • 2 ; third finger, 66 ; fifth finger, 55 ; ear from meatus, 
23; ear from crown, 19; width of ear, 14-6; tragus, 10-4. 
Adult female from Zay-Ugrocz, Hungary : head and body, 46 • 6 ; 
tail, 44 ; tibia, 22 ; foot, 9 ; forearm, 44 ; thumb, 9 ; third 
finger, 73 ; fifth finger, 59 ; ear from meatus, 25 ; ear from 
crown, 22-6; width of ear, 16; tragus, 11 8. For cranial 
measurements see Table, p. 183. 

Specimens examined. — Seventeen, from the following localities : — 
England : Henley-on-Thames, Oxfordshire, 1 ; New Forest, Hamp- 
shire, 6. 

Sweden: Skane, 1 (U.S.N.M.). 



182 



CHIROPTERA 



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184 



CHIROPTERA 



Fbance : Ktupes, Doubs, 1 (Mottaz) ; Ste. Baume, Var, 1 (Genoa) ; 
Cadillac, Gironde, 1 (Lataste). 

Switzeeland : Lugano, Ticino, 1 (Geneva: type of ghidinii Fatio). 
Spain: La Granja, Segovia, 1 (type of favonicus Thomas). 
Atjstbia-Hongaey : Zay-Ugr6cz, Trenesen, 4 (B.M. and U.S.N.M.). 

Remarks. — This animal is recognizable among European bats 
by its medium size and very large ears. On the basis of the 
material examined I am unable to distinguish a Spanish or 
western geographical race. 

<5. Henley-on-Thames, Ox- J. G. Millais (p). 6. 9. 14. 1. 

fordshire, England. 

1. New Forest, Hampshire. Dr. W. E. Leach (p). 56. A. 

2. New Forest, Hampshire. G.W.H.Blagg(c&p). 7.7.16.1-2. 
i al. La Granja, Segovia, M. de la Escalera (c). 6.11.4.1. 

Spain. (Type of M. favonicus Thos.) 

c5, ? al. Zay-Ugrocz, Trenesen, Budapest Museum 0. 6. 20. 1-2. 
Hungary. (e). 

1. Europe. Leyden Museum. 37. 4. 28. 23. 

MYOTIS DAUBENTONII Kuhl. 

1619. Vespertilio daubentonii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv 

(= Neue Ann., i), pt. 2, p. 195 (Hanau, Hessen, Germany). 
1839. Vespertilio sedilis Jenyns, Ann. Nat. Hist., in, p. 73, April, 1839 

(Aukland St. Andrew, Durham, England). 
1844. V[espertilio] lanatus Crespon, Faune Meridionale, I, p. 15 (South of 

Nimes, Gard, France). Type in Nimes Museum. 
1857. Vespertilio daubentonii Blasius, Saugethiere Deutschlands, p. 98. 
1871. Vespertilio capucinellus • ' Koch, Bayr. Fauna," Fitzinger, Sitzungsber. 

kais. Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1, 

p. 206 (Bavaria ?). 
1871. Vespertilio minutellus "Koch, Bayr. Fauna," Fitzinger, Sitzungsber. 

kais. Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1, 

p. 206 (Bavaria?). 
1871. Vespertilio daubentonii, albus Fitzinger, Sitzungsber. kais. Akad. 

Wissensch. Wien, Math.-Naturwiss. Classe, lxii, pt. 1, p. 210 

(Renaming of V. sedilis Jenyns). 
1878. Vespertilio daubentonii Dobson, Catal. Chiropt. Brit. Mus., p. 297. 
1890. Vespertilio staufferi Fatio, Faune Vert. Suisse, v, 3 mo suppi. aux 

Mamm., p. 6 (Lucerne, Switzerland). MS. synonym; repudiated. 
1900. Myotis daubentonii Mehely, Monogr. Chiropt. Hungarise, p. 164. 
1910. 2Ii/otis daubentoni Trouessart, Faune Mamm. d'Europe, p. 27. 

Type locality. — Hanau, Hessen-Nassau, Germany. 

Geographical distribution. — Europe from the Mediterranean 
north to central Norway, west to Ireland, east into Asia. 

Diagnosis. — Size nearly as small as in Myotis mystacinua 
(forearm about 35 mm., longest finger about 60 mm., condylo- 
basal length of skull, 13-4 to 13 -8); ear moderately long, 
extending about 2 mm. beyond tip of muzzle when laid forward, 
its posterior border with shallow inconspicuous emargination ; 
foot decidedly more than half as long as tibia ; wing membrane 
extending to side of metatarsus ; last caudal vertebra free. 



MYOTIS 185 

External characters. — Smallest of the large-footed European 
species. General form less slender and delicate than in M. 
mystacinus, the tail and legs relatively shorter. Ear rather 
.short, extending, when laid forward, about to tip of muzzle ; 
anterior border faintly and evenly convex from basal lobe to 
narrowly rounded-off tip ; posterior border with shallow though 
evident concavity above ; antitragus small and ill-defined ; tragus 
about half as high as conch, its greatest width contained three 
times in length of anterior border, the posterior border 
moderately and evenly convex, the anterior border 'nearly 
straight, the tip rather blunt ; posterior basal lobe relatively 
large and well denned. Wing slightly broader than in M. mysta- 
cinus, the three main metacarpals evidently graduated from 
third to fifth, the third very slightly shorter than forearm ; 
membrane inserted at middle of metatarsus. Foot large, 
appreciably more than half as long as tibia ; calcar slender, 
very long, without keel on posterior border and with slight 
terminal lobe, its length fully double that of free border of 
interfemoral membrane. Tail about as long as body without 
head, the terminal vertebra free from membrane except for an 
exceedingly narrow strip extending outward along each side. 

Fur and colour. — Fur slightly shorter and more dense than 
that of M. mystacinus, but with no peculiarities of distribution 
except that it tends to spread farther outward along dorsal 
surface of interfemoral membrane ; free border of uropatagium 
not fringed, but a slight fringe is usually present along basal 
half of calcar. Colour above essentially as in M. na/lereri, 
though usually inclining more definitely toward wood-brown ; 
underparts buffy grey usually less contrasted than in M. nattered, 
though sometimes pale enough to produce a distinct line of 
demarcation along sides of neck. Muzzle and cheeks dusky. 
Membranes and ears an indefinite brown. 

Skull. — The skull of Myotis daubentonii is smaller than that 
of any other European species except M. mystacinus. From this 
it is immediately distinguishable by its noticeably greater breadth 
both of rostrum, palate and brain-case, by the relatively lower 
occipital region, and relatively deeper rostrum. Posterior exten- 
sion of palate short and broad, the width just behind molars 
greater than length to tip of hamular ; median projection angular, 
seldom forming a distinct spine. Greatest breadth of brain-case 
slightly though appreciably more than greatest length of skull. 
Mandible with coronoid process low, scarcely rising above level 
of condyle. 

Teeth. — Teeth relatively smaller than those of Myotis mysta- 
cinus, a difference particularly noticeable in the crown areas of 
the first and second upper molars. Upper incisors as in mystacinus, 
but with cingulum less developed. Lower incisors very slightly 
imbricated, the form of the row vacillating between (J -shaped 
and broadly V-shaped, the cusps as in M. mystacinux but less 



186 



CHIKOPTERA 





FIG. 31. 
Mi/otis daubentonii. Teeth x 10. 



developed. Canines both above and below weaker and less 
trenchant than in M. myslacinus, their form essentially as in 

M. nattereri. Premolars 
and lower molars not 
obviously different from 
those of M. mystacinus. 
Upper molars peculiar as 
compared with those of 
the small-footed European 
species in the presence of 
an evident protoconule on 
anterior commissure of 
protocone, the small cusp 
provided with a small 
but distinct secondary 
commissure extending to 
base of paracone ; m 1 and 
m 2 with a small commis- 
sure extending from base 
of hypocone to base of 
metacone and partly fill- 
ing depression lying be- 
tween these cusps. 

Measurements. — Two 
adult males from Upsala, 
Sweden : head and body, 43 and 44 ; tail, 34 and 36 ; tibia, 17 
and 17-4 ; foot, 11 and 11 ; forearm, 37 and 37 ; thumb, 8 - 4 
and 8 ; third finger, 62 and 59 ; fifth finger, 49 and 49 ; ear 
from meatus, 13 and 13 ; width of ear, 10 and 9 - 6. Adult male 
and female from Lecco, Italy : head and body, 42 and 45 ; tail, 
36. and 39; tibia, 16 and 17; foot, 10 • 6 and 11; forearm, 
37 and 38 ; thumb, 8 and 8 ; third finger, 62 and 62 ; fifth finger, 
51 and 51 ; ear from meatus, 13 '6 and 13' 6; width of ear, 9*6 
and 9 - 4. For cranial measurements see Table, p. 190. 

Specimens examined. — Seventy-three, from the following localities : — 

Scotland : No exact locality, 1. 

England : Bowdon, Cheshire, 1 ; Knutsford, Cheshire, 1 ; Stratford- 
on-Avon, Warwickshire, 2 ; Hillingdon, Middlesex 1, ; Epping, Essex, 2 ; 
Northamptonshire, 1 ; Henley-on-Thames, Oxfordshire, 2 (B.M. and 
U.S.N.M.) ; Christchnrch, Hampshire, 1 ; Devonshire, 1. 

Sweden: Upsala, 3 (B.M. and U.S.N.M.); upland, 1; no exact locality, 2. 

Switzerland : Geneva, 1 (Mottaz). 

Pkance : Near Nimes, Gard, 1 (Nimes : type of lanatus Crespon). 

Italy : Lecco, Lombardy, 21 (U.S.N.M.) ; Pavia, 1 (U.S.N.M.); 
Florence, 30 (Mottaz). 

Remarks. — This species is immediately recognizable among 
the European members of the genus by its small size, large foot, 
and naked upper surface of legs. 

1. Scotland. Dr. J. Macgiilivray (p). 

6. Bowdon, Cheshire, Eng- T. A. Coward (c & p). 11. 1. 3. 60. 
land. 



MYOTIS 187 

?. Knutsford, Cheshire. T. A. Coward (c & p). 11. 1. 3. 22. 

2. Stratford-on- Avon, War- Tomes Collection. 7.1.1.486-487. 

wickshire. 

6 al. Hillingdon, Middlesex. 0. Thomas (c & p). 84. 1. 29. 1. 

2 al. Epping, Essex. H. Doubleday (c &p). 44. 10. 21. 1-2. 

al. Northampton. Mrs. Jenyns (p). 

9 st. Henley, Oxfordshire. J. G. Millais (o & p). 1. 11. 2. 2. 

1 al. Christohurch, Hamp- Lord Lilford (p). 87. 9. 1. 1. 

shire. 

S. Upland, Sweden. Lord Lilford (p). 11. 1. 1. 25. 

(G. Kolthoff.) 

2. Sweden. Stockholm Museum 46. 6. 2. 15. 

(e). 48. 6. 28. 3. 



MYOTIS CAPACCINII Bonaparte. 

1837. VespertiUo capaccinii Bonaparte, Iconogr. Faun. Ital., i, fasc. xx 

(Sicily). Type in British Museum. 
1839. VespertiUo megapodius Temminck, Monogr. de Mamm., n, p. 189 

(Sardinia). 
1841. Vesp[ertilio] dasypus de Selys-Longchamps, Atti della seconda 

Riunione degli Scienziati Italiani, Torino, 1840, p. 249 (Published 

as synonym of capaccinii). 
1844. Vesp[ertilio] pellucens Crespon, Faune Meridionale, i, p. 16 (Cave 

near Pont-du-Gard, Gard, Erance). 
1857. VespertiUo capaccinii Blasius, Saugethiere Deutschlands, p. 101. 

1877. VespertiUo blasii Major, Atti Soc. Tosc. Sci. Nat., Pisa, in, p. 108 

(New name for the capaccinii of Blasius should it prove to be 
different from that of Bonaparte) . 

1878. VespertiUo majori Ninni, Atti Beale Instit. Veneto, 5th ser., iv, 

pt. 1, p. 721 (Substitute for blasii Major). 
1878. VespertiUo capaccinii Dobson, Catal. Chiropt. Brit. Mus., p. 293. 
1901. Myotis capaccinii Thomas, Proc. Zool. Soc. London, p. 37. 
1910. Myotis (Leuconoe) capaccinii Trouessart, Paune Mamm. d'Europe, p. 26. 

Type locality. — Sicily. 

Geographical distribution. — Mediterranean region, north to 
Italian Switzerland, east into Asia. 

Diagnosis. — Not so small as Myotis daubentonii (forearm about 
42 mm., longest finger, about 68 mm., condylobasal length of 
skull, 14 "0 to 14 - 8 mm.), but similar in form and proportions ; 
wing membrane extending to ankle ; last caudal vertebra free ; 
tibia and adjacent membrane densely furred. 

External characters. — In all essential features the external 
form is as in M. daubentonii. The foot, however, is relatively 
larger, and the wing membrane is inserted at the ankle. 

Fur and colour. — Fur rather dense and short, the hairs at 
middle of back about 6 mm. in length; distribution peculiar 
among the European members of the genus in its tendency to 
spread on membranes, forming a distinct patch on upper surface 
of wing at elbow and extending over entire uropatagium to level 
of feet, the furry covering of tibia and immediately adjacent 
membrane (both above and below) especially dense. Colour 



188 



CHIEOPTERA 



above a light drab tinged with grey or with ecru-drab, the 
general effect paler and more greyish than in any of the other 
European species ; underparts pale buffy grey, rather strongly 
contrasted and with moderately well-defined line of demarcation 
along sides of neck. Underfur slaty black. Muzzle and 
cheeks faintly dusky. Ears and membranes an indefinite rather 
light brown. 

Skull. — In all respects the skull so closely resembles that of 
Myotis daubentonii that it is only distinguishable by its larger 
size. Erom the skulls of M. nattereri and M. emarginatus it 
differs in its greater breadth, a character perhaps most readily 
appreciable in the form of the post-palatal region. 

Teeth. — Except for their larger size the teeth resemble those 
of Myotis daubentonii. The crown area of upper molars is 
relatively less than in M. nattereri and M. emarginatus, though 
m 3 retains the broader outline characteristic of M. daubentonii 
and M. mystacinus. Upper molars with protoconule and its 
accessory small commissure, and m 1 and m 2 with commissure 
between hypocone and metacone as in M. daubentonii. 

Measurements. — Average and extremes of forearms in four 
males and eight females from Lugano, Ticino, Switzerland : 
males, 39-9 (38-8-41); females, 39-4 (38-8-40-4). Sicily 
(type): tibia, 15- 6; foot, 12; forearm, 39-2; thumb, 10 ; third 
finger, 56 ■ 2 ; fifth finger, 50. Adult male from Oorleone, Sicily : 
head and body, 49 ; tail, 38 ; tibia, 16 ; foot, 10 ; forearm, 41 ; 
thumb, 8 • 2 ; third finger, 68 ; fifth finger, 55 : ear from meatus, 
1 5 ; width of ear, 10-4. Two adult males from Sassari, Sardinia : 
head and body, 50 and 51 ; tail, 37 and 37 ; tibia, 16 -4 and 17 ; 
foot, 10-6 and 11 ; forearm, 39-4 and 41-6 ; thumb, 8-4 and 
8-6 ; third finger, 66 and 69 ; fifth finger, 51 and 56 ; ear from 
meatus, — and 14*4; width of ear, 10 and 10. For cranial 
measurements see Table, p. 191. 

Specimens examined. — Seventy-two, from the following localities : — 

Austria-Hungary : Herkulesbad, 1. 

Switzerland : Near Lugano, 49 (B.M., U.S.N. M., and Mottaz). 

Italy: Finalborgo, Liguria, 2 (Genoa); iPavia, 1 (U.S.M.M.) ; Ostia, 
Rome, 2; Marsala, Sicily, 1; Corleone, Sicily, 1 (U.S.N. M.); Sicily, no 
exact locality, 1 (type). 

Sardinia : Cagliari, 4 (B.M. and U.S.N.M.) ; Sassari, 5 (U.S.N.M.) ; 
Grotte de Sardale, 2. 

France : Marseilles, 1 (U.S.N.M.). 

Spain: Inca, Majorca, Balearic Islands, 1; Elche, Alicante, 1. 

Hemarlts. — In general appearance this species resembles 
Myotis daubentonii ; but it is immediately recognizable by its 
even larger foot, and by the densely pubescent upper surface of 
leg, the latter character unique among European bats. 

4 6, ?. Near Lugano, Ticino, O. Thomas (p). 2. 8. 4. 10-14. 

Switzerland. 

(E. H. Zollikofer.) 
2. Ostia, Rome. Dr. L. Sambon(c&p). 1.1.2.3-4. 



189 



skeleton Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 734. 

without parte.) (Type of species.) 
skull. 

9. Marsala, Sicily. 0. Thomas (p). 6. 8. 4. 21. 

(A. Robert.) 

2 al. Cagliari, Sardinia. FlorenceMuseum(E). 85.7.6.6-7. 

2 al. Grotte de Sardale, Sar- 0. Thomas (p). 0. 12. 3. 1-2. 
dinia. 

c5. Inca, Majorca, Balearic 0. Thomas and R. I. 0. 7. 1. 3. 

Islands. Pocock (c & p). 

6. Elche, Alicante, Spain. G. S. Miller (c). 8. 8. 4. 13. 



MYOTIS DASYCNEME Boie. 

1823. VespertiUo mystacinus Boie, Isis, p. 965. Not V. mystacinus Kuhl 

(Jutland, Denmark). 
1825. VespertiUo dasycneme Boie, Isis, p. 1200 (Renaming of mystacinus 

Boie). 
1839. VespertiUo limnophilus Temminck, Monogr. de Mamm., n, p. 176 

(Holland). 
1857. VespertiUo dasycneme Blasius, Saugethiere Deutschlands, p. 103. 
1878. VespertiUo dasycneme Dobson, Catal. Chiropt. Brit. Mus., p. 295. 
1904. Myotis dasycneme Trouessart, Catal. Mamm. Tam viv. quam foss., 

suppl., p. 88. 
1910. Myotis dasycneme Trouessart, Paune Mamm. d'Europe, p. 27. 

Type locality. — Chalk quarries at Dagbierg, near Wiborg, 
Jutland, Denmark. 

Geographical distribution. — Central and southern Europe, 
west to the Atlantic coast,* north to Sweden, east into Asia. 

Diagnosis. — Form and proportions essentially as in Myotis 
daubentonii and M. capaccinii, but size much larger (forearm 
about 47 mm., longest finger about 75 mm., condylobasal length 
of skull about 16 mm.) ; tibia and adjacent membrane naked. 

External characters. — Form not essentially different from the 
other European members of the large-footed group. Tragus 
relatively shorter than in any other European Myotis, its height 
distinctly less than half that of conch, its anterior border slightly 
concave, its posterior border slightly convex below, then more 
abruptly convex to bluntly rounded- off tip, the two margins 
essentially parallel through lower half. Free border of uro- 
patagium without fringe. 

Fur and colour. — Distribution of fur as in Myotis daubentonii ; 
quality not peculiar, the longest hairs on middle of back about 
8 mm. in length. Colour of upper parts a light yellowish wood- 
brown ; underparts strongly contrasted greyish white with a 
tinge of buff, the line of demarcation along sides of neck well 
defined. Muzzle and cheeks scarcely contrasted dusky. Under- 
fur slaty black. Ears and membranes an indefinite dark brown. 

* The supposed British record is probably erroneous (see Barrett- 
Hamilton, Hist. Brit. Mamm., I, pp. 157-158. December, 1910). 



190 



CHIROPTERA 



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CHIEOPTERA 



Skull. — The skull is large, slightly exceeding that of Myotis 
bechsteinii in length and noticeably surpassing it in breadth and 
robustness. Its general appearance is the least typically Myotis- 
like of any European member of the genus, a peculiarity 
heightened by the crowding of the small premolars and conse- 
quent shortening of anterior portion of tooth-row. Allowance 
being made for the great difference in size the skull is not very 
unlike that of Myotis daubentonii, but the rostrum is relatively 
shorter and the brain-case broader and more depressed. Sagittal 
crest barely indicated ; auditory bullae proportionately as in the 
smaller animal, less inflated than in M. bechsteinii ; lachrymal 
ridge well developed, its lower extremity separated from lachrymal 
foramen by an evident notch. 

Teeth. — In form the teeth resemble those of the small Myotis 
daubentonii ; molars decidedly more robust than those of M. 
bechsteinii, the width of the crown diminishing much less rapidly 
toward inner border ; small premolars much crowded, the second 
barely or not visible from outer side, the diameter of its crown, 
however, not much less than that of anterior tooth ; upper 
molars as in M. daubentonii and M. capaccinii ; mandibular teeth 
with no special peculiarities, the premolars and outer cusps of 
molars more slender than in M. bechsteinii and M. daubentonii. 

Measurements. — Two adult males from Maastricht, Holland : 
head and body, 57 and 58; tail, 49 and 51 ; tibia, 19 ■ 8 and 
20 -2 ; foot, 11 '8 and 12 ; forearm, 43 6 and 44 ; thumb, 9-6 
and 9 ; third linger, 72 and 77 ; fifth finger, 57 and 62 ; ear 
from meatus, 17 and 17 ' 2 ; width of ear, 10 - 6 and 11. Two 
adult females from the same locality : head and body, 60 and 61 ; 
tail, 47 and 46 ; tibia, 18 and 18 ; foot, 11-4 and 11 ; forearm, 
44 and 44 ; thumb, 9 and 9 ; third finger, 76 and 75 ; fifth 
finger, 61 and 60; ear from meatus, 16 • 6 and 17 ; width of ear, 
11 and 11. For cranial measurements see Table, p. 191. 

Specimens examined. — Six, from the following localities : — 
Holland : Leyden, 1 ; Maastricht, 4 (U.S.N.M.). 
Belgium : Near Namur, 1. 

1. Leyden, Holland. Tomes Collection. 7. 1. 1. 501. 

(H. Schlegel.) 
6 al. Namur, Belgium. Bev. D. B. Lebbe 9. 1. 11. 1. 

(c & p). 

MYOTIS MYOTIS Borkhausen. 

1775. Vespertilio murinus Schreber, Saugthiere, I, p. 165 (Not of Linnaeus, 

1758). 
1797. Vespertilio myotis Borkhausen, Deutsche Fauna, i, p. 80 (Germany). 
1797. Vespertilio myosotis B[orkhause]n, Der Zoologe (Compendiose 

Bibliothek gemeinnutzigsten Kenntnisse fur alle Stande, pt. xxi), 

Heft v-vin, p. 46 (Germany). 
1800. Vespertilio myosotis Bechstein, Pennant's Allgemeine Uebersicht der 

Vierfiissigen Thiere, p. 632 (Germany). 



MYOTIS 193 

1801. Vespertilio myotis Bechstein, Gemeinn. Naturgesch. Deutschl., i, 

2nd ed., p. 1154 (Described but not named in 1st ed., 1789, p. 164) 

Thiiringen, Germany. 
1827. Vespertilio submurinus Brehm, Ornis, Heft in, p. 24 (Renthendorf, 

Thiiringen, Germany). 
1844. V[espertilio] latipinnis Crespon, Paune Meridionale, I, p. 17 (Near 

Nimes, Gard, France). 
1857. Vespertilio murinus Blasius, Saugethiere Deutschlands, p. 82 (Not of 

Linnaeus, 1758). 
1863. [Myotus murinus'] var. typus Kocb, Jahrb. des Vereins fur Natur- 

kunde im Herzogthum Nassau, xvin, p. 415 (Wiesbaden, Nassau, 

Germany). 
1863. [Myotus murinus] var. alpinus Koch, Jahrb. des Vereins fur Natur- 

kunde im Herzogthum Nassau, xvm, p. 415 (St. Gothard, Uri, 

Switzerland). 
1878. Vespertilio murinus Dobson, Catal. Chiropt. Brit. Mus., p. 309 (Not 

of Linnaeus, 1758). 
1886. Myotis murma var. spelsea Bielz, Verhandl. u. Mittheilungen des 

Siebenbiirgischen Vereins fur Naturwissensch. in Hermannstadt, 

xxxvi, p. 83 (Homorod- Almas cave, Hungary). 
1897. Myotis myotis Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 383, 

October, 1897. 

1909. Myotis myosotis Miller, Ann. Mus. Zool. R. Univ. Napoli, N.S., in, 

No. 3, p. 1, April 26, 1909. 

1910. Myotis myotis and M. m. spelsea Trouessart, Faune Mamm. d'Burope, 

p. 32. 

Type locality. — Thiiringen, Germany. 

Geographical distribution. — Central and southern Continental 
Europe, west to Portugal, north to southern Sweden, eastward 
into Asia. One record of its occurrence in England.* 

Diagnosis. — Largest species of European Myotis (forearm, 
57 to 64 mm.; longest finger, 100 to 110 mm.; condylobasal 
length of skull, 22 to 23 ■ 6 mm.); form heavy, membranes thick 
and leathery ; ear moderately long, extending about 5 mm. 
beyond tip of muzzle when laid forward, its posterior margin 
scarcely or not emarginate above middle ; foot slightly more than 
half as long as tibia ; wing membrane extending to base of 
outer toe. 

External characters. — Although one of the largest European 
bats Myotis myotis does not differ conspicuously in form from 
the small M. mystacinus, except that its tail and legs are relatively 
shorter. The general build is not remarkably heavy as compared 
with other European species of approximately the same size, but 
the ears and membranes are rather thick and leathery. Ear 
moderately long, extending about 5 mm. beyond nostril when 
laid forward ; anterior margin moderately convex from base 
nearly to rather narrowly rounded-off tip ; posterior border with 
shallow ill-defined concavity above ; antitragus low and long, 
marked off posteriorly by a well-defined notch and not continuous 
with posterior border of conch ; tragus about half as high as 

* Bell, Hist. British Quadrupeds, p. 38, 1836 : "But in England it . . . 
has hitherto only been taken in the gardens of the British Museum." 

O 



194 



CHIEOPTBEA 



conch, its greatest width (slightly above level of anterior basey 
contained about 2J times in length of anterior margin, the 
posterior border convex to just below rather bluntly rounded tip, 
the anterior margin straight, posterior basal lobe small but well 
developed ; inner surface of conch with seven or eight ill-defined 
transverse ridges near posterior border. Wing rather broad but 
with no special peculiarities of form ; metacarpals somewhat 
elongated relatively to phalanges as compared with the smaller 
species, the third, fourth and fifth slightly but evidently 
graduated, the third scarcely shorter than forearm ; membrane 
inserted at side of metatarsus, but with a narrow strip extending 
to base of outer toe as in M. nattereri. Foot slightly more than 
half as long as tibia ; calcar heavy at base but tapering rapidly 
and terminating obscurely, its posterior border with slightly 
indicated keel, its length about equal to that of free border of 
interfemoral membrane. Tail about as long as body without 
head, only the extreme cartilaginous tip free from membrane. 

Fur and colour. — Relatively to size of animal the fur is rather 
short (longest hairs of back about 10 mm.); in distribution it 
shows no peculiarities ; free border of uropatagium without 
fringe. Colour an indefinite brown much like that of Myotis 
nattereri, the exact shade intermediate between the wood-brown 
and broccoli-brown of Ridgway, usually paler on head and neck 
than on back, and in immature individuals than in adults, the 
hairs slate-black through basal half, then light wood-brown 
followed by a darker though not strongly contrasted terminal 
area. Underparts strongly contrasted greyish white with a 

slight buffy tinge ; a well-defined 
line of demarcation along sides 
of neck to ear, emphasized in 
region of shoulder by a slight, 
diffused blackening of edge of 
dark area. Muzzle and cheeks 
dusky. Ears and membranes an 
indefinite brown. 

Skull. — Though much larger 
than that of any of the other 
European species of Myotis, M. 
oxygnathus excepted, the skull of 
Myotis myotis is one of the most 
slender in general outline. The 
brain-case is longer proportion- 
ately to its breadth and is less 
contrasted with rostrum than in 
the small members of the group ; 
greatest breadth of brain-case 
about one-half distance from 
lambda to posterior margin of nares. Rostrum relatively deep 
and interorbital concavity relatively shallow ; occipital region 






no. 32. 
Myotis myotis. Nat. size. 



195 



about on level with main portion of brain-case, and very slightly 
overhanging foramen magnum ; ventral profile scarcely elevated 
posteriorly ; palate rather narrow (essentially as in M. mystacinus) ; 
width of posterior extension of palate less than its length, median 
spine short but well developed ; posterior border of anteorbital 
foramen over anterior root of first molar ; mandible with coronoid 
process relatively higher and narrower than in the small species, 
its posterior border much more oblique. 

Teeth. — The dentition is of a less primitive type than in the 
small European species of Myotis. This is indicated by the 
general tendency to reduction shown especially in the lower 
incisors, the small premolars and the posterior lower molar. 
Upper incisors rather high and slender but not essentially 
different in form from those of M. mystacinus, the cingulum of 
inner tooth obsolete, that of outer slightly developed. Lower 
incisors very strongly im- 
bricated, the outline of the 
row U-shaped or broadly 
V-shaped ; cutting edge of 
ij and i 2 trifid, but decidedly 
oblique owing to reduction 
in size of outer cusp ; a 
minute cingulum cusp usu- 
ally present at extreme outer 
edge ; i 2 with small postero- 




FIG. 33. 
Myotis myotis. Teeth X 10. 



internal tubercle ; i 3 sub- 
terete, 4-tuberculate, the me- 
dian outer tubercle largest. 
Canines relatively smaller 
than in M. mystacinus and 
with less developed posterior 
cutting edge, their form 
essentially as in M. natter eri 
and JIT. emarginatus. Small 
upper premolars much 
crowded, the second usually 
forced inward from line of tooth-row, sometimes so much so that 
the first is practically in contact with large premolar. In form 
they show no special peculiarities ; shaft subterete ; cingulum 
well developed. Large upper premolar more reduced than in the 
small species, the inner margin of crown a mere cingulum at base 
of main cusp, and without trace of secondary cusps. Lower 
premolars not peculiar in form, but middle tooth usually more 
crowded between first and third than in the small members 
of the group. Molars showing no special peculiarities of form ; 
thickening that represents hypocone in m 1 and m 2 barely 
indicated ; m 3 more reduced than in the small species, its meta- 
cone scarcely more than a slight widening of terminal portion of 
third commissure. First and second lower molars essentially as 

o 2 



196 



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198 



CHIROPTEEA 



in the small species, except that cingulum does not form a postero- 
internal cusp behind entoconid ; m 3 with hypoconid and ento- 
conid more reduced than in the small species, the hypoconid 
displaced further inward, so that second triangle is barely half 
as large as first and conspicuously different from it in form. 

Measurements. — Average and extremes of four adult females 
from Tagerwilen, Thurgau, Switzerland: head and body, 75 "7 
(72-79); tail, 56"3 (54-60); tibia, 26-1 (25-4-26-6); foot, 
14-7 (13-16); forearm, 63 (63); thumb, 13-1 (12 -8-13 -4) ; 
third finger, 107-7 (106-109); fifth finger, 85-2 (84-86); ear 
from meatus, 27 • 6 (27-28) ; width of ear, 18-1 (17 ■ 6-19). Two 
adult females from Mte. Generoso, Ticino, Switzerland : head 
and body, 72 and 76 ; tail, 50 and 51 ; tibia, 25 and 25-4 ; foot, 
15 and 14- 4 ; forearm, 62 and 63-6 ; thumb, 13 and 12 ; third 
finger, 104 and 107 ; fifth finger, 83 and 84 ; ear from meatus, 
27 and 28; width of ear, 18 and 18-6. Adult male from 
Florence, Italy : head and body, 68 ; tail, 55 : tibia, 25 ; foot, 
13; forearm, 61 ; thumb, 12 ; third finger, 100 ; fifth finger, 78 ; 
ear from meatus, 27- 6; width of ear, 17-6. For cranial 
measurements see Table, p. 196. 

Specimens examined. — Ninety-five, from the following localities : — 

France : Cadillac, Gironde, 1 (U.S.N.M.) ; Nimes, Gard, 2 (B.M. and 
Nimes ; representing latipinnis Crespon, but not type). 

Germany : Hamburg, 1 ; Niesky, Silesia, 11 ; Strass, near Burgheim, 
Bavaria, 8 ; Heidelberg, Baden, 1. 

Aostria-Hungary : Herkulesbad, 1 ; Fiinfkirchen, S.W. Hungary, 2. 

Roumania: Bustenari, 3; Sinaia, 1 (U.S.N.M.). 

Switzerland : Geneva, 7 (Mottaz) ; Grotte de Vallorbe, Vaud, 1 
(Mottaz) ; Boudry, Neuchatel, 2 (Mottaz) ; St. Moritz, 1 ; Thayngen, Schaff- 
hausen, 2 (U.S.N.M.); Canton Thurgau, 5; Tagerwilen, Thurgau, 10 
(U.S.N.M.); Andermatt, Uri, 1 (U.S.N.M.); St. Gothard, Uri, 3 (B.M. 
and U.S.N.M.) ; Monte Generoso, Ticino, 4 (U.S.N.M.). 

Italy : Domodossola, 2 (U.S.N.M.) ; Finalborgo, Liguria, 1 (Genoa) ; 
Florence, 1 (U.S.N.M.); Rome, 2; Ostia, Borne, 2; Marsala, Sicily, 6. 

Sardinia : Oristano, Cagliari, 10. 

Spa™ : Seville, 2. 

Portu&al : Cintra, 2. 

Remarks. — Myotis myotis differs strikingly from the other 
European members of the genus, M. oxygnathus excepted, in its 
much larger size. From the large Vespertilionid.ee of other genera 
it is immediately recognizable by its long ears, extending notice- 
ably beyond nostril when laid forward, and by the greyish white 
colour of the underparts. In the Mediterranean region Myotis 
myotis is associated with M. oxygnathus ; but it is the only large 
species known to occur north of the Alps. 



lal. 


Nimes, Gard, Prance. 


G. E. Dobson (e). 


80. 12. 14. 2. 


1. 


Hamburg, Germany. 


Dr. J S E. Gray (p). 




2 6,8 9, 


Niesky, Silesia. (Dr. 


Dr. E. Hamilton (p). 


97. 12. 4. 7-17. 


? juv. 


W. Baer.) 






6 9. 


Strass, Burgheim, Ba- 
varia. (Korbitz.) 


Lord Lilford (p). 


11. 1. 1. 10-15. 


?. 


Strass, Burgheim, Ba- 


Lord Lilford (p). 


11. 1. 1. 124. 




varia. (Korbitz.) 




11. 1. 1. 131. 



MYOTIS 199 

9. Heidelberg, Baden. Hon. N. C. Botha- 10. 5. 29. 1. 

child (p). 

■i juv. Herkulesbad, Hungary. Hon. W. Rothschild 7. 9. 16. 8. 

(F. J. Cox.) (p). 

2 al. Piintkirchen. Budapest Museum (e). 94.7. 18. 11-12. 

2 (J, 9. Bustenari, Prahova, Lord Lilford (p). 4. 4. 6. 8-10. 
840 m. Boumania. 
(W. Dodson.) 

1. St. Moritz,Grisons,Swit- Leon 0. Galliard (p). 75.9.20.4. 

zerland. 

5 9. Thurgau. (E. H. Zolli- 0. Thomas (p). 4. 4. 5. 9-13. 

kofer.) 
2 9. Borne. (C. Goli.) G. Barrett-Hamilton 11. 1. 2. 26-27. 

2. Ostia, Borne. Dr.L. Sanation (c&p). 1.1.2.1-2. 

6 9. Marsala, Sicily. (A. O. Thomas (p). 6. 8. 4. 17-20. 

Robert.) 8. 9. 1. 3-4. 

10 al. Oristano, Cagliari, Sar- Hon. N. C. Boths- 7. 5. 24. 1-10. 

dinia. child (p). 

2 6 al. Seville, Spain. (Dr. A. Lord Lilford (p). 73. 1. 8. 1-2. 

Ruiz.) 

2 1. Cintra, Portugal. 0. Thomas (o & p). 98. 2. 2. 4-5. 

MYOTIS OXYGNATHUS Monfcicelli. 

1885. Vespertilio oxygnathus Monticelli, Ann. Accad. 0. Costa de Aspir. 
Nat., I, p. 82. Type in Naples Museum. 

1909. Myotis oxygnathus Miller, Ann. Mus. Zool. B. Univ. Napoli, N.S., 

in, No. 3, p. 1, April 26, 1909. 

1910. Myotis myotis oxygnathus Trouessart, Faune Mamm. d'Burope, p. 32. 

Type locality. — Matera, Basilicata, Italy. 

Geographical distribution. — Mediterranean region from Spain 
to Greece, north to Italian Switzerland ; Sardinia ; Malta ; 
Tunis. 

Diagnosis. — Similar to Myotis myotis but smaller and with 
shorter, narrower ears; condylobasal length of skull, 18 ■ 6 to 
21 • 4 instead of 22 to 23 • 6 mm. ; mandible, 15*2 to 17-2 instead 
of 17' 8 to 19 mm. ; maxillary tooth-row, 8 '2 to 9'4 instead of 
9-8 to 10-6 mm. 

Measurements. — Type (adult male) : head and body, 63 ; tail, 
54; tibia, 24-4; foot, 13; forearm, 57; thumb, 11-4; third 
finger, 98 ; fifth finger, 76 ; ear from meatus, 23 ; width of ear, 
13-6; tragus, 1 ■ 8. Two adult males from Velletri, Rome, 
Italy : head and body, 60 and 62 ; tail, 58 and 58 ; tibia, 23 
and 24 • 6 ; foot, 1 2 • 8 and 1 3 ; forearm, 53 • 6 and 57 ; thumb, 1 2 
and 11 ; third finger, 89 and 86 ; fifth finger, 73 and 76 ; ear 
from meatus, 23 and 24 ; width of ear, 14 and 15. Two adult 
females from the same locality : head and body, 62 and 66 ; 
tail, 58 and 57 ; tibia, 24 and 24- 6 ; foot, 13 and 12 ; forearm, 
56 and 58 ; thumb, 11 and 11 -8 ; third finger, 93 and 97 ; fifth 
finger, 74 and 78 ; ear from meatus, 23 and 22 ; width of ear, 
13 ■ 6 and 13-6. Adult male and female from Bozen, Tirol : head 
and body, 68 and 71 ; tail, 53 and 53 ; tibia, 25-4 and 24 ; foot, 



200 



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202 



CHIROPTEEA 



14 and 14 ; forearm, 59 and 60 ; thumb, 11 • 4 and 11 • 6 ; third 
finger, 97 and 98 ; fifth finger, 76 and 78 ; ear from meatus, 26 
and 25 ; width of ear, 17 and 15. For cranial measurements see 
Table, p. 200. 

Specimens examined. — Seventy-four, from the following localities : — 

Spain : Near Burgos, 2. 

Switzebland : Lugano, Tioino, 1 (U.S.N.M.). 

Austbia-Hungaby : Bozen, Tirol, 3 (U.S.N.M.). 

Italy : Finalborgo, Liguria, 2 (Genoa) ; Isoverde, 1 (Genoa) : Vallom- 
brosa, Florence, 1 (U.S.N.M.); Borne, 4; Velletri, Borne, 15 (U.S.N.M.); 
Matera, Basilicata, 1 (Naples, type). 

Sabdinia: Gagliari, 5 (U.S.N.M.); Monte Gennargentu, 3 (U.S.N.M.) ; 
no exact locality, 3 (U.S.N.M.). 

Malta : El Ghain, 5 ; Bubato, 3 ; no exact locality, 4. 

Montenegbo : Beri, 4. 

Gbeece: Patras, 2; Corinth, 7 (U.S.N.M.); Nauplia, 1 (U.S.N.M.); 
Lamia, Thessaly, 1 (U.S.N.M.) 

Cbete : Labyrinth, 4. 

Tunis : No exact locality, 3 (U.S.N.M.). 

Remarks. — In general appearance Myotis oxygnaihus resembles 
M. myotis, though the colour perhaps averages somewhat darker. 
It is readily distinguishable, however, by its smaller skull 
(distinctly smaller head in spirit specimens), and shorter, narrower 
ears. In form the skull is like that of M. myotis ; and the 
teeth are not peculiar except for their small size, a character 
readily appreciable on comparison of the canines or of the crown 
area of upper molars. The range of Myotis oxygnaihus is, so far 
as known, strictly Mediterranean, probably coincident with that 
of M. capaccinii and Pipistrellus kuhlii. North of the Alps 
ilf. myotis occurs alone. 

9. Burgos, Spain. G. S. Miller (c). 8. 8. 4. 10. 

4 <$. Borne, Italy. (Goli.) G. Barrett-Hamilton 11. 1. 2. 22-25. 

(*)■ 

3 i, 2 9. El Ghain, Malta. Lord Lilford (p.) 11. 1. 1. 16-20. 

(Micallef.) 

3 6, 9. Bubato, Malta. Lord Lilford (p). 11. 1. 1. 21-24. 

(Micallef.) 

4. Malta. (Micallef.) Lord Lilford (p). 95. 3. 2. 3-6. 

2 (J, 2 9. Beri, 50 m. Montenegro. O. Thomas (p). 5.8.4.1-4. 

(L. Filhrer.) 

2 9. Patras, Greece. Hon. N. C. Boths- 8. 10. 2. 15-16. 
(C. Mottaz.) child (p). 

3 S, 9. Labyrinth, Crete. Miss T>. Bate (c). 5. 12. 2. 5-8. 

Genus PIPISTRELLUS Kaup. 

1829. Pipistrellus Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, 
I, p. 98 (pipistrellus). 

1837. Pipistrellus Bonaparte, Iconogr. Faun. Ital., I, fasc. sx. 

1838. Bomicia Gray, Mag. Zool. and Bot., ii, p. 495, February, 1838 

(calcarata = kuhlii). 

1839. Vesperugo Keyserling and Blasius, Wiegmann's Archiv fur Natur- 

gesch., 1839, i, p. 312 (part). 
1856. Hypsugo Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden, 
neue Folge, II, p. 131 (maurus and krascheninikowii). 



PIPISTiJELLUS 



203 



1856. Nannugo Kolenati, Allgom. deutsohe Naturhist. Zeitung, Dresden, 

neue Folge, n, p. 131 (nathusU, pipistrellus, and kuMU). 

1857. Vesperugo Blasius, Saugethiere TJeutschlands, p. 49 (part). 
1878. Vesperugo Dobson, Oatal. Chiropt. Brit. Mus., p. 183 (part). 

1897. Pipistrellus Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 384, 

October, 1897. 
1899. Euvesperugo Acloque, Faune de France, Mammiferes, p. 35 (part, 

included noctula, leisleri, maurus, kuMii, pipistrellus, and 

abramus). 
1907. PipistrelVus Miller, Families and Genera of Bats, p. 204, June 29, 

1907. 

Type species. — Vespertilio pipistrellus Schreber. 

Geographical distribution. — Entire mainland of Eastern Hemi- 
sphere to limits of tree growth, also Malay Archipelago, New- 
Guinea, Solomon Islands, and northern Australia ; in America 
from northern United States (except in boreal zone) to southern 
Mexico. 

Characters. — Like Eptesicus (p. 224), but with 2 — 2 upper 
premolars ; dental formula : i ||, c t|, pm ^|, « jt| = 34. 

Bemarhs. — The genus Pipistrellus is widely distributed in the 
warmer portions of the Old World and of North America. 
About forty species are now known, four of which occur in 
Europe. Externally these may often be confused with the smaller 
Myotis, though they may usually be recognized by a certain 
heaviness of form, and more especially by the shorter ear and 
less slender tragus. Though often regarded as nearly related to 
Nyctalus, on account of the similarity of dental formula, this 
genus is much the more primitive of the two, its members 
showing no tendency to modify the ordinary vespertilionine wing 
structure. It is in reality not much more than a sub-genus of 
Eptesicus (see remarks under the latter), though for the sake of 
convenience the two groups are best treated as distinct. 

KEY TO THE EUROPEAN SPECIES OF PIPISTRELLUS. 

Anterior upper premolar excessively minute, some- 
times hidden by the gum, its crown area much 
less than that of outer incisor ; anterior lower 
premolar with crown area less than half that 
of posterior premolar ; greatest width of tragus 
nearly equal to length of anterior border ; hairs 
of back usually with noticeably contrasted light 

tips P. savii, p. 219. 

Anterior upper premolar not excessively minute, 
never hidden by the gum, its crown area about 
equal to that of outer incisor ; anterior lower 
premolar with crown area more than half that 
of posterior premolar ; greatest width of tragus 
much less than length of anterior border ; hairs 
of back without noticeably contrasted light tips. 
Outer upper incisor less than half as high as 

inner ; large upper premolar almost or quite 

in contact with canine, the small premolar 

forced inward from tooth-row and scarcely or 

not visible from outer side P. kuhlii, p. 215. 



204 



CHIROPTERA 



Outer upper incisor more than half as high as 

inner; large upper premolar separated from 

canine by distinct space in which the small 

premolar is clearly visible from outer side. 

Lower canine robust, the length of base along 

cingulum about equal to length of anterior 

border of shaft; oondylobasal length of 

skull 11 to 12 mm. ; thumb short, its length 

about equal to width of wrist ; fifth finger 

about 40 mm P. pipistrellus, t>. 204. 

Lower canine slender, the length of base along 

cingulum slightly more than half length 

of anterior border of shaft ; oondylobasal 

length of skull 12-6 to 13-4 mm. ; thumb 

long, its length much greater than width 

of wrist; fifth finger about 46 mm P. nathusii, p. 213. 



PIPISTRELLUS pipistrellus Schreber. 

1774. Vespertilio pipistrellus Schreber, Saugthiere, I, pi. liv. Described, i, 

p. 167, 1775, under name Die Zwergfledermaus. (Prance, based 

primarily on Daubenton.) 
1776. VespertiUo pipistrelle P. L. S. Miiller, Natursyst. Suppl. u. Regist.- 

Band, p. 16 (Prance, based on Schreber). 
1825. Vespertilio pygmseus Leach, Zool. Journ., i, p. 560, January, 1825 

(Dartmoor, Devonshire, England). 
1834. VespertiUo brachyotos Baillon, Mem. Soc. Royale d'Emulation 

d' Abbeville, 1833, p. 50 (Abbeville, Somme, Prance). 
1839. ? [VespertiUo pipistrellus] var. nigra de Selys-Longchamps, Etudes 

de Micromamm., p. 140 (nomen nudum). 

1839. ? [Vespertilio pipistrellus'] var. rufescens de Selys-Longchamps, Etudes 

de Micromamm., p. 140 (nomen nudum). 

1840. V[espertilio] pusillus Schinz, Europ. Pauna, I, p. 9 (Synonym of 

pipistrellus ; Brehm cited as authority). 

1840. V[espertilio] melanopterus Schinz, Europ. Pauna, I, p. 9. Brehm 
cited as authority, but name apparently published here for first 
time (Rhentendorf, Thiiringen, Germany). 

1840. V[espertilio] stenotus Schinz, Europ. Pauna, ±, p. 9. Brehm cited as 
authority, but name apparently published here for first time 
(Rhentendorf, Thiiringen, Germany). 

1840. Vespertilio minutissimus Schinz, Europ. Pauna, i, p. 9 (Zurich, 
Switzerland). 

1845. P[ipistrellus] nigricans Bonaparte, Atti della sesta Riunione degli 
Scienziati Italiani, Milano, 1844, p. 340. Described but not named 
in Atti della seconda Riunione degli Scienziati Italiani, Torino, 
1840, p. 247, 1841. (Sardinia.) 

1845. Pipistrellus genei Bonaparte, Atti della sesta Riunione degli Scien- 
ziati Italiani, Milano, 1844, p. 340 (Alternative name for nigricans). 

1845. P[ipistrellus] typus Bonaparte, Atti della sesta Riunione degli Scien- 
ziati Italiani, Milano, 1844, p. 340 (Substitute for pipistrellus 
Schreber). 

1857. Vesperugo pipistrellus Blasius, Saugethiere Deutschlands, p. 61. 

1862. Vesperugo pipistrellus var. macrcpterus Jeitteles, Verhandl. der k. k. 

Zool. Bot. Gesellsch., Wien, hi, p. 250 (Kaschau, Hungary). 

1863. [Nannugo pipistrellus] var. typus Koch, Jahrb. des Vereins fur 

Naturkunde im Herzogthum Nassau, xvin, p. 490. Not of 
Bonaparte, 1845 (Wiesbaden, Hessen-Nassau, Germany). 



PIPISTRELLUS 205 

1863. [Nannugo pipistrellus] var. flavescens Koch, Jahrb. des Vereins fur 
Naturkunde im Herzogthum Nassau, xviii, p. 491 (Nassau, 
Germany). 

1863. [Nannugo pipistrellus] var. nigricans Koch, Jahrb. des Vereins fur 

Naturkunde im Herzogthum Nassau, xviii, p. 491. Not of 

Bonaparte, 1845 (Nassau, Germany). 
1863. [Nannugo pipistrellus'] var. Umbatus Koch, Jahrb. des Vereins fur 

Naturkunde im Herzogthum Nassau, xviii, p. 491 (Siegen, Nassau, 

Germany). 

1878. Vesperugo pipistrellus Dobson, Catal. Ghiropt. Brit. Mus., p. 223. 
1897. Pipistrellus pipistrellus Miller, Ann. and Mag. Nat. Hist., 6th ser., 
xx, p. 384, October, 1897. 

1904. Pipistrellus pipistrellus mediterraneus Cabrera, Mem. Soc. Espafi. 

Hist. Nat., II, p. 273 (Valencia, Spain). 
1910. Pipistrellus pipistrellus and P. pipistrellus mediterraneus Trouessart, 

Paune Mamm. d'Europe, pp. 14-15. 

Type locality. — France. 

Geographical distribution. — Europe from the Mediterranean 
north to Scotland and Scandinavia, west to Ireland and the 
Hebrides, east into Asia. 

Diagnosis.- — -Smallest European member of the genus (forearm, 
27-6 to 32 mm. ; condylobasal length of skull, 11 to 12 mm.) ; 
outer upper incisor more than half as high as inner incisor ; 
large upper premolar separated from canine by a distinct space, 
the small tooth visible from outer side, its crown area about 
equal to that of outer incisor ; anterior lower premolar with 
crown area equal to more than half that of succeeding tooth ; 
lower canine robust, the length of base along cingulum about 
equal to length of anterior border of shaft ; tragus with greatest 
width less than length of anterior border ; thumb short, its 
length about equal to width of wrist ; length of fifth finger about 
40 mm. ; posterior edge of wing membrane usually dark. 

External characters. — General form robust, the tail and legs 
rather short, the membranes relatively thick and opaque. Muzzle 
with very noticeable glandular swellings extending back to 
beneath eye. Ear extending about to nostril when laid forward, 
its general form rather short and broad, though with narrowly 
rounded tip ; anterior border abruptly convex at base, then 
essentially straight almost to tip ; posterior border faintly and 
irregularly concave above, evenly convex below, the antitragal 
lobe represented by a thickened ridge extending along margin of 
ear and turning abruptly inward without producing any notice- 
able break in outline of conch. Inner surface of conch slightly 
rugose, but without evident cross ridges. Tragus erect, scarcely 
half as high as conch, its tip broadly rounded, its greatest width 
(slightly above level of anterior base) about half length of anterior 
border ; except near tip, both borders are nearly straight or very 
slightly convex ; posterior basal lobe small but well defined. 
Wing rather narrow, with no special peculiarities of form, the third, 
fourth and fifth metacarpals sub-equal (fifth slightly shorter than 



206 



CHIEOPTBRA 



the others) and extending nearly to point of elbow ; fifth finger 
extending beyond elbow to a distance equal to less than one- 
third length of forearm ; thumb short, its length about equal to 
width of wrist ; membrane inserted at base of outer toe. Foot 
about half as long as the short, robust tibia ; calcar considerably 
longer than free border of interfemoral membrane, robust at base, 
but tapering rapidly and terminating without lobe, its keel well 
developed, with evenly convex margin. Tail about as long as 
body without head and 2^ times as long as tibia, the snort 
terminal vertebra free from membrane. 

Fur and colour. — The fur is closely confined to the body, 
showing no tendency to spread on membranes. On wing it 
extends, both above and below, to line joining knee and basal 
third of humerus ; lower surface of interfemoral membrane 
essentially naked except at extreme base, upper surface furred 
nearly to middle. Colour of upper parts a uniform brown, in 
most specimens nearly intermediate between the wood-brown and 
cinnamon of Ridway but sometimes darker, with a strong tinge 
of prouts-brown or raw umber, this especially noticeable in 
immature specimens, though occasionally evident in adults ; 
under parts essentially like back though slightly less dark ; hairs 
everywhere slaty brown at base, those of upper parts with tips 
darker than sub-terminal band, but not enough so to produce a 
definitely tricolor effect. Ears and membranes blackish. 

Skull. — Notwithstanding its small size, less than that of any 
other European bat, the skull is robust and heavily built as 
compared with that of the small species of Myotis. Dorsal profile 
rising gradually from nares to lambda, with slight concavity in 
interorbital region and slight convexity over middle of brain-case ; 
occipital region scarcely produced backward except for a median 
swelling between foramen magnum and lambda, 
on each side of which a condyle is just visible 
when skull is viewed from above ; ventral profile 
nearly flat except for a slight upward bend pos- 
teriorly. Brain-case ovate in general outline, 
its region of greatest breadth distinctly behind 
middle, its surface smooth or with faintly indi- 
cated sagittal crest and lateral portion of 
lambdoid crest ; greatest breadth of brain-case 
"fig 34 noticeably exceeding that of rostrum and slightly 

Pipistreiius pipi*- though evidently more than half greatest length 
trains. Nat. size, of skull ; floor of brain-case flat, without vacui- 
ties ; a distinct groove between cochlea and 
median portion of floor, this groove bounded antero-externally 
by a slight though usually evident longitudinal ridge ; auditory 
bulla? moderately large, not peculiar in form ; interorbital region 
broadly hour-glass shaped, its least breadth about equal to 
breadth across roots of canines ; between constriction and ante- 
orbital foramen the orbital margin is slightly but evidently 




PIPISTEELLUS 207 

inflated, the inflated region with a median angle suggesting 
a rudimentary postorbital process ; rostrum short and broad, 
narrowing gradually in front, a slight concavity at each side 
bordering lachrymal inflation, and an evident median longi- 
tudinal groove, most noticeable posteriorly ; nasal emargina- 
tion slightly deeper than wide, extending less than half way to 
interorbital constriction ; anteorbital foramen small, over point of 
contact between large premolar and first molar ; palate broad, 
distinctly concave both longitudinally and laterally ; anterior 
emargination small, wider than deep, its posterior border on line 
with posterior edge of canine ; mesopterygoid fossa squarish, 
encroached on anteriorly by broadly triangular median palatal 
spine ; hamulars slightly turned inward. Mandible robust, the 
ramus much deeper at symphysis than behind tooth-row, the 
coronoid process so low that upper edge of posterior portion of 
mandible is squarely and horizontally truncate, parallel with 
alveolar line ; angular process short but well developed, on level 
with alveolar line, its extremity slightly bent inward. 

Teeth. — Relatively to size of skull the teeth are rather large 
and robust, though inclined to be low, tendencies especially 
noticeable in the canines. Inner upper incisor robust, its shaft 
nearly half as high as that of canine, and directed strongly 
forward and slightly inward, its crown irregularly elliptical-oval 
in outline, with main axis nearly in line of tooth-row ; secondary 
cusp large and conspicuous, about half as high as main shaft, 
from the postero-external surface of which it projects ; cingulum 
well developed, often forming a minute postero-basal cusp. Outer 
upper incisor slightly but evidently smaller than inner, its shaft 
more than half as high as that of inner, to secondary cusp of 
which its extremity is closely approximated ; crown outline 
essentially as in inner tooth but main axis lying at right angles 
to tooth-row ; posterior surface of shaft broadly concave ; inner 
margin with small though distinct secondary cusp ; cingulum 
moderately well doveloped. The main cusps of the two teeth lie 
in line of general curve of anterior portion of tooth-row. Space 
between outer incisor and canine about equal to greatest diameter 
of incisor. Lower incisors forming a continuous, broadly U -shaped 
row between canines, their crowns very slightly imbricated ; 
crowns much longer than high, trifid, that of i x narrowest, 
longest and lowest, that of i 2 and i 3 widened posteriorly but 
without additional cusps or tubercles. Upper canine robust, 
the greatest diameter of its crown about three-quarters length 
of anterior border of shaft, the cross section of shaft broadly 
triangular with longest side formed by nearly flat postero-internal 
surface ; a sharply defined antero-exteral longitudinal groove, and 
less definite postero-external concavity ; anterior edge narrow 
but not strictly trenchant ; posterior edge trenchant, with well 
marked angle slightly below middle, this angle frequently becom- 
ing a distinct secondary cusp ; cingulum well developed but not 



208 CHIEOPTBRA 

forming true basal cusps. Mandibular canine low and heavy, 
its apex scarcely rising above level of highest molar cusps, its 
greatest diameter measured along cingulum nearly or quite equal 
to length of anterior border of shaft ; cingulum well developed, 
forming a distinct antero-basal cusp, the apex of which rises to 
level of middle of posterior border of shaft. Anterior upper 
premolar with area of crown approximately equal to that of upper 
incisor and about one-fifth that of canine. It is somewhat 
crowded inward from tooth-row, though about half of its crown is 
visible from outer side in space between canine and large 
premolar ; main cusp short but well developed, lying somewhat 
in front of middle of crown, the general form of the tooth much 
like that of canine but proportionally lower. Large upper 

premolar with crown area about equal to that 

) of canine or slightly greater, the inner portion 

/?V^ ^ narrow and flattened-concave, with evident 

-^?T\l elevated rim, the posterior border strongly 

Cx 3 . „ f\ concave, the anterior border usually convex 

30 but occasionally a little concave ; height of 

main cusp slightly greater than that of highest 
fig. 35. molar cusps and about equal to length of 

*K£Mh x ""*■ tooth along outer cingulum, posterior cutting 

edge well developed ; secondary cusp low but 
evident, rising from cingulum at antero-internal base of main 
cusp. Lower premolars with crown areas not conspicuously 
unequal, though that of second perceptibly greater than that of 
first ; outline of crown of each tooth rhombic, the outer border 
somewhat convex, the anterior border of second relatively shorter 
than that of first ; cingulum well developed, forming a slight 
antero-internal basal cusp ; main cusp triangular in outline when 
viewed from the side, that of second as high as molar cusps, that 
of first shorter, the antero-external surface of each tooth convex, 
the internal and posterior surfaces concave. First and second 
upper molars sub-equal, though transverse diameter is relatively 
greater in latter than in former ; inner border rather narrowly 
rounded, the region of greatest convexity a little in front of 
middle ; anterior and posterior borders straight or slightly 
concave ; protocone robust though rather low ; hypocone small 
but well developed, though not completely distinct from posterior 
commissure of protocone ; metacone higher than paracone ; styles 
well developed ; VV-p a ttern normal ; m 3 with crown area about 
two-thirds that of m 1 , the hypocone absent, the metacone smaller 
than paracone ; no trace of metastyle or fourth commissure. 
Lower molars with no special peculiarities ; protoconid higher 
than hypoconid in all three teeth ; hypoconid with greater basal 
area than protoconid in m. 1 and m 2 , but with less in m 3 ; cingulum 
well developed, forming a slight postero-internal cusp behind 
entoconid. 

Measurements. — Adult male from Henley-on-Thames, Oxford- 



PIPISTRELLUS 



209 



shire, England : head and body, 44 ; tail, 32 ; tibia, 10- 4; foot, 
6 ; forearm, 30 • 4 ; thumb, 4 • 4 ; third finger, 52 ; fifth finger, 
38. Two adult males from Sorrento, Italy : head and body, 42 
and 43 ; tail, 29 and 32 ; tibia, 10 ■ 8 and 10 ; foot, 5 ■ 8 and 6 • 2 ; 
forearm, 30-4 and 31 ; thumb, 4 - 8 and 4-2; third finger, 53 
and 53; fifth finger, 39 and 38; ear from meatus, ll - 4 and 
11-4; width of ear, 8*2 and 8 - 2. Two adult females from the 
same locality : head and body, 39 and 40 ; tail 32 and 33 ; tibia, 
10-8 and 10' 6 ; foot, 6 and 6 ; forearm, 32 and 30-2 ; thumb, 
5 and 4-4 ; third finger, 58 and 52 ; fifth finger, 41-6 and 40 ; 
ear from meatus, 12 and 11 - 2 ; width of ear, 8-4 and 8. Adult 
female from Burgos, Spain, and adult female from Ciudad Real, 
Spain : head and body, 49 and 39 ; tail, 31 and 30; tibia, 10 '6 
and 9-6 ; foot, 4-8 and 5-6 ; forearm, 32 and 28-8 ; thumb, 5 
and 4 ■ 6 ; third finger, 57 ■ 6 and 51 ; fifth finger, 42 and 38 ; ear 
from meatus, 10 ■ 4 and 10 • 4. Extremes of twenty males from 
Florence, Italy : head and body, 33-38 ; tail, 26-31 ; tibia, 9- 2- 
9-6; foot, 5-0-5-2; forearm, 27 • 6-30 ; thumb, 5 ■ 0-5 • ; third 
finger, 49-53 ; fifth finger, 34 • 6-40. For cranial measurements 
see Table, p. 210. 

Specimens examined. — Two hundred and seventy -nine, from the following 
localities :— 

Ireland : Co. Longford, 1 ; Co. Antrim, 1. 

England : Alnwick, Northumberland, 1 ; Bowdon, Cheshire, 1 ; Great 
Grimsby, Lincolnshire, 1; Henley-on-Thames, Oxfordshire, 1 (U.S.N.M.) ; 
Tring, Hertfordshire, 4 : Lilford, Northamptonshire, 2 ; Aberia, Merioneth- 
shire, 1 ; Chelmsford, Essex, 2 ; London, 1 ; Wimbledon, Surrey, 1 ; Twig- 
worth, Gloucestershire, 1 ; New Forest, Hampshire, 1 ; Netley, Hamp- 
shire, 1 ; Loddiswell, Devonshire, 1. 

Sweden : Upsala, 6. 

Denmark : Hillerod, Zealand, 1. 

France : Boulogne -sur-Mer, Pas-de-Calais, 2 ; Etupes, Doubs, 1 
(Mottaz) ; Nimes, Gard, 3 (Mottaz and Nlmes, the last wrongly marked 
type of migrans Crespon) ; St. Genies, Gard, 1 (Mottaz) ; Marseilles, 1 
(U.S.N.M.) 

Germany : Bonn, 4 ; Ingelheim, Bheinhessen, 1 ; Magdeburg, Saxony, 1 ; 
Berlin, 1 ; Budolstadt, Bavaria, 3 ; Niesky, Silesia, 1. 

Austria-Hungary : Transylvania, 1 ; Zara, Dalmatia, 1. 

Switzerland : Geneva, 15 (Mottaz) ; Buchillon, Vaud, 4 (Mottaz) ; 
Morat, Pribourg, 1 (Mottaz); Neuchatel, 1 (Mottaz); Cortivallo, Ticino, 
1 (U.S.N.M.) ; Mt. San Salvatore, Ticino, 13 (U.S.N.M.). 

Italy : Campiglio, Tirol, 1 ; near Genoa, 10 (U.S.N.M. and Genoa) ; 
Isola Giglio, 2 (Genoa) ; Florence, 103 (U.S.N.M. and Mottaz) ; Rome, 2 
(U.S.N.M.); Sorrento, 21 (U.S.N.M.); Mondulo, Sicily, 1 (U.S.N.M.); 
Palermo, Sicily, 1 (U.S.N.M.) ; Sicily, no exact locality, 1 ; Ustica Island, 1 
(U.S.N.M.). K ' . 

Sardinia : Cagliari, 17 (B.M. and Genoa) ; no exact locality, 2. 

Greece : Athens, 1 (U.S.N.M.) ; Tatoi, near Athens, 11 ; Kephissia, 
near Athens, 4. 

Spain : Villalba, Lugo, 1 ; Burgos, 3 ; Silos, Burgos, 2 ; La Granja, 
Segovia, 4 ; Ciudad Seal, Madrid, 2 ; Granada, 1 ; Seville, 1 ; Alcala, near 
Seville, 1. , 

Remarks. — Pipistrellus pipistrellus is the smallest as well as 
one of the commonest and most generally distributed of European 



210 



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212 



OHIEOPTBKA 



bats. Superficially it may be distinguished from the almost 
equally small Myotis mystacinus by its smaller ears and shorter 
legs ; but for positive discrimination from the members of the 
genus Pipistrellus recourse to the more technical characters of 
skull and teeth is necessary. On superficial examination it 
may usually be recognized among its congeners by its small size, 
and by the shortness of the fifth finger. 



<5al. 


Go. Longford, Ireland. 


Dr. G. E. Dobson (p). 


76. 2. 12. 1. 


9 st. 


Co. Antrim. 


Hon. N. C. Rothschild (p). 


1. 9. 3. 6. 


?. 


Alnwick, Northum- 
berland, England. 


W. E. de Winton (p). 


11. 1. 3. 390. 


2 6, 2 9. 


Tring, Hertfordshire. 


Hon. N. C. Rothschild (r). 


9. 2. 19. 1-4. 


9. 


Lilf ord, Northampton- 
shire. 


■ Lord Lilf ord (p). 


11. 1. 1. 125. 


2 9. 


Chelmsford, Essex. 


M. Christy and E. L. 
Thompson (p). 


11. 1. 3. 23-24. 


9 juv. al. 


London. 


Dr. A. Giinther (p). 


74. 7. 6. 1. 


S. 


Wimbledon, Surrey. 


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


11. 1. 3. 27. 


juv. 


New Forest, Hamp- 
shire. 


Col. J. W. Yerbury (c & p). 


11. 1. 3. 389. 


Sal. 


Netley, Hampshire. 


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


76. 11. 3. 1, 


9. 


Loddiswell, Devon- 
shire. 


Col. J. W. Yerbury (c&p). 


11. 1. 3. 25. 


3 9. 


Upsala, Sweden. 
(Kolthoff.) 


Lord Lilf ord (p). 


11. 1. 1. 26-28. 


S. 


Hillerod, Zealand, 
Denmark. 


0. Thomas (c & p). 


96. 6. 7. 1. 


,5,9. 


Boulogne, Pas - de - 
Calais, France. 


0. Thomas (c & p). 


98. 1. 9. 1-2. 


S, 9 al. 


, Bonn, Rhineland, 


Dr. A. Giinther (p). 








Germany. 






9. 


Ingelheim, Rhein- 
hessen. 


C. H. Hilgert (c). 


8. 11. 2. 3-4. 


Sal. 


Magdeburg, Saxony. 


Dr. W. Wolterstorff (p). 


92. 12. 1. 2. 


dal. 


Berlin. 


Dr. A. Giinther (p). 

C. G. Danford and J. A. 


66. 2. 1. 22. 


Sal. 


Transylvania, Hun- 


74. 7. 4. 6. 




gary- 


Brown (c & p). 




i. 


Zara, Dalmatia. 


Lord Lilf ord (p). 


11. 1. 1. 29. 


2al. 


Ciudad Real, Spain. 


A. Cabrera (p). 


8. 7. 23. 2-3. 


$. 


Burgos, Prov. Burgos 


. G. S. Miller (c). 


8. 8. 4. 14. 


S. 


Silos, Burgos. 


G. S. Miller (c). 


8. 8. 4. 15. 


lal. 


Campiglio, Tirol, 
Italy. 


G. C. Champion (c & p). 


96. 8. 7. 1. 


6 juv. al 


. Sicily. 




46. 6. 15. 6. 




Ual. 


Aristano, Cagliari, 
Sardinia. (C. 
Krausse.) 


Hon. N. C. Rothschild (p) 


. 7. 5. 24. 11-24. 


2 juv. al 


. Sardinia. (P.Bonomi. 


) E. N. Buxton (p). 


95. 4. 16. 4-5. 


3$. 


Tatoi, Athens, Greece. 
(C. Mottcus.) 


, Hon. N.C. Rothschild (p). 


8. 10. 2. 19-21. 


$, 8 9. 


Kephissia, Athens. 


C. Mottaz (c). 


8. 11. 3. 4-7. 



PIPISTRELLUS 213 



PIPISTRELLUS NATHUSH Keyserling and Blasius. 

1839. V[espertilio~] nathusii Keyserling and Blasius, Wiegmann's Archiv 

fur Naturgesch., 1839, I, p. 320 (Berlin, Germany). 
1857. Vesperugo nathusii Blasius, Saugethiere Deutschlands, p. 58. 
1878. Vesperugo abramus Dobson, Catal. Ohiropt. Brit. Mus., p. 226 (Part : 

not of Temminck). 
1900. Pipistrellus nathusii Mehely, Monogr. Chiropt. Hungarise, p. 276. 
1905. Vesp[erugo] nathusii var. unicolor Fatio, Arch. Soi. Phys. et Nat., 

Geneve, 4th ser., xrx, p. 510, May, 1905 (Geneva, Switzerland). 

Type in Geneva Museum. 
1910. Pipistrellus abramus Trouessart, Paune Mamm. d'Europe, p. 16. 

Type locality. — Berlin, Germany. 

Geographical distribution. — Central and southern Continental 
Europe ; exact limits of range not known. 

Diagnosis. — Not so small as Pipistrellus pipistrellus (forearm, 
32 to 35 mm. ; condylobasal length of skull, 12-6 to 13 • 4 mm.), 
which in general it resembles, but : small upper premolar better 
developed, the greatest diameter of its crown nearly half that of 
canine ; canines both 'above and below much more slender, the 
length of base of lower tooth measured along cingulum slightly 
more than half length of anterior border ; tragus more slender, 
its greatest width much less than length of anterior border ; 
thumb long, its length much greater than width of wrist ; length 
of fifth finger about 46 mm. ; posterior edge of wing membrane 
always pale, though never sharply defined white. 

External characters. — In general and apart from the animal's 
less diminutive size, the external characters are essentially as in 
Pipistrellus pipistrellus. Ear larger and broader with more 
obtuse apex, more evidently concave posterior border, the inner 
surface of conch more rugose and with about four irregular cross 
striations behind tragus ; antitragus small but well defined, 
projecting distinctly beyond border of conch ; tragus about as 
high as in P. pipistrellus and similarly blunt at tip, but with 
posterior border more evidently convex ; posterior basal lobe 
small, usually less well defined than in P. pipistrellus. Wing 
larger and relatively broader than in P. pipistrellus, the meta- 
carpals as in the smaller animal, but fifth finger extending beyond 
elbow to a distance equal to decidedly more than one-third length 
of forearm ; thumb less shortened than in the other European 
members of the genus, its length noticeably greater than width 
of wrist ; membrane inserted at base of outer toe. Foot, calcar 
and tail as in P. pipistrellus. 

Fur and colour. — Fur slightly more loose in texture than that 
of P. pipistrellus, the individual hairs somewhat longer, those at 
middle of back about 7 mm. in length. In distribution it shows 
no peculiarities, though it extends perhaps less widely on dorsal 
surface of interfemoral membrane. Colour essentially like that 
of Pipistrellus pipistrellus, though usually distinguishable by a 



214 



CHIROPTEEA 



tendency away from the cinnamon and raw-umber tints toward 
a clearer brown more resembling Ridgway's mars-brown. Mem- 
branes less blackish than in P. pipistrellus, the wing from foot 
nearly to fifth finger with a noticeable pale border about 1 mm. 
in width, similar to that present in P. huhlii, but less sharply 
denned and less nearly white. 

Skull. — The skull is less diminutive than that of Pipistrellus 
pipistrellus, its general size about as Myotis mystacinus. General 
form less robust than in P. pipistrellus, the width of brain-case 
barely one-half greatest length, but more contrasted with that of 
rostrum. Dorsal profile as in the smaller species, but with more 
evident anterior concavity and posterior convexity, the anterior 
edge of interparietal indicated by a slight transverse constriction. 
Other details of form essentially as in P. pipistrellus. 

Teeth. — As compared with those of Pipistrellus pipistrellus the 
teeth throughout show a tendency toward slenderness and height. 
Inner upper incisor noticeably more slender than that of P. 
pipistrellus, and with less well developed secondary cuspj outer 
upper incisor distinctly larger than inner, its apex extending 
noticeably beyond secondary cusp of inner tooth, its general form 
essentially as in P. pipistrellus, but inner margin without evident 
secondary cusp. Lower incisors less crowded than in P. pipis- 
trellus, a slight space usually present in 
median line, another between i 2 and i s , and 
another between i s and canine ; outer edge 
of i x slightly overlapping « 2 ; in form the 
teeth are not peculiar. Upper canine like 
that of P. pipistrellus, except that the 
greatest diameter of its crown is only about 
half length of anterior border of shaft. 
Fig. 36. Mandibular canine high and slender, its apex 

Pipistrellus nathusii. rising distinctly above that of highest molar 
Anterior teeth x 5. cusps, its greatest diameter measured along 
cingulum equal to a little more than half 
length of anterior border ; apex of anterior cingulum cusp not 
rising above level of basal third of posterior border. Upper 
premolars as in P. pipistrellus, except that the small tooth is 
relatively higher and more perfectly in the tooth-row, and the 
posterior border of its shaft usually shows some indication of an 
angular secondary cusp corresponding to that of canine. Lower 
premolars with crown area more nearly equal than in the 
smaller species, but without special peculiarities of form. Molars 
both above and below essentially similar to those of P. pipistrellus. 
Measurements. — Adult male from Berlin, Germany (topotype) : 
head and body, 45 ; tail, 35-4 ; tibia, 13 ; foot, 6'8 ; forearm, 
33; thumb, 5-2; third finger, 65; fifth finger, 47; ear from 
meatus, 1 2 ; width of ear, 1 1 . Average and extremes of six 
adults from Buchillon, Vaud, Switzerland : tibia, 1 2 • 9 ( 1 2 • 6-1 4) ; 
foot, 7-3 (6-8-7-6); forearm, 33 (32-4-34-6); thumb, 5-7 




PIPISTRELLUS 215 

(5-2-6); third finger, 61-3 (58-65); fifth finger, 44-6 (41-47). 
Adult male and female from Florence, Italy : head and body, 
46 and 47; tail, 40 and 38; tibia, 12-8 and 12-6; foot, 6-8 
and 7 ; forearm, 35 and 33 ; thumb, 6 • 2 and 6 • 6 ; third finger, 
63 and 62 ; fifth finger, 47 and 46 ; ear from meatus, 12-6 and 
12-6 ; width of ear, 11 and 11-4. For cranial measurements see 
Table, p. 222. 

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

Fbance : St. Gilles, Gard, 1. 

Gebmany: Berlin, 1; Bavaria, 1 (U.S.N.M.); Ingelheim, Bhein- 
hessen, 1. 

Switzerland: Geneva, 6 (Mottaz and Geneva, including type of 
unicolor Patio) ; Montreux, Vaud, 1 (Mottaz) ; Buchillon, Vaud, 6 (Mottaz) ; 
Neuchatel, 1 (U.S.N.M.) ; Canton TJri, 1 ; St. Gothard, Uri, 1 (U.S.N.M.). 

Aostbia-Hungary : Palics, Bacser, southern Hungary, 2. 

Italy: Siena, 1 (U.S.N.M.); Florence, 3 (U.S.N.M.); Rome, 5 (B.M. 
and U.S.N.M.) ; Borzoli, Liguria, 1 ; Catanzaro, Calabria, 1 (U.S.N.M.). 

Remarks. — Though readily distinguishable from the other 
European members of the genus by its cranial and dental 
characters, Pipistrellus nathusii is superficially much like P. pipis- 
trellus. It is usually recognizable, however, by its slightly less 
diminutive size, more robust form, and by the constant presence 
of an ill-defined light (though never actually whitish) border to 
the wing.* As pointed out by Mehely in 1900 it has no very 
near relationship to the Oriental P. abramus. 

6. St. Gilles, Gard, France. G. S. Miller (c). 8. 8. 4. 128. 

6 al. Berlin, Germany. Dr. Giinther (c & p). 66. 2. 1. 22. 

9. Ingelheim, Bheinhessen. C. Hilgert (c). 8. 11. 2. 3. 

2 al. Palics, Bacser, Hungary. Budapest Museum (b). 0. 4. 9. 1-2. 

<5. Canton Uri, Switzerland. Tomes Collection. 7. 1. 1. 398. 

i, 9. Rome. (C. Coli.) G. Barrett-Hamilton (p). 11. 1. 2. 20-21. 

9. Borzoli, Liguria, Italy. Marquis G. Doria (c & p). 5. 12. 15. 7. 

PIPISTRELLUS KOHLII Kuhl. 

1819. Vespertilio kuhlii Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv 

(= Neue Ann., i), pt. 2, p. 199 (Triest). 
1835. Vesp[ertilio~] alboUmbatus Kiister, Isis, p. 75 (Cagliari, Sardinia). 
1837. Vespertilio vispistrelhis Bonaparte, Iconogr. Faun. Ital., ±, fasc. xx 

(near Borne, Italy). Type in British Museum. 

1837. Vespertilio alcythoe Bonaparte, Iconogr. Faun. Ital., I, fasc. xxi 

(Sicily.) Type in British Museum. 

1838. Bomicia calcarata Gray, Mag. Zool. and Bot., n, p. 495 (locality 

unknown). 
1841. Pipistrellus marginatus Bonaparte, Iconogr. Fauna. Ital., Indie. 

distrib., nomencl. mod. (Substitute for alboUmbatus). 
1844. Vespertilio marginatus Wagner, Schreber's Saugthiere, Suppl., i, 

p. 503, pi. lv A. No description. Name occurs in synonymy of 

kuhlii with Michahelles as authority, and on plate. Apparently 

not previously published. 



* A light border sometimes occurs in P. pipistrellus, but is rare. That 
constantly present in P. kuhlii is more sharply defined and more truly 
whitish than in P. nathusii. 



216 



CHIROPTEKA 



1840. V[espertilio] Ursula Wagner, Schreber's Saugthiere, Suppl., i, p. 505 

(Morea, Greece). 
1857. Vesperugo kuhtti Blasius, Saugethiere Deutschlands, p. 63. 
1878. Vesperugo kuhlii Dobson, Catal. Chiropt. Brit. Mus., p. 230. 
1886. [Vesperugo kuhlii] var. albicans Montioelli, Atti Soo. Ital. Soi. Nat., 

Milano, xxvu, p. 200, March, 1886 (Caivano, Naples, Italy). 
1886. [Vesperugo kuhlii] var. pullatus Monticelli, Atti Soc. Ital. Sci. Nat., 

Milano, xxvn, p. 200, March, 1886 (Bella Vista, near Portici, 

Naples, Italy). 
1900. Pipistrellus kuhlii M6hely, Monogr. Chiropt. Hungariae, p. 261. 
1910. Pipistrellus kuhli Trouessart, Faune Mamm. d'Europe, p. 17. 

Type locality. — Trieste, Austria-Hungary. 

Geographical distribution. — Mediterranean region and eastward 
into Asia. 

Diagnosis. — Size about as in Pipistrellus nathusii (forearm, 31 
to 35 mm.; condylobasal length of skull, 12-0 to 13-2 mm.); 
outer upper incisor less than half as high as inner incisor ; large 
upper premolar almost or quite in contact with canine, the small 
premolar forced inward from tooth-row and scarcely or not visible 
from outer side, its greatest diameter about equal to that of 
outer incisor ; canines less robust than in P. pipistrellus, less 
slender than in P. nathusii ; tragus with greatest width less than 
length of anterior border ; thumb short, its length about equal to 
width of wrist ; posterior edge of wing membrane with sharply 
denned whitish border. 

External characters. — General form very similar to that of 
Pipistrellus pipistrellus, the wing similarly narrow as compared 
with that of P. nathusii. Ear narrowly rounded at tip, the posterior 
border slightly concave ; antitragus slightly developed, producing 
an evident break in contour of conch; inner surface of conch 
somewhat rugose, without well defined transverse striations ; 
tragus essentially as in P. nathusii, the posterior border noticeably 
convex. Wing, foot, calcar and tail as in P. pipistrellus. 

Fur and colour. — Quality and distribution of fur essentially 
as in Pipistrellus pipistrellus, but dorsal surface of interfemoral 
membrane haired scarcely beyond basal third. Colour not very 
different from that of Pipistrellus pipistrellus, but somewhat 
lighter and more yellow, often approaching raw-siena. Mem- 
branes blackish, the wing between foot and fifth finger with a 
sharply defined very narrow (less than 1 mm.) nearly white 
border. 

Shull. — The skull resembles that of Pipistrellus nathusii in 
size, but its form is even more robust than that of P. pipistrellus. 
Dorsal profile with very slight interorbital concavity and barely 
perceptible convexity over middle of brain-case. Breadth of 
brain-case about half greatest length of skull. Dorsal surface 
of rostrum less rounded off at sides than in P. pipistrellus and 
P. nathusii, but not so much flattened as in P. savii. Narial 
emargination more abruptly narrowed posteriorly than in the 



PIPISTRELLUS 



217 




other European species. Mesopterygoid space slightly longer 
than wide. Mandible with coronoid process distinctly higher 
than articular process, so that upper edge of posterior portion is 
oblique and not parallel with alveolar line. 

Teeth. — Inner upper incisor essentially as in Pipistrellus 
■pipistrellus except that secondary cusp is reduced to a minute, 
sometimes obsolete, projection from cingulum at posterior base of 
shaft ; outer incisor very small, less than half as high as inner, 
its apex about on level with highest point of cingulum of larger 
tooth, its crown area about two-thirds that of latter, its structure 
essentially as in P. pipistrellus, though with very small secondary 
cusp ; small tooth situated directly exterior to 
large, so that a line perpendicular to main 
axis of skull would pass through middle of 
all four incisors ; lower incisors essentially as 
in P. pipistrellus, though relatively larger and 
more strongly imbricated. Canines both above 
and below intermediate in form between those 
of P. pipistrellus and P. naihusii. Anterior , . ... , 

upper premolar crowded inward from tooth- 
row and closely wedged between canine and H , |( , 
large premolar which are nearly or quite in Pivistre uZkubZn. 
contact ; the small tooth is usually though not Anterior teeth x 5. 
always invisible from outer side, its cusp is 
very low, nearly terete, its crown area about equal to that 
of outer incisor ; large premolar with no special peculiarities. 
Lower premolars as in the related species, but disproportion in 
size more marked, the crown area of first a little more than 
half that of second. Molars both above and below essentially 
as in P. pipistrellus and P. naihusii, but somewhat more robust. 

Measurements. — Two adult males from near Genoa, Italy : 
head and body, 43 and 46 ; tail, 37 '4 and 38 ; tibia, 12-4 and 
13 ; foot, 6 and 6 ; forearm, 33 • 6 and 34 ; thumb, 5 and 5-2 ; 
third finger, 60 and 60 ; fifth finger, 45 and 44 ; ear from meatus, 
12 6 and 13; width of ear, 10 '4 and 10. Two adult females 
from the same locality : head and body, 44 and 47 ; tail, 39 and 
40 ; tibia, 1 2 and 13 ; foot, 6 • 2 and 6 • 2 ; forearm, 33 • 4 and 35 ; 
thumb, 5 - 2 and 5 - 4; third finger, 60 and 63 ; fifth finger, 45 
and 47 ; ear from meatus, 13 and 13 ; width of ear, 10 and 10. 
Adult male and female from Palermo, Sicily : head and body, 
42 and 44; tail, 36 and 35-4; tibia, 12-4 and 12-4 : foot, 5-6 
and 5 • 4 ; forearm, 31 and 34 ; thumb, 5 ■ 4 and 5 • 2 ; third finger, 
56 and 61 ; fifth finger, 41 and 43 • 6 ; ear from meatus, 12- 4 and 
13 ; width of ear, 10 and 10. Adult male and female from 
Cagliari, Sardinia : head and body, 44 and 45 ; tail, 35 and 37 ; 
tibia, 12-6 and 12-4; foot, 6-2 and 6-8; forearm, 33 and 33; 
thumb, 5 and 5 ; third finger, 58 and 62 ; fifth finger, 43 and 45 ; 
ear from meatus, 13 and 12 ; width of ear, 10 and 10. For 
cranial measurements see Table, p. 218. 



218 



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PIPISTRELLUS 



219 



Specimens examined. — One hundred and eight, from the following 
localities : — 

France : St. Gilles, Gard, 2 ; Nimes, Gard, 3 (Mottaz) ; St. Genies, 
Gard, 4 (Mottaz) ; Marseilles, 1 (U.S.N.M.). 

Switzerland : Coremmo, Tioino, 1 (Mottaz) ; Lugano, Ticino, 1 (Mottaz). 

Italy: Near Genoa, 14 (U.S.N.M. and Genoa); Siena, 6 (B.M. and 
U.S.N.M.); Florence, 1 (U.S.N.M.); Rome, 6 (B.M. and U.S.N.M.); 
near Rome 1 (type of vispistrellus Bonaparte) ; Sorrento, 3 (U.S.N.M. 
Catanzaro, Calabria, 6 (U.S.N.M.) ; Palermo, Sicily, 23 
Corleone, Sicily, 1 (U.S.N.M.) ; Ustica Island, Sicily, 4 
Sicily, 1 (type of alcythoe Bonaparte). ' 

Sardinia : Gagliari, 4 (U.S.N.M.); Mt. Gennargentu, 7 (U.S.N.M.); 
no exact locality, 2. 

Greece : Corfu, 2 ; Cephalonia, 10 ; Patras, 3. 

Spain : San Cristobal, Minorca, Balearic Islands, 2. 



IS: 



S.N.M.) 
S.N.M.) 



Bemarhs. — Pipistrellus Ttuhlii is easily recognizable by tbe 
form and relative size of tbe upper incisors. Externally it may 
usually be known by tbe sharply denned wbitisb border to tbe 
wing membrane, tbough too mucb reliance sbould not be placed 
on this character alone. Many specimens from Sardinia are 
lighter in colour than those from the mainland. These represent 
the albolimbatus of Kiister. Normally coloured examples also 
occur ; and in the absence of adequate material it has seemed 
preferable for the time being not to attempt to define the insular 
form. 



6. St. Gilles, Gard, 

France. 
i. Siena, Italy. 

(S. Brogi.) 
3 6, 9. Rome. (C. Coli.) 

skeleton with- Sicily. (Prince Bona- 

out skull. parte.) 

skeleton with- Near Rome, Italy, 
out skull. (Prince Bonaparte.) 

S, ?. Corfu, Greece. 

(G. Mottaz.) 
3 <5. Argostoli, Cephalonia. 

(C. Mottaz.) 
Patras. (C. Mottaz.) 



<5, ?. 
2 S. 



San Cristobal, 
Minorca, Balearic 
Islands. 



G. S. Miller (c). 8. 8. 4. 127. 

Dr. E. Hamilton (p). 98. 10. 2. 2. 

G. Barrett-Hamilton 11. 1. 2. 16-1P. 

Tomes Collection. 7. 1. 1. 730. 

(Type of V. alcythoe Bonaparte.) 
Tomes Collection. 7. 1. 1. 729. 

(Type of V. vispistellus Bonaparte.) 
J. I. S. Whitaker (p). 8. 10. 1. 4-5. 

J. I. S. Whitaker (p). 8. 10. 1. 1-3! 

Hon. N. C. Rothschild 8. 10. 2. 17-18. 

O. Thomas and R. I. 0. 7. 1. 29-30. 
Pocock (c & p). 



PIPISTRELLUS SAVH Bonaparte. 

1837. Vespertilio savii Bonaparte, Iconogr. Faun. Ital., i, fasc. xx (Pisa). 

Type in British Museum. 
1837. VespertiUo aristippe Bonaparte, Iconogr. Faun. Ital., I, fasc. xxi 

(Sicily). 

1837. Vespertilio leucippe Bonaparte, Iconogr. Faun. Ital., i, fasc. xxi 

(Sicily). Type in British Museum. 

1838. Vespertilio bonapartii Savi, Nuovo Giorn. de' Letterati, Pisa, xxxvn, 

p. 226 (Tuscany). 



220 



CHIEOPTBBA 



1844. Vesp[ertilio] nigrcms Greapon, Paune Meridionale, i, p. 24 (Nimes, 

Gard, Prance). 
1853. Vesperugo maurus Blasius, Wiegmann's Arohiv fur Naturgesch., 

1853, 1, p. 35 (Central chain of the Alps). 
1857. Vesperugo maurus Blasius, Saugethiere Deutschlands, p. 67. 
1872. V[espertilio] agilis Patio, Paune Vert. Suisse, Append, au vol. I, 

p. iii (Alternative name for V. savii Bonaparte, ex Savi MS.). 
1878. Vesperugo maurus Dobson, Catal. Chiropt. Brit. Mus., p. 218. 
1904. Vespertilio ochromixtus Cabrera, Mem. Soc. Espafi. Hist. Nat., n, 

p. 267 (Sierra de Guadarrama, Madrid, Spain). 
1910. Pipistrellus savii and P. savii ochromixtus Trouessart, Paune Mamm. 

d'Europe, pp. 13-14. 

Type locality. — Pisa, Italy. 

Geographical distribution. — Southern Europe, west to the 
Iberian Peninsula, north to the Alps ; also the Canary Islands, 
northern Africa and southern Asia. Limits of range very 
imperfectly known. 

Diagnosis. — Largest European member of the genus (condy- 
lobasal length of skull, 13 to 14 mm. ; forearm, 31 to 33 mm.) ; 
outer upper incisor more than half as high as inner ; large 
premolar broadly in contact with canine, the small tooth very 
minute, crowded inward from axis of tooth-row, invisible from 
outer side and occasionally covered by the gum, its diameter 
much less than that of outer incisor ; anterior lower premolar 
with crown area less than half that of succeeding tooth ; lower 
canine robust ; tragus with greatest width nearly equal to length 
of anterior border ; thumb short ; hairs of back usually with 
contrasting light brown tips. 

External characters. — Ear broad, its general form about as in 
Pipistrellus nathusii, the posterior border slightly but evidently 
concave above middle, the inner surface of conch noticeably 
rugose and with faint, irregular transverse ridges behind tragus ; 
antitragus small and ill-defined, but producing an evident break 
in outline of conch ; tragus less than half as high as conch, very 
wide (greatest width, at level of middle of anterior border, nearly 
equal to length of anterior border), the anterior border nearly 
straight, the posterior border strongly and evenly convex from 
tip to notch above small basal lobe. Wing, foot, calcar and tail 
as in P. pipistrellus. 

Fur and colour. — The fur resembles that of Pipistrellus kuhlii 
in quality and distribution. Colour differing from that of the 
other European species in the evident contrast between light 
tips of hairs of back and darker ground tint. It is also the only 
species in which there is much individual variation in colour. 
Four specimens from the neighbourhood of Genoa are coloured as 
follows : male, not fully adult, uniform very dark vandyke-brown, 
the extreme tips of hairs of back faintly lighter, underparts a 
light brown faintly overlaid on blackish under colour ; adult male : 
light tips on back well developed, giving general colour to region, 
between raw-umber and clay-colour ; adult male : light tips very 



PIPISTRELLUS 



221 




Pipistrellus sari 
Nat. size. 



conspicuous, a peculiar dull brownish ochraceous-buff; adult female : 
light tips as conspicuous as in last, dull brownish cream-buff. 

Skull. — The skull is slightly larger than that of Pipistrellus 
nathusii and P. huhlii, and is immediately distinguishable among 
the European species by the flatness of dorsal surface of rostrum, 
prominence of ridge along edge of orbit, and 
relatively small size of narial emargination, 
characters the first two of which suggest 
Verpertilio murinus. Dorsal profile of skull as 
in P. pipistrellus and P. lcuhlii, but brain-case 
slightly more depressed. Breadth of brain-case 
about half greatest length of skull. Rostrum 
relatively broader than in the other European 
species, its dorsal surface more flattened and 
orbital ridges more prominent ; narial emar- 
gination scarcely larger than in P. pipistrellus. 
Mesopterygoid space about as wide as long, its 
general outline, aside from the notch caused 
by median spine of palate, broadly barrel shape, 
the hamulars distinctly turned inward. Mandible with coronoid 
process and upper border of posterior portion as in P. Jnihlii, 
but with angular process less curved and relatively longer than 
in the other European species. 

Teeth. — Incisors both above and below essentially as in 
Pipistrellus pipistrellus, except that inner upper tooth has the 
secondary cusp somewhat better developed. 
Canines with no special peculiarities, not 
essentially different from those of P. pipi- 
strellus. Anterior upper premolar very 
minute, sometimes hidden in the gum or 
occasionally absent, its crown area never 
much more one-sixth that of outer incisor ; 
large premolar always strongly in contact 
with canine, its form peculiar in the absence 
or slight development of the antero-internal 
cusp. Lower premolars strongly contrasted 
in size, the crown area of first decidedly less than half that 
of second, its cusp relatively lower and less developed than in 
any of the other European species. Molars both above and 
below with no special peculiarities. 

Measurements. — Two adult males from Palermo, Sicily : head 
and body, 43 and 47 ; tail, 34 and 35 ; tibia, 12-6 and 12-8; 
foot, 6-4 and 7; forearm, 31 and 32-6; thumb, 5 and 5-6; 
third finger, 54 and 56 ; fifth finger, 42 and 41 ; ear from meatus, 
12 - 4 and 12 • 6 ; width of ear, 12 and 12. Two adult females 
from the same locality : head and body, 46 and 47 ; tail, 35 and 
39 ; tibia, 13 and 13 '4 ; foot, 6-6 and 7 ; forearm, 33 and 33 ; 
thumb, 5 and 5 • 4 ; third finger, 56 and 57 ; fifth finger, 42 and 
43 ; ear from meatus, 12*4 and 13 ; width of ear, 12 and 11-6. 




C2» 



Flfl. 39. 



Pipistrellus savii. 
Anterior teeth, x 5. 



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224 



CHIROPTERA 



Adult male from Escorial, Spain (paratype of ochromixtus Cabrera) : 
head and body, 48 • 6 ; tail, 34 ; tibia, 13 • 2 ; foot, 5 • 6 ; forearm, 
57 ; thumb, 5 - 4; third finger, 43 ; ear from meatus, 13 ; width 
of ear, 11. For cranial measurements see Table, p. 223. 

Specimens examined. — Twenty-four, from the following localities : — 

Switzerland : St. Gothard, Uri, 1 ; no exact locality, 1. 

Italy : Near Genoa, 10 (B.M. and Genoa) ; Florence, 3 (Mottaz) ; 
Sorrento, 1 (U.S.N.M.) ; no exact locality, 1 (type) ; Palermo, Sicily, 8 
(U.S.N.M.) ; Sicily, no exact locality, 1 (type of leucippe Bonaparte) ; 
Ustica Island, Sicily, 1 (U.S.N.M.). 

Prance : Near Nimes, Gard, 2 (U.S.N.M. and Nimes ; the latter 
agreeing with description of nigrans Crespon, though not marked type) ; 
St. Gilles, Gard, 1 ; no exact locality, 1. 

Spain: El Escorial, Madrid, 1 (paratype of ochromixtus Cabrera). 

Remarks. — This species is readily distinguishable among the 
European members of the genus Pipistrellus by the peculiar form 
of the tragus, apart from its very pronounced cranial and dental 
characters. Its colour gives it a superficial resemblance to 
Ejptesicus nilssoni, a likeness that is so heightened by the exces- 
sively small size of the anterior upper premolar that the animal 
has been once and perhaps twice described as a member of the 
genus Eptesicus* 

1. St. Gothard, Uri, Swit- Tomes Collection. 7. 1. 1. 397. 

zerland. 
Switzerland. Purchased (Brandt). 45. 11. 1. 3. 

4 al. Genoa, Liguria, Italy. Genoa Museum (e). 86. 11. 3. 14- 

17. 
<5. Borzoli, Liguria. Marquis G. Doria 5. 12. 15. 6. 

(c & p). 
skeleton with- Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 732. 

out skull. parte.) (Type of species.) 

skeleton with- Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 731. 

out skull. parte.) (Type of V. leucippe Bonaparte.) 

9 al. Prance. Purchased (Lefebre). 46.1.2.12. 

S. El Escorial, Madrid, A. Cabrera (p). 8. 7. 23. 1. 

Spain. (Paratype of V. ochromixtus Cabrera.) 

Genus EPTESICUS Rafinesque. 

1820. Eptesicus Rafinesque, Annals of Nature, p. 2 (melanops = fuscus). 
1829. Cnephseus Kaup, Entw.-Gesch. u. Natiirl Syst. Europ., Thierwelt, i, 

p. 103 (serotinus). 
1839. Vesperugo Keyserling and Blasius, Wiegmann's Archiv fur Natur- 

geseh., 1839, i, p. 312 (part). 
1839. Vesperus Keyserling and Blasius, Wiegmann's Archiv fur Natur- 

gesch., 1839, i, p. 313 (Sub-genus of Vesperugo, part). Not Vesperus 

Latreille, 1829. 



* This is certainly the case with the Vespertilio ochromixtus of Cabrera. 
In the paratype of this species (B.M. no. 8. 7. 23. 1), which I carefully 
examined before removal of the skull, in company with Mr. Knud Andersen, 
no trace of the small premolar could be found. When the skull was cleaned, 
however, the presence of the tooth in its normal position was revealed, 
thus showing the animal's true identity. It. seems not improbable, so far 
as can be judged from the original description, that Satunin's Vesperugo 
caucasicus (Zool. Anzeiger, xxiv, p. 462, August 5, 1901) was based on 
similar specimens. 



EPTESICUS 



225 



1841. Noctula Bonaparte, Iconogr. Faun. Ital., I, faso. xxi, in account of 

Vespertilio alcythoe (serotinus). 
1856. Cateorus Kolenati, AUgem. deutsche Naturhist. Zeitung, Dresden, 

neue Folge, n, p. 131 (serotinus). 

1856. Meteorus Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden, 

neue Folge, n, p. 131 (part). 

1857. Vesperus Blasius, Saugethiere Deutschlands, p. 51 (Sub-genus of 

Vesperugo), part. 

1858. Amblyotus Kolenati, Sitzungsber. kais. Akad. Wissensch. Wien, 

Math.-Naturwissenseh. Classe, xxix, p. 252 (atratus = nilssoni). 
1863. Aristippe " Kolenati, Beitriige zur Kenntniss der Phthiriomyiarien, 

Petersburg, 1863 " (part, included both discolor = murinus and 

nilssoni). 
1866. Pachyomus Gray, Ann. and Mag. Nat. Hist., 3rd ser., xvn, p. 90, 

February, 1863 (pachyomus). 
1870. Nyctiptenus Fitzinger, Sitzungsber. kais. Akad. Wissensch. Wien, 

Math.-Naturwiss. Classe, lxii, p. 424 (smithii). 
1878. Vesperus Dobson, Oatal. Chiropt. Brit. Mus., p. 184 (Sub-genus of 

Vesperugo), part. 
1892. Adelonycteris H. Allen, Proc. Acad. Nat. Sci. Philadelphia, 1891, 

p. 466, January 19, 1892 (part; substitute for Vesperus, pre- 
occupied). 
1897. Vespertilio Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 384, 

October, 1897 (part). 
1900. Eptesicus Mehely, Monogr. Chiropt. Hungariee, p. 219 (part). 
1907. Eptesicus Miller, Families and Genera of Bats, p. 207, June 29, 1907. 

Type species. — Eptesicus melanops Rafinesque = Verpertilio 
fuscus Beauvois. 

Geographical distribution. — Europe, Asia (except Malay region), 
Australia, Africa, Madagascar ; America from southern Canada 
southward (except Lesser Antilles). 

Characters. — Dental formula : i £?. c J4, P m iS; m S = 32. 
Teeth strictly normal throughout, and showing no special pecu- 
liarities. Both upper incisors well developed, the inner larger 
than the outer and usually with distinct secondary cusp, the 
outer separated from canine by a space equal to its greatest 
diameter ; m 3 variable in form, usually with well developed 
metacone and three commissures in the smaller species, but with 
metacone and third commissure obsolete in larger forms. Skull 
without special peculiarities of form or structure, the rostrum 
flattish or more usually rounded off above, the nares and palatal 
emargination not specially enlarged, the latter at least as deep 
as wide. Ear of moderate size, not peculiar in form ; wing 
broad (normal). 

Bemarlis. — Among European bats the members of the genus 
Eptesicus may be distinguished by their dental formula combined 
with a simple Pipistrellus-like ear and not specially modified 
skull. The group is nearly related to Pipistrellus through 
P. savii, in which the small premolar is occasionally absent and 
not infrequently so minute as to be concealed by the gum. About 
forty-five species are known, three of which occur in Europe. 

Q 



226 CHIEOPTEKA 

KEY TO THE EUROPEAN SPECIES OP EPTESICUS. 

Porearm less than 40 mm. ; condylobasal length of 
skull less than 16 mm. ; a distinct line of demar- 
cation between colours of upper and lower surfaces 
of neck E. nilssonU, p. 234. 

Porearm more than 45 mm. ; condylobasal length of 
skull more than 17 mm. ; no line of demarcation 
between colours of upper and lower surfaces of 
neck. 

Condylobasal length of skull 19 to 21 - 6 mm E. serotinus, p. 226. 

Condylobasal length of skull about 18 mm E. sodalis, p. 231. 

eptesicus serotinus Schreber. 

1774. Vespertilio serotinus Schreber, Saugthiere, i, pi. liii (Description, i, 

p. 167, 1775, under name: Die Blasse Pledermaus). Prance, 

based primarily on "La Se>otine" of Daubenton, Hist. Acad. 

Royale des Sci., 1759, p. 377. 1765. 
1776. Vespertilio serotine P. L. S. Miiller, Natursyst. Suppl. u. Regist.- 

Band, p. 16 (Based on "die Blasse Pledermaus " of Schreber). 
1827. Vespertilio wiedii Brehm, Ornis, Heft in, p. 24 (Renthendorf, 

Thviringen, Germany). 
1827. Vespertilio okenii Brehm, Ornis, Heft in, p. 25 (Renthendorf, 

Thiiringen, Germany). 
1844. Vesp\eartilio\ incisivus Crespon, Paune Meridionale, I, p. 26 (Nimes, 

Gard, Prance). 
1857. Vesperugo serotinus Blasius, Saugethiere Deutschlands, p. 76. 
1863. [Cateorus serotinus'] var. typus Koch, Jahrb. des Vereins fur Natur- 

kunde im Herzogthum Nassau, xvni, p. 466 (Wiesbaden, Nassau, 

Germany). 
1863. [Cateorus serotinus] var. rufescens Koch, Jahrb. des Vereins fiir 

Naturkande im Herzogthum Nassau, xvm, p. 466 (Preiburg, 

Breisgau, Germany). 
1878. Vesperugo serotinus Dobson, Catal. Chiropt. Brit. Mus., p. 191. 

1885. Vesperus serotinus var. transylvanus Daday, Orvos-Termeszettudo- 

manyi Erteseto', Kolozsvar, x, p. 275 (Also-Szoes, Szolnok-Doboka, 
Hungary). 

1886. Vesperus serotinus var. transsylvanus Daday, Verhandl. u. Mittheil- 

ungen des Siebenbiirgischen Vereins fur Naturwissensch. in 

Hermannstadt, xxxvi, p. 81. 
1900. Eptesicus serotinus M6hely, Monogr. Chiropt. Hungarise, p. 209. 
1904. Vespertilio serotinus insularis Cabrera, Mem. Soc. Espafl. Hist. Nat., 

n, p. 263 (Minorca, Balearic Islands). 
1904. Vespertilio isabellinus Cabrera, Mem. Soc. Espafl. Hist. Nat., n, 

p. 264 (southern Spain). Not of Temminck. 
1904. Vespertilio boscai Cabrera, Mem. Soc. Espafl. Hist. Nat., n, p. 265 

(Muchamiel, Alicante, Spain). 
1910. Eptesicus serotinus, E. serotinus transsylvanus, and E. boscai Troues- 

sart, Paune Mamm. d'Europe, pp. 20-22. 

Type locality. — France. 

Geographical distribution. — Central and southern Europe 
from England and Denmark to the Mediterranean ; eastward 
into Asia. 

Diagnosis. — Size rather large (forearm more than 45 mm., 



EPTBSICUS 227 

condylobasal length of skull more than 19 mm.) ; colour of upper 
parts a yellowish brown without noticeably contrasted light tips 
to the hairs ; under parts essentially similar, so that there is no 
line of demarcation along sides of neck. 

External characters. — General form robust, though less so 
than in Nyctalus, the tail and legs rather short, the membranes 
thick and opaque. Muzzle with moderately prominent glandular 
swellings, its greatest width across this region less than distance 
from nostril to ear ; nostrils projecting very slightly, the concavity 
between them not conspicuous, the orifice crescentic. Ear 
moderately long, e±tending slightly more than half way from eye 
to nostril when laid forward, its breadth when flattened about 
equal to height above crown ; anterior border of conch abruptly 
convex below, then nearly straight to narrowly rounded off tip ; 
posterior border straight or irregularly concave from just below 
tip to level of anterior base, then convex to abrupt angle under 
meatus marking posterior limit of small but well defined anti- 
tragus, the anterior border of which terminates obscurely about 
3 mm. behind angle of mouth ; inner surface of conch obscurely 
papillose, the region behind tragus marked by about six faint 
cross ridges ; tragus short, its height less than half that of ear 
conch, its anterior border straight, its posterior border gently 
convex from narrowly rounded tip to upper edge of small but 
distinct basal lobe, its greatest width, at level of middle of anterior 
border, equal to slightly more than half length of anterior border. 
Wing broad, the fifth finger exceeding forearm by one-quarter 
to one-third length of forearm, the membrane leathery and 
opaque, though perhaps less so than in Nyctalus noctula, joining 
leg at base of outer toe ; third and fourth metacarpals sub-equal, 
nearly as long as forearm, fifth about 2 mm. shorter. Leg rather 
slender, the foot less than half as long as tibia ; calcar about 
four-fifths as long as tibia and slightly exceeding length of free 
border of uropatagium, its keel ill-defined, its terminal lobe well 
developed though small. Tail extending to between shoulders 
when laid forward, the last vertebra and distal third of penulti- 
mate vertebra free from membrane. 

Fur and colour. — Pur soft and dense, the longest hairs on 
back about 10 mm. in length, those of underparts shorter. It 
is strictly confined to body, only extending as a thin pubescence 
on extreme base of membranes and along a narrow line bordering 
under surface of forearm ; free edge of uropatagium naked. 
Ground colour of upper parts ranging from prouts-brown to a 
light wood-brown, the basal portion of the hairs not essentially 
different, the tips of the hairs of back behind shoulders with 
inconspicuous lighter (buffy) tips ; underparts slightly paler, 
sometimes approaching ochraceous-buff, but never sufficiently 
contrasted to produce a line of demarcation along sides of neck. 
Muzzle, cheeks, ears and membranes blackish. The variation 
in general colour appears to be strictly individual. 

Q 2 



228 



CHIEOPTERA 



Skull. — General aspect of skull robust and flattened, with 
widely spreading zygomata, but rather narrow brain-case and 
rostrum. Dorsal profile rising gradually from nares to over- 
hanging lambda, essentially straight throughout, though with 
slight concavity over lachrymal region. Ventral profile very 
slightly elevated posteriorly. Brain-case ovate in general outline, 
narrower than in Nyctalus noctula, the straight, well developed 
lambdoid crests which form its posterior border meeting in median 
line almost at right angles, depth at middle about half mastoid 
breadth ; sagittal crest low but evident, the region at each side 
of it not depressed ; floor of brain-case smooth, with no evident 
ridges or depressions, a very narrow slit between cochlea and 
basioccipital ; auditory bullae small, the transverse diameter con- 
siderably less than distance between bullse. Interorbital region 
moderately constricted, hour-glass shaped, the lachrymal region 

decidedly less wide than brain-case, 
with slight tubercular projection close 
to anterior rim of orbit ; rostrum 
flattened, with shallow but evident 
lateral concavity on each side, dis- 
tinctly narrower anteriorly than pos- 
teriorly, the narrowly obovate narial 
emargination extending about half 
way back to level of lachrymal fora- 
men ; rostral depth at front of orbit 
less than distance from orbit to outer 
incisor ; anteorbital foramen less re- 
duced than in Nyctalus noctula, its 
posterior border over region of con- 
tact between large premolar and first 
molar, lachrymal foramen directly 
behind it, on inner side of orbital 
rim. Palate long and narrow, slightly concave both laterally and 
longitudinally, the anterior emargination small, squarish, extend- 
ing back to level of middle of canine. Posterior extension of 
palate nearly parallel sided, though narrowing a little posteriorly, 
its width at level of posterior molar considerably less than its 
length ; hamulars slightly turned inward ; median spine well 
developed. Mandible robust, but with lower border nearly 
parallel to alveolar line ; posterior portion high in front, low 
behind, the height of coronoid process above level of alveolar 
line about equal to horizontal diameter, the upper border sloping 
abruptly from coronoid to articular process ; angular process 
moderately long, about on level with alveolar line, its main axis 
directed gradually outward and downward, its distal extremity 
slightly expanded and hooked upward. 

Teeth. — Relatively to size of skull the teeth are large and 
robust, rather more so than in Nyctalus noctula. Inner upper 
incisor about half as high as canine, its crown area about one- 




FIG. 40. 
Epte&ieus serotinus. Nat. size. 



BPTBSICUS 



229 



quarter that of canine, the subterete shaft directed inward and 
slightly forward, its secondary cusp large and prominent, situated 
on outer side of shaft near tip ; cingulum well developed, but 
without cusps. Outer upper incisor much shorter than inner 
and with about half its crown area, the apex of its shaft slightly 
exceeding level of cingulum of larger tooth ; outer and posterior 
surfaces flattened or double-concave, tbe two concavities some- 
times separated by a low but evident ridge ; inner margin with 
a low secondary cusp on well developed cingulum. A line 
perpendicular to main axis of skull would pass through centre 
of shafts of all four incisors ; the outer tooth is separated from 
canine by a space about equal to breadth of its own crown. Lower 
incisors large, much crowded, and very conspicuously imbricated, 
i 1 and i 2 overlapping more than half of front surface of the 
succeeding tooth, the general outline of the entire series V-shaped ; 
front surface of crowns about as high as wide, the edge obliquely 
trifid (occasionally a low, rudimentary fourth cusp at outer 
margin of i 2 ) ; cross section of crown somewhat triangular, the 
posterior angle occupied by a low tubercle in i 2 and i 3 . Upper 
canine large, its shaft decidedly the highest of the upper series, 
its cross section sub-triangular, the posterior cutting edge well 
developed, the anterior less trenchant than in Nyctalus noctula ; 
inner surface divided by a low ridge into two shallow con- 
cavities, the posterior of which is the larger ; antero-outer surface 
convex ; postero-outer surface with deep longitudinal groove ; 
cingulum narrow but complete, without cusps. Lower canine 
very robust, the diameter of crown noticeably greater than least 
distance between canines, the shaft decidedly higher than main 
cusps of molars, smoothly rounded in front, flattened-concave 
behind and on inner side, the cingulum narrow but complete 
except at point of contact with i 3 , where it becomes abruptly 
obsolete, terminating in a slight tubercle corresponding to the 
cusp present in Nyctalus and Pipistrellus. Upper premolar with 
crown area about equal to that of canine and about two-thirds 
that of first molar, its main cusp robust, nearly as long as canine 
(measured along cingulum), sharply trenchant posteriorly, flat- 
tened-concave on inner side, a well developed external and antero- 
internal longitudinal groove ; crown with slight anterior and 
more marked posterior emargination, the inner side narrow, with 
slight concave crushing surface and well developed cingulum, 
which rises to a small cusp anteriorly. Lower premolars closely 
crowded, the crown area of first about half that of canine, that 
of second nearly four-fifths that of canine ; cusp of first a little 
more than half as high as second, which slightly exceeds main 
cusps of molars ; cingulum of each tooth well developed, tending 
to form a slight postero-internal tubercle. First and second 
upper molars sub-equal in crown area, the second wider but more 
constricted at middle ; protocone robust, not very high ; no true 
hypocone, but region which it would occupy indicated by slight 



230 



CHIEOPTEEA 



columnar thickening of posterior base of protocone ; paracone 
lower and smaller than metacone, the contrast unusually notice- 
able ; styles and commissures well developed, the W-pattern 
normal ; m 3 with crown area less than half that of m 1 , its longi- 
tudinal diameter through metacone much less than half transverse 
diameter, the mesostyle, metacone, and second and third com- 
missures greatly reduced, though not sufficiently to lose their 
identity. Lower molars with no special peculiarities ; angles in 
commissures between outer and inner cusps rather wide and 
shallow, especially that between protoconid and metaconid ; area 
of second V in. m 3 scarcely half that of first. 

Measurements. — Adult female from Herrnhut, Saxony : head 
and body, 62 ; tail, 54 (its free tip, 6-6) ; tibia, 20 ; foot, 10 • 4 ; 
forearm, 50 • 4 ; thumb, 9 ; third finger, 84 : fifth finger, 63 ; ear 
from meatus, 18 ; width of ear, 15. Adult male from Barsac, 
Gironde, France : head and body, 69 ; tail, 54 (free tip, 7 ■ 6) 
tibia, 22 ; foot, 11 -4 ; forearm, 51 ; thumb, 9 ; third finger, 91 
fifth finger, 67 ; ear from meatus, 18 ■ 4 ; width of ear, 1 6. Adult 
male and female from Seville, Spain : head and body, 67 and 64 
tail, 46 and 51 ; tibia, 21-6 and 22-4 ; foot, 11 and 10-4 ; fore 
arm, 49 and 50 ; thumb, 9 ■ 2 and 8 ■ 4 ; third finger, 89 and 90 
fifth finger, 64 and 68 ; ear from meatus, 17 "6 and 18. Adult 
male and female from Rome, Italy : head and body, 72 and 73 
tail, 54 and 54 ; tibia, 21-4 and 21 ; foot, 11 and 11 -4 ; fore 
arm, 51 • 6 and 53 ; thumb, 9 • 6 and 9 ■ 8 ; third finger, 91 and 92 
fifth finger, 66 and 67 ; ear from meatus, 19 and 19 ; width of 
ear, 15 ■ 6 and 15 • 4. For cranial measurements see Table, 
p. 232. 

Specimens examined. — Seventy-eight, from the following localities : — 

England: Kenley, Surrey, 1 ; Hawkhurst, Kent, 1 ; Whitstable, Kent, 2; 
Wingham, Dover, Kent, 1 ; Yalding, Kent, 5 (B.M. and U.S.N.M.) ; Isle 
of Wight, 5. 

France: Barsao, Gironde, 1 (U.S.N.M.); near Nlmes, Gard, 2 (Nlmes; 
type of incisivus Crespon, and a specimen wrongly marked type of paluslris 
Crespon). 

Germany : Ingelheim, Bheinhessen, 2 ; Magdeburg, Saxony, 2 (B.M. 
and U.S.N.M.) ; Herrnhut, Saxony, 1 (U.S.N.M.) ; Strass, near Burgheim, 
Bavaria, 3 ; Bavaria, no exact locality, 2 (U.S.N.M.) ; Tubingen, Silesia, 2. 

Austria-Hungary : Moravia, 1 ; Csall6kbz-Somorja, Pressburg, Hun- 
gary, 1 ; Budapest, Hungary, 2 ; Transylvania, 5 ; Zara, Dalmatia, 2. 

Eoumania : Bustenari, Prahova, 3. 

Greece : Patras, 14 ; near Athens, 1. 

Italy : Siena, 3 (B.M. and U.S.N.M.) ; Florence, 2 (B.M. and U.S.N.M.) ; 
Vallombrosa, 1 (U.S.N.M.) ; Volterra,l (U.S.N.M.) ; Eome,3; SanMartino 
al Cimino, Borne, 1 (Genoa) ; Ustica Island, Sicily, 1 (U.S.N.M.). 

Sardinia : Cagliari, 2 (Genoa). 

Spain: Pa] ares, Leon, 1; Seville, 3; Muchamiel, Alicante, 1 (Madrid; 
type of boscai Cabrera). < 

Bernarks. — Among the bats of Europe this species is recogniz- 
able by its rather large size, noticeable free tip to the tail, 
moderately long, narrow ear, and straight, erect tragus. With 
the material now available for study it seems impossible to 



EPTESICHS 



231 



distinguish any local geographical forms. Specimens from Seville 
representing the isabellinus of Cabrera I am unable to separate 
from true serotinus ; the type of boscai Cabrera is a young of the 
same animal ; insularis I have not seen, but there is nothing in 
the original description to indicate that it is distinct.* 



i. Keuley, Surrey, England. 



2 9. 
Sal. 
1st. 
3?. 

Sal. 
2. 
1. 

1. 

6,9. 



2<5, S. 

2al. 
lal. 



2al. 

5<5al. 

<?,$• 
<5,S. 

IS. 

S. 

?. 

lal. 
t. 
S. 

Sal. 

lal. 

1. 



Whitstable, Kent. 
Wingham, Kent. 
Yalding, Kent. 
Yalding, Kent. 

Isle of Wight. 

Freshwater, Isle of Wight. 

Freshwater, Isle of Wight. 
(F. Bond.) 

Bembridge, Isle of Wight. 

Ingelheim, Rheinhessen, 
Germany. 

Magdeburg, Saxony. 
(Wolterstorff.) 

Strass, Burgheim, Bavaria. 
(KOrbitz.) 

Tubingen, Silesia. 

Moravia, Hungary. 

Csall6k6z-Somorja, Press- 
burg. 

Budapest. 

Transylvania. 

Zara,Dalmatia. (E.Blos.) 
Bustenari, Prahova, 840 m. 
Roumania. (W.Dodson.) 
Patras, Greece. (C.Mottaz.) 



W. R. Ogilvie-Grant 

(c&p). 
G. H. B. Grant (c & p). 
G. Donker (c & p). 
H. Reid (c & p). 
W. R. Ogilvie-Grant 

(c & p). 
Rev. C. Bury (c & p). 
P. Bond (c&p). 
Tomes Collection. 

Tomes Collection. 
C. Hilgert (c). 



Patras. 
Athens. 



(0. Mottaz.) 



Florence, Italy. 
Rome. (C. Coli.) 
Pajares, Leon, 

(N. Gonzalez.) 
Seville. 
Seville. 
Europe. 



Spain. 



Lord Lilford (p). 

Lord Lilford (p). 

Dr. A. Giinther (p). 
Purchased (Parreys). 
Budapest Museum (e). 

Budapest Museum (e). 
0. G. Danford and J. A. 

Brown (o & p). 
Lord Lilford (p). 
Lord Lilford (p). 

Hon. N. C. Rothschild 

ro- 
cs. Mottaz (c). 

Hon. N. C. Rothschild 

Florence Museum (e). 
G.Barrett-Hamilton (p). 
0. Thomas (p). 

Dr. V. L. Seoane (p). 
Seville Museum (e). 
Leyden Museum (e). 



11. 1. 3. 45. 

11. 1. 3. 46-47. 
90. 4. 17. 1. 
97. 8. 27. 1. 
11.1.3.42-44. 

44. 6. 15. 7. 
61. 11. 5. 1-2. 
7. 1. 1. 352. 

7. 1. 1. 353. 

8. 11. 2. 1-2. 

11. 1. 1. 38. 

11. 1. 1. 33-35. 

66. 2. 1. 7-8. 
46. 6. 15. 54. 
94. 3. 1. 12-13. 

94. 7. 18. 6-7. 
74. 7. 4. 1-5. 

11. 1. 1. 36-37. 
4. 4. 6. 2-3. 

8. 10. 2. 1-13, 

15. 
8. 11. 3. 3. 
8. 10. 2. 14. 

85. 7. 6. 13. 
11. 1. 2. 28. 
8. 2. 9. 1. 

94. 1. 1. 6. 
94. 5. 8. 1. 
37. 4. 28. 58. 



EPTESICUS SODALIS Barrett-Hamilton. 

1910. Vespertilio sodalis Barrett-Hamilton, Ann. and Mag. Nat. Hist., 

8th ser., v, p. 291, March, 1910. Type in British Museum. 
1910. Eptesicus sodalis Trouessart, Faune Mamm. d'Europe, p. 22. 

Type locality. — Bustenari, Prahova, Roumania (in Carpa- 
thians, alt. 840 m.). 

Geographical distribution. — Known only from the type locality 
and St. Gothard, Switzerland. 

* Mr. Cabrera has come to the same conclusion (Bol. Real Soc. Espafi. 
Hist. Nat., vi, p. 449, December, 1908). 



232 



CHIEOPTEEA 









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233 





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234 



CHIBOPTEEA 



Diagnosis. — Similar to Eptesicus serotinus but smaller, condylo- 
basal length of skull about 18 mm. instead of 19 to 21 6 mm. 

Measurements. — Type (young-adult male) : head and body, 
63; tail, 42; tibia, 18-6; foot, 9-8; forearm, 45-4; third 
finger, 79-0; fifth finger, 58-0; ear (fresh), 18. Adult from 
St. Gothard, Switzerland : tibia, 19 ; forearm, 48. For cranial 
measurements see Table, p. 233. 

Specimens examined. — Two, from the following localities : — 
Switzerland : St. Gothard, 1 (U.S.N.M.). 
Roumania : Bustenari, Prahova, 1 (type). 

Remarks. — The two specimens on which this species is based 
indicate the existence of an animal bearing much the same 
relationship to Eptesicus serotinus as Nyctalus noctula to N. 
maximus. 

6. Bustenari, Prahova, 840 m. Lord Lilford (p). 4. 4. 6. 1. 

Boumania. (W. Dodson.) (Type of species.) 



eptesicus nilssonii Keyserling and Blasius. 

1836. Vespertilio kuhlii Nilsson, Ilium. Pig. Skand. Fauna, pt. 17, pi. 34 
upper figure. Not of Kuhl, 1819. 

1838. Vespert[ilio] borealis Nilsson, Tllum. Pig. Skand. Fauna, pt. 19, 

pi. 34 (renumbered 36) upper figure. Not of P. L. S. Miiller, 1776 
(Scandinavia). 

1839. V[espertilw] nilssonii Keyserling and Blasius, Wiegmann's Archiv 

fur Naturgesch., 1839, p. 315 (Mountains of Scandinavia. Based 
on the V. kuhlii of Nilsson, 1836). 

1857. Vesperugo nilssonii Blasius, Saugethiere Deutschlands, p. 70. 

1858. Amblyotus atratus Kolenati, Sitzungsber. kais. Akad. Wissensch. 

Wien, Math.-Naturwissensch. Classe, xxix, p. 252 (Altvater, 

Austrian Silesia, alt. 2400-4600 ft.). 
1878. Vesperugo borealis Dobson, Catal. Chiropt. Brit. Mus., p. 203. 
1894. Vesperugo nillsoni (sic) Bhoads, Beprint Ord's N. Amer. Zoology, 

Append., p. 3. 
1907. E[ptesicus~\ nilssoni Miller, Families and Genera of Bats, p. 209, 

June 29, 1907. 
1910. Eptesicus nilssoni Trouessart, Faune Mamm. d'Europe, p. 23. 

Type locality. — Sweden. 

Geographical distribution. — Continental Europe, from northern 
Norway to the Alps. 

Diagnosis. — Size medium (forearm less than 40 mm., condylo- 
basal length of skull less than 16 mm.) ; colour of upper parts a 
rich dark brown, the hairs of back with noticeably contrasted 
light tips ; underparts light yellowish brown ; a well defined 
line of demarcation along sides of neck. 

External form. — In general the external form agrees with 
that of Eptesicus serotinus, due allowance being made for the less 
robust stature of the smaller animal. Ear relatively longer, 



EPTESICUS 



235 



extending to nostril when laid forward, though of the same 
general form as in E. serotinus ; tip less narrowly rounded off, 
and flattened or concave portion of posterior border less con- 
spicuous ; tragus relatively shorter and wider, though not 
essentially different in form, its greatest width nearly equal to 
length of anterior border ; transverse striations on inner surface 
of conch obsolete. Wings and feet essentially as in E. serotinus. 
Tail slightly longer than in the related animal, extending nearly 
to head when laid forward, its terminal vertebra free. 

Fur and colour. — In quality the fur resembles that of E. 
serotinus except that it is softer and more silky in texture, the 
hairs fully as long as in the larger animal. In distribution it 
shows no special peculiarities ; dorsal surface of uropatagium 
thinly furred to about middle instead of on extreme base only. 
Colour above a rich dark brown, ranging from burnt-umber nearly 
to seal-brown, the hairs everywhere with slightly darker, faintly 
slaty bases, those of median dorsal region from crown to base of 
tail tipped with light glossy ochraceous-buff in evident contrast 
with ground colour, the light tips most numerous behind shoulders, 
and forming a noticeable mantle over middle of back ; under- 
pays rather strongly contrasted light yellowish brown, between 
the wood-brown and ochraceous-buff of Ridgway, the basal 
portion of the hairs similar to ground colour of back, the yellowish 
brown area extending over sides of head and completely encircling 
base of ear, the line of demarcation between it and the dark 
brown of upper parts sharply defined along sides of neck. Muzzle, 
cheeks, ears and membranes blackish. 

Skull. — In general the skull differs from that of Eptesicus 
serotinus, apart from its smaller size, in a general tendency to 
greater depth, less elongation, and smoother, more evenly 
rounded surfaces. Dorsal profile with evident convexity at middle. 
Lambda not overhanging ; low, ill-defined 
lambdoid crests curving slightly toward point 
of contact at middle, the rounded posterior 
outline of the occiput plainly visible behind 
them when skull is viewed from above ; 
sagittal crest essentially absent. Brain-case 
sub-spherical or broadly ovate in outline, 
its depth slightly more than half mastoid 
breadth ; floor of brain-case marked by a 
wide lateral groove between each cochlea 
and the median line ; auditory bullae slightly 
larger than in E. serotinus. Interorbital fig. 41. 

region relatively less constricted than in Epteswm nOnomi. 
E. serotinus, but of essentially the same form ; ^«*- size - 

lachrymal swelling present but less noticeable 
than in the larger animal. Rostrum rounded off at sides, with 
only the faintest trace of lateral concavities ; narial and palatal 
emarginations essentially as in E. serotinus. Palate showing no 




236 



CHIEOPTBRA 



noteworthy peculiarities, its general outline less narrow than in 
the related species. 

Teeth. — Inner upper incisor as in Eptesicus serotinus, but more 
robust in proportion to its height ; outer incisor noticeably higher 
than in the related species, its apex reaching level of secondary 
cusp of larger tooth, its secondary cusp more prominent ; no 
marked contrast between crown areas of the two teeth; each 
pair in line of general curve of anterior portion of palate, instead 
of at right angles to main axis. Lower incisors less crowded 
than those of E. serotinus, their imbrication distinct but not 
unusual, the general outline of the row broadly V-shaped ; in form 
the individual teeth show no special peculiarities ; i 2 without 
postero-internal tubercle. Canines and premolars with no special 
peculiarities. First and second upper molars essentially as in 
Eptesicus serotinus, but disproportion between paracone and 
metacone less evident ; m 3 with crown area about two-thirds that 
of m l , its longitudinal diameter through metacone slightly more 
than half transverse diameter, the mesostyle and metacone well 
developed, the second and third commissures more than half as 
long as first ; lower molars like those of E. serotinus in form, but 
angles in commissures between outer and inner cusps deeper ; 
area of second V in ™3 nearly equal to that of first. 

Measurements. — Adult male from Grotte de Vallorbe, Vaud, 
Switzerland: head and body, 68 • 5 ; tail, 47; tibia, 17; foot, 10; 
forearm, 38 ■ 2 ; thumb, 9 • 8 ; third finger, 68 ; fifth finger, 49. 
Forearm in adult male from Upsala, Sweden, 38 ■ 4. Three 
females from the same locality: forearm, 39, 39, and 39 - 6; 
third finger in the four specimens from Upsala : 66, 68, 70 and 
68. For cranial measurements see Table opposite. 

Specimens examined. — Fifteen, from the following localities : — 

Norway : No exact locality, 1. 

Sweden: Upsala, 8 (B.M. and U.S.N.M.); Upland, 2 (U.S.N.M.). 

Germany : Wernigerode, Saxony, 2. 

Austeia-Hungaby : Csall6k6z-Somorja, Pressburg, 1. 

Switzerland : Grotte de Vallorbe, Vaud, 1 (Mottaz). 

Remarks. — This species is readily distinguishable from Eptesicus 
serotinus by its smaller size and by the conspicuous pale tips to 
the hairs of back. From Vesperiilio murinus, which resembles it 
in colour pattern, the narrow ear, more yellowish (less whitish) 
hair-tips on back, and slightly smaller size distinguish it super- 
ficially. Pipistrellus savii, which often has almost exactly similar 
coloration, is a much smaller animal (forearm, 31 to 33 instead 
of 38 to 40). 

1. Norway. (Collett.) E. B. Alston (p.) 81. 6. 9. 2. 

4?. Upsala, Sweden. (Eolthoff.) Lord Lilford (p). 11.1.1.39-42. 

6, 9. Wernigerode, Saxony, Ger- Dr. W. Wolterstorfl 0. 2. 8. 3-4. 
many. (c & p). 

1, Csall6koz-Somorja, Press- Budapest Museum (e.) 94.3.1.14. 
burg, Hungary. 



BPTESICUS 



237 



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CHIROPTEKA 



Genus VESPERTILIO Linnaeus. 



1758. Vespertilio Linnaeus, Syst. Nat., i, 10th ed., p. 31 (murinus by 

tautonymy). 
1839. Yesperugo Keyserling and Blasiua, Wiegmann's Archiv fur Natur- 

gesoh., 1839, 1, p. 312 (part). 
1839. Vesperus Keyserling and Blasius, Wiegmann's Archiv fur Naturgesch., 

1839, i, p. 313. Sub-genus of Yesperugo (part). 

1856. Meteorus Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden, 

neue Folge, n, p. 131 (part). 

1857. Yesperus, misprinted Yesperugo Blasius, Saugethiere Deutschlands, 

p. 69 (Sub-genus of Yesperugo). 
1863. " Aristippe Kolenati, Beitrage zur Kenntniss der Phthiriomyiarien, 

Petersburg, 1863" (part). 
1872. Marsipoleemus Peters, Monatsber. k. preuss. Akad. Wissenseh., 

Berlin, p. 260 (Sub-genus of Yesperugo for albigularis = murinus). 
1878. Yesperus Dobson, Catal. Chiropt. Brit. Mus., p. 183. Sub-genus of 

Yesperugo (part). 
1897. Vespertilio Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 384, 

October, 1897 (part). 
1900. Yespertilio Mehely, Monogr. Chiropt. Hungarise, p. 219 (part). 
1907. Vespertilio Miller, Families and Genera of Bats, p. 209, June 29, 1907. 

Type species. — Vespertilio murinus Linnaeus. 

Geographical distribution. — Forested northern portion of the 
Palsearctic region from the Atlantic coast eastward through 
Continental Asia. 

Characters. — Like Eptesicus, but ear much shortened and 
broadened, rostrum flattened above, with deep concavity on 
each side between nares and lachrymal region ; nares very large, 
extending back nearly half way to interorbital constriction, and 
palatal emargination extended so far laterally that its width is 
distinctly greater than its depth. 

Remarks. — The genus Vespertilio as now restricted contains 
only two species, V. murinus Linnaeus, and V. superans Thomas, 
the former European, the latter Asiatic. The form of the skull 
suggests in certain respects that of Nyctalus, and in others the 
North American Lasionycteris. 

VESPERTILIO MURINUS Linnasus. 

1758. [Vespertilio] murinus Linnseus, Syst. Nat., i, 10th ed., p. 32 (Sweden). 

1819. Yespertilio discolor Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv 
(= Neue Ann., i), pt. 2, p. 187 (Vienna, Austria). 

1857. Yesperugo discolor Blasius, Saugethiere Deutschlands, p. 73. 

1872. Vesperus (Marsipoleemus) albigularis Peters, Monatsber. k. preuss. 
Akad. Wissenseh., Berlin, p. 260 (Type supposed to have been 
taken in Mexico ; for its reference to this species see Mehely, 
Monogr. Chiropt. Hungarise, pp. 229, 341, 1900). 

1878. Yesperugo discolor Dobson, Catal. Chiropt. Brit. Mus., p. 204. 

1885. Vesperus siculus Daday, Orvos-Termeszettudomanyi Erteseto, Kolozs- 
var, x, p. 275 (Homorod-ALmas cave, Hungary). See Mehely, 
Monogr. Chiropt. Hungarise, pp. 229, 346, 1900. 



VESPERTILIO 



239 



1886. Vesperus siculus Daday, Verhandl. u. Mittheilungen des Siebenbiirg- 
ischeniVereins fiir Naturwissensch. in Hermannstadt, xxxvi, p. 82. 

1897. Vesjoertilio murinus Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, 
p. 382, October, 1897. 

1910. Eptesicus siculus and Vespertilio murinus Trouessart, Faune Mamm. 
d'Burope, pp. 23 and 25. 

Type locality.- — Upsala, Sweden. 

Geographical distribution, — Northern and central Europe, 
from southern Norway and central Sweden south to the Alps. 
One record of its occurrence in England.* 

Diagnosis. — General characters as in the genus ; length of 
forearm about 43 to 45 mm. 

External characters. — General appearance much as in Eptesicus 
nilssonii but slightly larger and more robust. Ear extending 
nearly to nostril when laid forward, its width when flattened 
about 1£ times height from crown ; anterior basal lobe so much 
reduced as to suggest the keel of the Molossidse, the anterior 
border of conch nearly straight from forehead to broadly rounded 
tip ; posterior border shallowly sinuous-concave from just below 
tip nearly to level of meatus ; here it turns forward almost at 
right angles and extends nearly to angle of mouth, where it 
terminates at a well developed wart, the terminal portion often 
forming a slight fold or pocket ; antitragus low and keel-like, 
nearly 2 mm. from margin of conch ; inner surface of conch 
slightly papillose, without evident cross ridges ; tragus low, 
scarcely rising above level of outer angle of anterior lobe, its 
greatest width (slightly above middle) nearly equal to length of 
straight or slightly concave anterior border, its tip broadly 
rounded, the convexity continuous behind with that of upper 
portion of posterior border ; posterior basal lobe obsolete. Wing 
rather narrow, the fifth finger exceeding forearm by one-sixth to 
one-fifth length of forearm, the membranes thin, extending to 
base of outer toe ; third and fourth metacarpals sub-equal, nearly 
5 mm. shorter than forearm, fifth about 3 mm. shorter than 
third. Leg rather robust ; foot about half as long as tibia ; 
calcar slightly longer than tibia and free border of uropatagium, 
its keel ill defined, its termination obscure. Tail extending to 
between shoulders when laid forward, the last vertebra free. 

Fur and colour. — The fur is rather short and dense, less silky 
than that of Eptesicus nilssonii, the hairs at middle of back about 
7 mm. in length. In distribution it shows no special peculiarities ; 
upper surface of interfemoral membrane furred on basal third. 
Colour above essentially as in Eptesicus nilssonii, but light tips to 
hairs very pale, almost whitish cream-buff, producing a decidedly 
" frosted " appearance ; underparts cream-buff, the dusky under 
colour showing through on chest and anterior portion of belly ; 
line of demarcation on sides of neck sharply defined ; muzzle and 
chin dusky ; ears and membranes blackish. 

* A single specimen, undoubtedly a straggler, taken at Plymouth. 



240 



CHIROPTERA 




Skull. — The skull combines the short, smooth brain-case of 
Eptesicus nils8onii with the broad, flat rostrum of E. serotinus, but 
differs conspicuously in aspect from both in the very deep, 
Nyctalus-like narial and palatal emarginations. Dorsal profile 
essentially straight from nares to rounded, not overhanging 
lambda. Sagittal crest obsolete ; lambdoid crest low but evident. 
Depth of brain-case at middle decidedly more than half mastoid 
breadth ; floor with wide conspicuous slit between cochlea and 
basioccipital ; a flattened pit-like depression at front of each slit ; 
auditory bullae moderately large, their trans- 
verse diameter about equal to space between 
them. Interorbital region broadly hour-glass 
shaped, flattened above ; edge of orbit from 
lachrymal region nearly to most constricted 
portion distinctly and irregularly swollen. 
Dorsal surface of rostrum flattened, with broad, 
shallow lateral depressions ; narial emargina- 
tion broadly triangular, its apex extending 
back to level of lachrymal region ; rostral depth 
at front of orbit equal to distance from orbit 

Ves P e,-tiKo Zurinm. to f ront of inner incisor J . anteorbital foramen 
Nat. size. moderately large, over point of contact between 

large premolar and first molar ; lachrymal fora- 
men directly behind it on inner rim of orbit. Palate broad and 
short, noticeably concave both laterally and longitudinally, the 
anterior emargination large, much wider than deep, but extend- 
ing back to level of middle of premolar ; posterior extension of 
palate parallel sided, its length slightly greater than width 
behind molars ; hamulars slightly bent inward ; median spine 
well developed. Mandible robust, much deeper at symphysis 
than behind tooth-row, the coronoid process low (height less than 
least width of posterior section) with gradually sloping upper 
border ; angular process rather slender, its extremity slightly 
curved inward. 

Teeth. — While in general resembling those of the European 
species of Eptesicus, especially those of E. nilssonii, the teeth of 
Vespertilio murinus show certain notable peculiarities. Inner 
upper incisor similar to that of Eptesicus serotinus when viewed 
from in front, but with well developed postero-basal cusp which 
in some specimens rises nearly to half the height of main cusp, 
producing a distinctly trifid tooth ; outer incisor with crown area 
slightly less than that of inner, the cusp rising nearly to middle 
of shaft of inner tooth ; secondary cusp slightly developed ; 
position of incisors relatively to tooth-row about as in E. nilssonii. 
Lower incisors as in E. nilssonii. Upper canine with transverse 
diameter slightly greater than longitudinal diameter, the reverse 
of the condition in the two species of Eptesicus ; lower canine 
with same peculiarity in outline of crown though to a less degree. 
Upper premolar with longitudinal diameter of crown relatively 



VESPEBTILIO 



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CHIKOPTERA 



less than in the European species of Eptesicus, and antero-internal 
basal cusp better developed ; lower premolars more crowded and 
compressed than in E. nilssonii, and crown area of first relatively- 
smaller. Molars as in E. nilssonii except that columnar hypocone 
of m 1 and m 2 stands out more prominently from outline of 
protocone. 

Measurements. — Three adult males from Denmark : head and 
body, 62, 59 and 62; tail, 43, 40 and 44; tibia, 17-4, 16-8 and 
16 ■ 8 ; foot, 9 ■ 6, 9 • 2 and 10 ; forearm, 43 • 4, 44 and 43 ; thumb, 
7, 7-4 and 7-6; third finger, 73, 76 and 74; fifth finger, 52, 54 
and 53 ; ear from meatus, 15, 15 and 15" 6; width of ear, 16, 
16 and 17. Adult female from the same locality: head and 
body, 63 ; tail, 41 ; tibia, 16-4 ; foot, 9; forearm, 45; thumb, 8 ; 
third finger, 78; fifth finger, 54; ear from meatus, 16; width 
of ear, 17. Adult male from Morat, Fribourg, Switzerland : 
head and body, 56 ; tail, 42 ; tibia, 16 ; foot, 8 - 8 ; forearm, 45 ; 
thumb, 5 - 4 ; third finger, 76 ; fifth finger, 55 ; ear from meatus, 
12; width of ear, 12 '4. For cranial measurements see Table, 
p. 241. 

Specimens examined. — Twelve, from the following localities : — 

England : Plymouth, 1. 

Sweden : No exact locality, 1. 

Denmark: No exact locality, 4 (U.S.N.M.). 

Germany : Gross Hennersdorf, Saxony, 1 ; Bavaria, 2 (U.S.N.M.). 

Austria-Hungary : Csallokoz-Somorja, Pressburg, 1. 

Switzerland : Morat, Fribourg, 1 (Mottaz). 

Locality unknown : One ; type of albigularis Peters (Berlin). 

'Remarks. — Superficially this bat somewhat resembles Eptesicus 
nilssoni. It is readily distinguishable, however, by its slightly 
larger size (forearm, 43 to 45 instead of 38 to 40), low, rounded 
ear, and by the whitish rather than yellowish wash on back. 

1. Plymouth, Devonshire, Dr. Leach (p). 37. a. 

England. 
— Sweden. Stockholm Museum (e). 46. 6. 2. 21. 

?. Gross Hennersdorf, Lord Lilford (p). 99.1.9.5. 

Saxony (W. Baer.) 
1. Csallokoz-Somorja, Press- Budapest Museum (e). 94. 3. 1. 15. 

burg, Hungary. 

Genus NYCTALUS Bowdich. 

1825. Nyctalus Bowdich, Excursions in Madeira and Porto Santo, p. 36 

(verrucosus). 
1829. Pterygistes Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierwelt, 

i, p. 99. 
1839. Vesperugo Keyserling and Blasius, Wiegmann's Archiv fur Natur- 

gesch., 1839, i, p. 312 (part). 
1842. Noctulinia Gray, Ann. and Mag. Nat. Hist., x, p. 258 (part ; contained 

proterus = noctula anAfulvus = Scotophilus kuhUi). 

1856. Panugo Kolenati, Allgem. deutsche Naturhist. Zeitung, Dresden, 

neue Polge, ii, p. 131 (noctula and leisleri). 

1857. Vesperugo Blasius, Saugethiere Deutschlands, p. 49 (part). 



NYCTALUS 243 

1878. Vesperugo Dobson, Oatal. Ohiropt. Brit. Mus., p. 183 (part). 

1898. Noctulinia H. Allen, Proo. U.S. National Museum, xvi, p. 30, 

June 13, 1893. 
1897. Pterygistes Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 384, 
October, 1897. 

1899. Euvcsperugo Acloque, Faune de France, Mammiferes, p. 32 (part, 

inoluded noctula, leisleri, maurus, kuhlii, pipistrellus, and 
abramus). 

1907. Pterygistes Miller, Families and Genera of Bats, p. 207, June 27, 

1907. 

1908. Nyctalus Andersen, Ann. and Mag. Nat. Hist., 9th ser., i, p. 434, 

May, 1908. 

Type species. — Nyctalus verrucosus Bowdich. 

Geographical distribution. — Northern portion of Eastern Hemi- 
sphere from the Azores and Madeira to Japan. 

Characters. — Dental formula as in Pipistrellus; skull with 
nares extending unusually far back, half way to interorbital con- 




a bed 

FIG. 43. 
Nyctalus Trmxvmus (a), N. noctula, (b), N. leisleri (c), and AT. azoreum (d). Nat. size. 

striction, and with large anterior palatal emargination ; fifth finger 
much shortened, scarcely exceeding metacarpal of fourth or third. 
Remarks.- — This genus is well differentiated from allied groups 
by the peculiar narrowing of the wing due to the shortness of the 
fifth finger. Three of the half dozen known species occur on the 
continent of Europe, while a fourth inhabits the Azores. The 
most obvious character by which the European forms are dis- 
tinguished is the size of the skull (see fig. 43). 

KEY TO THE EUBOPEAN SPECIES OP NYCTALUS. 

Condylobasal length of skull more than 17 mm. ; fore- 
arm more than 45 mm. ; hairs of back without 
dark bases. 
Condylobasal length of skull 22 to 23 mm. ; forearm 

64 to 68 mm. (Southern Europe) N. maximus, p. 244. 

Condylobasal length of skull 17 ■ 4 to 19 ■ 4 mm. ; fore- 
arm 47 to 55 mm. (Distribution general) N. noctula, p. 245. 

Condylobasal length of skull less than 17 mm. ; fore- 
arm less than 45 mm. ; hairs of back with notice- 
able dark bases. 
Condylobasal length of skull 15 to 16 mm. ; forearm 

39 to 43 mm. (Distribution general) N. leisleri,^. 252. 

Condylobasal length of skull 13 to 14-2 mm. ; fore- 
arm 35 to 42 mm. (Azores) N. azoreum, p. 254. 

R 2 



244 CHIROPTEHA 

NYCTALUS MAXIMUS Fatio. 

1781. ? VespertiHo lasiopterus Schreber, Saugethiere, pi. lvii b. See 

Thomas, Ann. and Mag. Nat Hist., 8th ser., vin, pp. 379-380, 

September, 1911. 
1827. ?? Vespertilio ferrugineus Brehm, Ornis, Heft III, p. 26 (Benthendorf, 

Thiiringen, Germany). 
1869. [Vesperugo noctula'] var. maxima Patio, Paune Vert. Suisse, i, p. 57. 

Co-type in Geneva Museum. 
1900. Pterygistes maximus Miller, Proo. Biol. Soc. Washington, xiir, 

p. 156, June 13, 1900. 
1910. Nyctalus maximus Trouessart, Faune Mamm. d'Europe, p. 19. 

Type locality. — Amsteg, Uri, Switzerland. 

Geographical distribution. — At present known from a few 
localities in Switzerland and Italy. 

Diagnosis. — Essentially similar to Nyctalus noctula but much 
larger ; condylobasal length of skull, 22 to 23 mm. ; forearm, 64 
to 68 mm. 

External characters. — Except for the conspicuously greater 
size and consequent more robust form, there appears to be no 
tangible character by which the animal can be distinguished 
from N. noctula. 

Fur and colour. — In the few specimens examined the furred 
area on under surface of wing membrane behind forearm appears 
to be better defined than in N. noctula. Colour as in the common 
species. 

Skull and teeth. — The skull and teeth so closely resemble those 
of Nyctalus noctula that in general they differ in their greater 
size only. The posterior portion of occiput is, however, more 
elevated above base of cranium than in the smaller animal, so 
that lower edge of condyle is about on level with anteorbital 
foramen and alveolus of canine instead of distinctly below them. 
Correlated with this character is a more abrupt rising of lambdal 
region above level of anterior portion of brain-case, and a less 
nearly horizontal truncation of occipital region. Palate appar- 
ently more concave longitudinally than in any of the smaller 
European species. 

Measurements. — Adult male and female from Pisa, Italy : 
head and body, 78 and 87 ; tail, 59 and 66 ; tibia, 23-6 and 24; 
foot, 12 and 13 ; forearm, 65 and 68 ; thumb, 11 and 11 ; third 
finger, 119 and 123 ; fifth finger, 76 and 77 ; ear from meatus, 
21 and 22 ; width of ear, 23 and 24. Adult male from Venice : * 
head and body, 92 ; tail, 65 ; tibia, 26 ; foot, 14 ; forearm, 67. 
Adult female from Amsteg, Uri, Switzerland : head and body, 90± ; 
tibia, 21-6 ; foot, 13 ; forearm, 64; thumb, 9'4; third finger, 
114; fifth finger, 72. For cranial measurements see Table, p. 250. 

Specimens examined. — Pive, from the following localities : — 
Switzeeland: Amsteg, Uri, 1 (Geneva; co-type). 
Itam : Pisa, 3 (B.M. and U.S.N.M.) ; Ravenna, 1. 



Ninni, Atti Soc. Ital. Sci, Nat., Milano, xxvi, p. 109, 1883. 



NYCTALUS 245 

Bemarks. — Nijctalus maximus is readily distinguished among 
the bats of Europe by its large size. It is at present a rare and 
little known animal. 

9 al. Pisa, Italy. Dr. A. Senna (b). 94. 6. 3. 3. 

<5 al. Ravenna. Florence Museum (b). 85. 7. 6. 9. 

6 al. Lidth de Jeude Coll. 67. 4. 12. 337. 

NYCTALUS NOCTULA Schreber. 

1774. Vespertilio noctula Schreber, Saugthiere, I, pi. lii ; description, i, 

p. 166, under name : Die Speckmaus (Prance ; based primarily 

on "La Noctule" of Daubenton, Hist. Acad. Boyale des Sci., 

Paris, 1759, p. 376. 1765). 
1776. Vespertilio lardarius P. L. S. Miiller, Natursyst. Suppl. u. Regist.- 

Band, p. 15 (Prance; based primarily on Schreber's Speckmaus). 
1789. [Vespertilio] magnus Borkenhaut, Syn. Nat. Hist. Great Britain and 

Ireland, i, p. 1 (Cambridge, England; based on Pennant, Brit. 

Zool., No. 38). 
1789. Vespertilio altivolans White, Nat. Hist, and Antiquities of Selborne, 

p. 93 (Selborne, Hampshire, England). 
1816. ? Vespertilio major Leach, Syst. Catal. Spec. Indig. Mamm. and 

Birds Brit. Mus. (Willoughby Society reprint, 1892), p. 5 Nomen 

nudum : " Great Bat." 
1818. Vespertilio proterus Kuhl, Ann. Wetterau, Gesellsch. Naturk, rv 

(= Neue Ann., i), pt. 1, p. 41 (Substitute for noctula). 
1829. Vespertilio rufescens Brehm, Isis, p. 643 (Jena, Thiiringen, Germany). 
1841. Vespertilio noctula Bonaparte, Iconogr. Faun. Ital., i, Ind. Distrib. 

(sp. illustr.). 
1841. Noctula serotina Bonaparte, Iconogr. Faun. Ital., I, Ind. Distrib. 

(nomencl. moderna). 
1844. Vesp[ertMo] palustris Orespon, Faune Meridionale, i, p. 22 (marshes 

near Nlmes, Gard, France). 
1857. Vesperugo noctula Blasius, Saugethiere Deutschlands, p. 53. 
1869. [Vesperugo noctula] var. minima Fatio, Faune Vert. Suisse, i, p. 58 

(Geneva, Switzerland). 
1878. Vesperugo noctula Dobson, Catal. Chiropt. Brit. Mus., p. 212. 
1897. Pterygistes noctula Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, 

p. 384, October, 1897. 
1910. Nyctalus noctula Trouessart, Faune Mamm. d'Europe, p. 18. 

Type locality. — France. 

Geographical distribution. — Europe from southern Norway 
and central Sweden to the Mediterranean, and from England 
eastward into Asia. 

Diagnosis. — Condylobasal length of skull, 17 ■ 4 to 19 -4 mm. ; 
forearm, 47 to 55 mm. ; general colour dark yellowish brown, the 
hairs not darker at base. 

External characters. — General form robust and heavy. Muzzle 
broad, with conspicuous glandular swelling between eye and 
nostril, the greatest width across this region decidedly more than 
distance between nostril and ear ; nostrils projecting forward and 
outward with evident median concavity between them, the orifice 
crescentic. Ear short, extending when laid forward about half 



246 



CHIEOPTBRA 



way from eye to nostril, its breadth when flattened decidedly 
greater than height above crown ; anterior border of conch 
abruptly convex below, then nearly straight to broadly rounded 
off extremity ; posterior border convex throughout, most strongly 
at middle ; antitragus long and low, well marked off posteriorly, 
its anterior border extending to just below angle of mouth ; inner 
surface of conih finely papillose, without evident cross ridges ; 
tragus very short, scarcely rising above level of anterior base of 
conch, much wider above than below, its greatest width about 
equal to height ; anterior and posterior borders concave, the 
anterior more so than posterior, their length about equal to that 
of expanded upper portion, which is more abruptly rounded 
anteriorly than posteriorly. Wing long and slender, the fifth 
finger exceeding forearm by only ^ to ^ length of latter, the 
membrane leathery and opaque, joining leg at ankle ; third and 
fourth metacarpals sub-equal, the third slightly the longer and 
about equal to forearm, fifth abruptly shorter by slightly more 
than one-fifth forearm. Leg short and strong, the broad foot 
about one-half as long as tibia ; calcar 1^ to twice as long as 
tibia or as free border of interfemoral membrane, its distal 
termination obscure but basal portion strong, well defined and 
\vith keel about 2-5 mm. wide supported by a well defined 
thickening and terminating in an abrupt convexity on side 
nearest heel. Tail rather short, extending to between shoulders 
when laid forward, the tip of last vertebra (about 2 mm.) free. 

Fur and colour. — Fur dense and velvety, the hairs at middle 
of back only about 5 mm. in length ; on dorsal surface of wing 
it extends to line joining knee with middle of humerus, on ventral 
surface to line from elbow to knee ; beyond this line it spreads 
thinly on under surface of antebrachial membrane and also 
behind forearm and across bases of metacarpals along an area 
about 10 mm. wide. Interfemoral membrane furred both above 
and below at extreme base only. Colour of upper parts a rather 
dark yellowish brown, near the wood-brown and cinnamon of 
Ridgway, the hairs showing a faint clouding of prouts-brown in 
certain lights, their basal portion a light dull isabella-colour ; 
underparts scarcely different from back, though usually a little 
lighter and more dull ; muzzle and cheeks dusky ; ears and 
membranes blackish. 

Skull. — General aspect of skull broad and robust, rather high 
posteriorly, low anteriorly, with conspicuous narial emargination. 
Dorsal profile rising rather rapidly from incisors to lambda, with 
slight convexity over lachrymal region and a slighter though 
more abrupt concavity just in front of lambda. Ventral profile 
elevated posteriorly. Brain-case ovate, but with wide mastoid 
and paroccipital region and squarely truncate occiput which 
together produce a distinctly truncate-cuneate outline, slightly 
though evidently wider than long ; depth at middle distinctly 
more than half mastoid breath ; sagittal crest low but evident, 



NYCTALUS 



247 




the region on each side of it flat or slightly depressed ; lambdoid 
crest high, curved abruptly forward at point of junction with 
sagittal crest ; floor of brain-case with small but evident lateral 
pits, well denned anteriorly but communicating posteriorly with 
vacuity between cochlea and basioccipital ; auditory bullae well 
developed but of moderate size, the transverse diameter about 
equal to distance between bullae. Interorbital region deeply 
constricted, short hour-glass shaped, the lachrymal region widen- 
ing abruptly to a breadth nearly equal to that of brain-case, and 
forming a slight though evident tuber- 
cular projection over anterior rim of orbit ; 
rostrum squarish, somewhat narrower 
anteriorly than posteriorly, the oval or 
ovate narial emargination extending back 
to level of lachrymal foramen ; rostral 
depth at front of orbit about equal to 
distance from orbit to front of inner 
incisor ; anteorbital foramen small, over 
space between large premolar and first 
molar, lachrymal foramen slightly above 
and behind it, on orbital rim. Palate 
rather short, owing to the large size of 
the anterior emargination, the posterior , """ "'. , 

, ,, .... ° -. , ... ■ i ii j? Nyctalus noctvZa. 

edge ot which is on level with middle ot Nat. size, 

large premolar, rather evidently concave 

laterally, less so longitudinally. Posterior extension of palate 
nearly parallel sided, its width at level of posterior edge of third 
molar slightly less than length ; hamulars barely turned inward ; 
median spine large, acute. Mandible robust, noticeably deeper 
at symphysis than immediately behind tooth-row ; posterior 
portion rather low and long, the height of coronoid process 
above alveolus less than least length, the upper border sloping 
gradually from coronoid to articular process ; angular process 
moderately long, about on level with alveolar line, its main 
axis directed rather abruptly outward, its distal extremity 
obliquely widened. 

Teeth. — Relatively to size of skull the teeth are large, though 
the cusps are rather low. Inner upper incisor slender, with 
slightly developed cingulum, its crown area barely one quarter 
that of canine, the nearly terete shaft directed obliquely inward, 
its apex extending slightly beyond level of cingulum of canine, 
its secondary cusp small but evident, situated on postero-external 
surface of shaft ; outer upper incisor with crown area distinctly 
greater than that of inner tooth, the shaft deeply concave, its 
concavity directed outward and backward, the main cusp lying close 
against secondary cusp of inner tooth, the well-defined se-.iondary 
cusp relatively much larger than that of inner incisor, cingulum 
well developed, bearing a small but distinct antero-external cusp 
on opposite side of concavity from secondary cusp. The two 



248 



CHIROPTEEA 



incisors are closely crowded against each other ; outer tooth 
separated from canine by a very narrow space. Lower incisors 
rather strongly imbricated, forming a very broadly (J shaped 
row between canines, the crowns of i l and i 2 compressed, trifid, 
that of i 3 sub-terete, flattened anteriorly, with two low posterior 
tubercles in addition to the three cusps corresponding to those of 
other teeth. Upper canine robust, with well developed anterior 
and posterior cutting edge, the shaft triangular in cross-section ; 
cingulum well developed but without true secondary cusps. 
Lower canine with posterior surface of shaft strongly concave, 
the well developed cingulum forming a postero-internal basal 
cusp and a conspicuous secondary cusp near middle of antero- 
internal border of shaft ; diameter of crown distinctly less than 
distance between canines. Anterior upper premolar minute, closely 
wedged in space between canine and posterior premolar, its cross- 
section less than half that of incisors, its crown reniforni in 
outline, its cusp low but distinct ; posterior upper premolar with 
crown area somewhat more than half that of first molar, the 
protocone well developed, nearly as high as in molars, the main 
cusp rather high but much shorter than canine (measured along 
cingulum), its two cutting edges well developed. Lower pre- 
molars closely crowded, their crowns about equal in cross-section 
and somewhat more than half that of canine, the shaft of first 
lower and more robust than that of second and about half as 
high as canine ; cingulum well developed, in each tooth forming 
a small but evident antero-internal and postero-internal cusp. 
First and second upper molars with crowns rather broad on 
inner side, the posterior emargination slight ; protocone robust 
but low ; hypocone small but well developed and distinct from 
commissure of protocone ; outer cups without special peculiarities, 
the styles low but well developed, the W -pattern normal, m 3 with 
crown area distinctly more than half that of m 1 , its longitudinal 
diameter through metacone about half transverse diameter, its 
three cusps, two styles and three commissures well developed, 
but proportion between paracone and metacone reversed as 
compared with other molars. Lower molars with unusually 
robust hypoconid and entoconid, producing a strong contrast in 
size between the anterior and posterior V in ™i and m 2 an£ l 
approximate equality in m 3 ; cingulum well developed but not 
forming basal cusp behind entoconid. 

Measurements. — Adult female from Kew Gardens, Surrey, 
England : head and body, 71 ; tail, 51 ; tibia, 18 ; foot, 10 ; 
forearm, 52 ; thumb, 9 ; third finger, 94 ; fifth finger, 58 ; ear 
from meatus, 15 ; width of ear, 16. Adult female from Herrn- 
hut, Saxony, Germany : head and body, 79 ; tail, 53 ; tibia, 19 ; 
foot, 10 - 6; forearm, 53; thumb, 9 - 4; third finger, 97; fifth 
finger, 60 ; ear from meatus, 17 ; width of ear, 17. Two adult 
males from Pisa, Italy : head and body, 70 and 71 ; tail, 49 and 
52-6 ; tibia, 18 and 19 ; foot, 10 and 11 ; forearm, 51 and 53 ; 



NYCTALUS 249 

thumb, 8 and 9 ; third finger, 92 and 97 ; fifth finger, 56 and 59 ; 
ear from meatus, 14*6 and 16 ; width of ear, 14 and 15 '6. Two 
adult females from the same locality : head and body, 74 and 75 ; 
tail, 50 and 53 ; tibia, 18 • 6 and 18 ; foot, 10 and 11 ; forearm, 
51-6 and 52 ; thumb, 9 and 10 ; third finger, 93 and 97 ; fifth 
finger, 57 and 58; ear from meatus, 16 and 16; width of ear, 
16 and 16. Two adult females from Corinth, Greece : head and 
body, 69 and 76 ; tail, 51 and 54; tibia, 18 and 19; foot, 11 
and 10 ; forearm, 52 and 54 ; thumb, 9 ■ 4 and 9 • 4 ; third finger, 
96 and 102 ; fifth finger, 58 and 62 ; ear from meatus, 15" 6 and 
16 ; width of ear, 15 and 16. For cranial measurements see 
Table, p. 250. 

Specimens examined. — One hundred and fourteen, from the following 
localities : — 

England : Bowdon, Cheshire, 1 ; Stoke, Staffordshire, 1 ; Oundle, 
Northamptonshire, 4 ; Bedfordshire, 1 ; Trumpington, Cambridgeshire, 1 ; 
Cambridgeshire, no exact locality, 1 ; Henley-on-Thames, Oxfordshire, 1 ; 
Sandringham, Norfolk, 1 ; Kingsbury, Middlesex, 1 ; Chelmsford, Essex, 1 ; 
Wellyn, Hertfordshire, 1 ; Tring, Hertfordshire, 3 ; Wandsworth Common, 
Surrey, 1 ; Shalford, Surrey, 2 ; Earlsfield, Surrey, 5 ; Kew Gardens, 
Surrey, 1 (U.S.N.M.) ; Fulham Park, London, 1 ; Eastwell, Kent, 2 ; St. 
Leonards, Sussex, 1 ; Farnborough, Hampshire, 1 ; Wareham, Dorset- 
shire, 1. 

Holland : Oosterbeek, Guelderland, 1. 

Germany : Moritzburg, Saxony, 3 (U.S.N.M.) ; Herrnhut, Saxony, 2 
(B.M. and U.S.N.M.) ; Niesky, Silesia, 1 ; Schwarzburg, Thiiringen, 2 ; 
Strass, near Burgheim, Bavaria, 3 ; Bavaria, no exact locality, 8 (U.S.N.M.) ; 
Stuttgart, 1. 

Austeia-Hungary : Csall6k6z-Somorja, Pressburg, Hungary, 2. 

Roumania : Gageni, 1 ; Bustenari, 1 ; no exact locality, 1 (U.S.N.M.). 

Switzerland : Geneva, i (Geneva and Mottaz) ; St. Margarethen, 
Appenzell, 8 (B.M. and U.S.N.M.). 

Spain : Seville, i ; no exact locality, 2. 

Italy : Pisa, 18 (U.S.N.M.) ; Florence, 4 (Mottaz) ; Vallombrosa, near 
Florence, 1 (U.S.N.M.) ; Siena, 1 ; Rome, 2. 

Greece : Corinth, 12 (U.S.N.M.). 

Remarks. — This is one of the most widely distributed and 
abundant of the larger European bats. From Eptesicus serotinus, 
the only species with which it is likely to be confused, it is easily 
distinguished by the narrow wing, short, recurved tragus, and 
low, rounded ear. 

$. Aberia, Merionethshire, Col. Yerbury (c & p). 

"Wales. 

?. Bowdon, Cheshire, Eng- G. Barrett-Hamilton 

land. (T. A. Coward.) (p). 

2 <$, 2 9. Oundle, Northampton- Lord Lilford (p). 
shire. 

9.' Bedfordshire. J. S. Elliot (c & p). 

1. Trumpington, Cambridge- G. Barrett-Hamilton 
shire. (p). 

9 al. Cambridgeshire. Rev. L. Jenyns (c & p). 



11. 


1.3. 


26. 


11. 


1.2. 


100. 


11. 


1.1. 


126- 


127. 




11. 


1.1. 

L30. 


129- 


U. 


1. 3. 


391. 


11. 


1.2. 


101. 



250 



CHIEOPTERA 



SO 

5 

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co 

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Teeth slightly worn. 

„ not worn. 
„ slightly worn. 

„ moderately worn. 
„ slightly worn. 
„ moderately worn. 
„ much worn. 
,, not worn. 
„ slightly worn. 

„ not worn. 

„ moderately worn. 

„ slightly worn. 




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N. maximus. 

Switzerland : Amsteg, Uri . 
Italy: Pisa .... 

Ravenna . 

N. noctula. 

England : Wellyn, Hertfordshire 
Shalford, Surrey 
Earlsfield „ 

33 33 ■ 
33 33 • 

Kingsbury, Middlesex 
Oundle, Northampton 

33 33 

Chelmsford, Essex 

Holland : Oosterheek, Guelder-1 

land . . . ) 

Germany: Sohwarzburg, Thiir-1 

ingen . . ./ 

33 33 



NYCTALUS 



251 



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252 



CHIROPTEEA 



<5. Henley-on-Thames, Ox- 
fordshire. 

£ st. Sandringham, Norfolk. 

9. Chelmsford, Essex. 

?. Wellyn, Hertfordshire. 

2 i, 9 st. Tring, Hertfordshire. 

2 S. Shalford, Surrey. 

3 <5, 2 9. Barlsfield, Surrey. 



(J st. 



Fulham Park, London. 
Eastwell, Kent. 



9. St. Leonards, Sussex. 

6. Farnborough, Hampshire. 

9. Wareham, Dorset. 

6. Oosterbeek, Guelderland, 

Holland. 
9. Niesky, Silesia, Germany. 

(W. Boer.) 
6. Schwartzburg, Thiiringen, 

200 m. 
3 c?. Strass, Burgheim, 

Bavaria. (Kiirbitz.) 

1 al. Stuttgart, Wurtemburg. 
2. Csall6k6z-Somorja, Press- 
burg, Hungary. 

S. Gageni, Prahova, Bou- 

mania. (W. Dodson.) 

9. Bustenari, Prahova. (W. 

Dodson.) 

2<5,39. St. Margarethen, 

Appenzell, Switzer 

land. (E.H.Zollikofer. 

4 al. Seville, Spain. (A. Ruiz. 

2 al. Spain. 
S juv. al. Borne. 



Heatley Noble (c & p). 
H.M. King Edward VII. 

M. Christy and L. E. 

Thompson (o & p). 
C. H. B. Grant (c & p). 
Hon. N. C. Rothschild 

(o & p). 
W. B. Ogilvie - Grant 

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

J. Saunders (o & p). 
C. H. Caton - Haigh 

(c & p). 
B. Butterfield (c & pi. 
H. S. Soott (c & p). 
W. T. Blanford (c & p). 
0. Thomas (o & p). 

Lord Lilford (p). 

Lord Lilford (p). 

Lord Lilford (p). 

Dr. A. Giinther (p). 
Budapest Museum (e). 

Lord Lilford (p). 

Lord Lilford (p). 

0. Thomas (p). 



Seville Museum (e). 
Lord Lilford (p). 
Florence Museum (e). 



11. 1. 3. 40. 
96. 7. 27. 1. 
11. 1. 3. 41. 

11. 1. 3. 33. 

I. 5. 22. 1-3. 

II. 1. 3. 34- 
35. 

11. 1. 3. 28- 

32 
87. 7. 22. 1. 
11. 1. 3. 36- 

37. 
11. 1. 3. 392. 
11. 1. 3. 39. 
11. 1. 3. 38. 

98. 2. 1. 5. 

99. 1. 9. 7. 
95. 4. 18. 1. 

11. 1. 1. 30- 

32. 
66. 2. 1. 6. 
94. 3. 1. 16- 

17. 
4. 4. 6. 4. 

4. 4. 6. 5. 

4. 4. 5. 15-19. 



94. 5. 8. 1-4. 
72. 8. 21. 1-2. 
85. 7. 6. 8. 



NYCTALUS LEISLERI Kuhl. 

1818. Vespertilio leisleri Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv 

(= Neue Ann., I), pt. 1, p. 46. 
1818. Vespertilio dasycarpos Kuhl, Ann. Wetterau. Gesellsch. Naturk., iv 

(= Neue Ann., i), pt. 1, p. 49 (Alternative name for leisleri). 
1839. Vespertilio pachygnathus Michahelles in Wagner, Schreber's Saug- 

thiere, Suppl., i, pi. lv b (Dalmatia. See Pitzinger, Sitzungsber. 

kais. Akad. Wissensch. Wien, Math.-Naturwiss. Classe, lxii, 

Abth. I, p. 222, 1870). 
1857. Vesperugo leisleri Blasius, Saugethiere Deutschlands, p. 56. 
1878. Vesperugo leisleri Dobson, Catal. Chiropt. Brit. Mus., p. 215. 
1910. Nyctalus leisleri Trouessart, Paune Mamm. d'Europe, p. 19. 

Type locality. — Hanau, Hessen-Nassau, Germany. 
Geographical distribution. — Central Europe, west to Ireland. 
Diagnosis. — Like Nyctalus noctula but not so large ; condylo- 



NYCTALUS 253 

basal length of skull, 15 to 16 mm. ; forearm, 39 to 43 mm. ; 
basal portion of fur conspicuously darkened. 

Colour. — The colour is usually a brown, darker than in N. 
noctula, nearly the prout-brown of Ridgway, with or without a 
lighter, wood-brown tinge, and usually showing drab or isabella 
reflections in certain lights, the underparts not so dark as back ; 
hairs everywhere blackish (seal-brown) through basal half. Ears 
and membranes (dry) blackish. Two skins are a pale buffy wood- 
brown throughout except for the usual dark bases of the hairs. 

Slcull and teeth. — Except for its smaller size and more delicate 
structure the skull does not differ appreciably from that of 
Nyctalus noctula. Teeth essentially as in the larger species, but 
crown area of upper incisors nearly equal, and lower incisor row 
forming a deeper, almost V-shaped convexity. 

Measurements. — External measurements of adult male from 
Co. Wicklow, Ireland : head and body, 60 ; tail, 39 ; tibia, 16-2 ; 
foot, 8 • 2 ; forearm, 42 ; third finger, 78 ; fourth finger, 48 ; ear 
from meatus, 13. Adult female from Co. Armagh, Ireland : 
head and body, 63 ; tail, 42 ; tibia, 16 • 6 ; foot, 8 ; forearm, 42 ; 
third finger, 76 ; fifth finger, 49 ; ear from meatus, 14 ; width 
of ear, 14. Two adults from Welford, Warwickshire, England : 
forearm, 40 • 4 and 41-4. Adult from Meiringen, Bern, Switzer- 
land : forearm, 40. For cranial measurements see Table, 
p. 255. 

Specimens examined. — Eleven, from the following localities : — 
Ibeland : Belfast, 1 ; Bray, Co. Wicklow, 2 ; Co. Armagh, 2. 
England : Mexbro', Yorkshire, 1 ; Welford, Warwickshire, 2 ; Cleeve 
Priory, 1 ; no exact locality, 1. 

Roumania : Bustenari, Prahova, 1. 
. Switzerland : Meiringen, Bern, 1. 

Remarks. — Though its smaller size is the most obvious 
character of this species as compared with N. noctula, the 
bicolored fur, the different relative sizes of the upper incisors, 
and the more abrupt convexity of the mandibular incisor series 
would by themselves be quite sufficient to distinguish it. 



9. 


Belfast, Ireland. 


G. Barrett-Hamilton (p.) 


1. 3. 15. 1. 


<5al. 


Bray, Wicklow. 


J. E. Harting (c & p). 


90. 2. H. 1. 


Sal. 


Armagh. 


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


89. 11. 12. 5, 


Sal. 


Armagh. 


R. M. Barrington (c & p). 


74. 5. 28. 8. 


2 9. 


Welford, Warwickshire, 


Tomes Collection. 


7. 1. 1. 386- 




England. 




387. 


6. 


Cleeve Priory, Warwick- 
shire. 


Tomes Collection. 


7. 1. 1. 388. 


1. 


England. 


Dr. Leach (p). 


63. A. 


i. 


Bustenari, Prahova, 840 m. 
Roumania. (W.Dodson.) 


Lord Lilford (p). 


4. 4. 6. 6. 


1, 


Meiringen, Bern, Switzer- 
land. 


Tomes Collection. 


7. 1. 1. 389. 



254 



CHIEOPTEEA 



NYCTALUS AZOREUM Thomas. 



1901. Pterygistes azoreum Thomas, Ann. and Mag. Nat. Hist., 7th ser., vm, 
p. 33, July, 1901. Type in British Museum. 

Type locality. — St. Michael, Azores. 

Geographical description. — Azores. 

Diagnosis. — Smaller than Nyctalus leisleri (condylobasal length 
of skull, 13 to 14 • 2 mm. ; forearm, 35 to 42 mm.) ; colour darker 
than in the Continental European species. 

External characters. — Except for its conspicuously smaller 
size Nyctalus azoreum does not differ appreciably in external 
features from N. noctula. The fifth finger shows a tendency to 
be less shortened relatively to length of forearm, but this is appa- 
rently not constant enough to be regarded as a specific character. 

Colour. — Upper parts prout-brown, the hairs with darker, 
ill-defined slaty basal area, and usually with lighter tips, the 
lighter colour ranging in different individuals from raw-umber 
through wood-brown to a dull light buff, and usually producing 
a noticeable wash from shoulders backward, particularly near 
edge of membranes ; underparts not essentially different from 
back ; muzzle, cheeks, ears, and wings blackish. 

Shull and teeth. — The skull does not differ appreciably in form 
from that of N. noctula, though it is perhaps slightly narrower 
and deeper, and the surface is more smoothly rounded off; 
lambdoid crest slightly developed. Teeth essentially as in the 
larger species, but transverse diameter of m 3 relatively greater, 
crown area of upper incisors nearly equal, and lower incisor row 
forming a more abrupt convexity. 

Measurements. — Type (adult male, St. Michael) : head and 
body, 54 • tail, 42 ; tibia, 17 ; foot, 7 ■ 7 ; forearm, 37 ; thumb, 6 ; 
third finger, 62 ; fifth finger, 43 • 7 ; ear from meatus, 12. Adult 
male from Terceira : head and body, 50 ; tail, 43 ; tibia, 16 ■ 6 ; 
foot, 7 ; forearm, 42 ; thumb, 6 ; third finger, 72 ; fifth finger, 
48 ; ear from meatus, 12 ; width of ear, 11. For cranial measure- 
ments see Table, p. 255. 

Specimens examined. — Seventeen, from the following localities in the 
Azores : — Teroeira, i (B.M. and U.S.N.M.) ; above Magdalena, Pico, 5 ; St. 
Michael, 6 (B.M. and U.S.N.M.) ; St. George, 2. 

Remarks. — The Azorean Nyctalus is slightly more differentiated 
from N. leisleri than the Continental forms are among themselves, 
since it differs from the other members of the genus rather 
noticeably in colour as well as in size. The specimens show 
considerable variation in the extent of the light wash on upper 
parts caused by the pale tips to the longer hairs, but this appears 
to be independent of locality. 

<5 al. St. Michael, Azores. F. Du Cane Godman 65. 10. 2. 1. 

(o & p). (Type of species.) 
2(5,2?. St. Michael. (W. B. 0. Hon. W. Eothschild 3.6.5.1-3,7. 
Chant.) (p). 



255 



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St. Michael 



256 



CHIROPTEEA 



<5, 2 9. Reguinho, Teroeira. Hon. W. Rothschild 3.6.5.4-6. 
(W. B. 0. Grant.) (p). 

2 1, 3 9. Magdalena,Pioo. (W.B.O. Hon. W. Rothschild 3.6.5.10-14. 
Grant.) (p). 

<5,S. Calheto, St. George. Hon. W. Rothschild 3.6.5.8-9. 
(W. B. 0. Grant.) (p). 



Genus PLECOTUS Geoffroy. 

1816. ? Macrotus Leach, Catal, Spec. Indig. Mamm. and Birds Brit. Mus. 
(Willoughby Society reprint, 1882), p. 1. Nomen nudum: "Euro- 
pean Longear, Macrotus europasus." 

1818. Plecoius Geoffroy, Description de l'Egypte, n, p. 112. For date see 
Sherborn. Proc. Zool. Soc. London, 1897, p. 288. 

1857. Plecotus Blasius, Saugethiere Deutschlands, p. 38. 

1878. Plecotus Dobson, Catal. Chiropt. Brit. Mus., p. 177 (part). 

1907. Plecotus Miller, Families and Genera of Bats, p. 224, June 29, 1907. 

Type species. — Vespertilio auritus Linnaeus. 

Geographical distribution. — Temperate Europe, Asia and 
northern Africa. 

Characters. — Dental formula : i *=|, c Jd, pm ||, m || = 36. 
Skull with large, elongate and rounded brain-case, and slender 
but normally formed rostrum ; auditory bullse larger than in any 
other European member of the family, their greatest diameter 
equal to about three times the distance between them. Ears 
very large, much longer than head, joined across forehead ; 
nostrils opening upward, their orifices continued backward by 
slit-like prolongations, the wart-like outgrowths on muzzle not 
specially prominent. 

Remarks. — The genus Plecotus is at once recognizable among 
European bats by the very large ears, joined across forehead. 
In the only other European genus with 36 teeth (Miniopterus) 
the ears are unusually short. Seven species are known, one of 
which occurs in Europe. 

PLECOTUS AURITUS Linnaeus. 

1758. Vespertilio auritus Linnseus, Syst. Nat., i, 10th ed., p. 32 (Sweden). 

1816. ? Macrotus europseus Leach, Catal. Spec. Indig. Mamm. and Birds 
Brit. Mus. (Willoughby Society reprint, 1882), p. 1. Nomen 
nudum: " European Longear, Macrotus europseus." 

1818. Plecotus auritus Geoffrey, Description de l'Egypte, n, p. 118. 

1825. Vespertilio otus Boie, Isis, p. 1206 (Copenhagen, Denmark). 

1826. Vespertilio cornutus Eaber, Isis, p. 515 (Jutland, Denmark). 

1827. Plecotus communis Lesson, Man. de Mammal., p. 95 (Prance). 

1828. P[lecotus] brevimanus Jenyns, Trans. Linn. Soc, London, xvi, p. 55 

(Grunty Pen, Isle of Ely, Cambridgeshire, England). 

1829. Plecotus vulgaris Desmarest, Paune Prancaise, p. 18 (France). 



PLECOTUS 257 

1829. [Vespertilio auritus] y austriacus Fischer, Synops. Mamm., p. 117. 

Based on the " second variety" of Desmarest, Diet, des Sci. Nat., 

lviii, p. 51, 1829 (Vienna, Austria). 
1832. Plecotus velatus I. Geoffroy, Guerin's Mag. de Zool., n, CI. i, pi. 2, 

p. 5 (not numbered), footnote (Name applied by lapsus calami to 

the common Plecotus of England). 
1840. Plecotus megalotos Schinz, Europ. Fauna, i, p. 19 (Synonym of 

auritus ; Brehm cited as authority). 
1857. Plecotus auritus Blasius, Siiugethiere Deutschlands, p. 39. 
1860. Plecotus kirschbaumii Koch, Achter Ber. Oberhess. Gesellseh. 

Natur.- u. Heilkunde, Giessen, p. 40, May, 1860 (Dillenburg, 

Oberhessen, Germany). 
1863. [Plecotus auritus] var. typus Koch, Jahrb. des Vereins flir Natur- 

kunde im Herzogthum Nassau, xviii, p. 406 (Wiesbaden, Nassau, 

Germany). 
1863. [Plecotus auritus] var. montanus Koch, Jahrb. des Vereins fiir Natur- 

kunde im Herzogthum Nassau, xviii, p. 406 (Westerwald, Nassau, 

Germany). 
1863. [Plecotus auritus] var. brevipes Koch, Jahrb. des Vereins fiir Natur- 

kunde im Herzogthum Nassau, xviir, p. 407 (Substitute for kirsch- 
baumii Koch). t 
1878. Plecotus auritus Dobson, Catal. Chiropt. Brit. Mus., p. 178. 
1910. Plecotus auritus Trouessart, Faune Mamm. d'Europe, p. 12. 

Type locality. — Sweden. 

Geographical distribution. — Prom Ireland eastward into Asia, 
and from the Mediterranean north to the Scandinavian Peninsula. 

Diagnosis. — Characters as in the genus ; auditory bulla? 
not excessively enlarged ; forearm about 37 mm. 

External characters. — General form slender and delicate, the 
most conspicuous feature the greatly enlarged ears, the super- 
ficial area of which together when outstretched is considerably 
greater than that of entire body. Muzzle rather narrow, its 
width at middle about equal to distance from tip to point on 
forehead at middle of base of joining membrane of ears ; glandular 
swellings not conspicuous, but extending up over sides to median 
line behind nostrils, and backward to eye. Nostrils opening 
upward and slightly forward, their orifices crescentic with slit- 
like posterior elongation ; space between nostrils flat, crossed 
at middle by a transverse groove with overhanging anterior edge. 
Lips moderately full, smooth. Ear very large, extending about 
20 mm. beyond tip of muzzle when laid forward, its length from 
meatus nearly equal to that of forearm : general outline of conch 
a simple, rater elongate oval, the anterior and posterior borders 
evenly convex, the tip rather narrowly rounded ; anterior bases 
joined across forehead, the membrane at point of junction about 
2-5 mm. high ; outline of anterior border broken about 3 mm. 
above base by conspicuously projecting, evenly rounded lobe 
2 mm. long by 3 mm. wide at base ; antitragus scarcely indicated, 
the posterior border of conch terminating abruptly a little behind 
level of eye ; inner surface of conch with about twenty faint 
cross striations ; joining membrane thinly furred ; a hairy line 



258 



CHIROPTEKA 



parallel with anterior border, and minute scattered tufts on other 
parts of inner surface ; tragus simple, erect, about half as high 
as conch, the anterior border straight below, slightly convex 
above, the tip narrowly rounded, the posterior border faintly 
concave above, distinctly convex below, greatest width (through 
anterior base) equal to a little less than half length of anterior 
border ; posterior basal lobe well developed though not large. 
Wing broad, the fifth finger exceeding forearm by more than 
one-third length of latter, the membrane thin and semi-trans- 
parent, joining foot at base of outer toe ; third and fourth meta- 
carpals equal, about 2 mm. shorter than forearm, fifth slightly 
shorter than fourth ; leg slender ; foot about half tibia ; calcar 
slender but sharply defined, equal in length to tibia and to free 
border of interfemoral membrane, its distal termination marked 
by a slight lobe, its keel obsolete. Tail extending about to point 
of juncture of ears when laid forward, the extreme tip (about 
1 ■ 5 mm.) free. 

Skull. — Among the European members of the order the 
skull of Plecotus is noticeable for its large, rather low, smoothly 
rounded brain-case, relatively small, weak rostrum, and much 
inflated auditory bullae. Dorsal profile rising gradually from front 
of nares to middle of anterior portion of brain-case, with slight 
concavity at interlachrymal region, then falling gradually to 
low, strongly convex, slightly overhanging lambda ; ventral 
profile scarcely elevated posteriorly. Brain-case large but rather 
low, the depth at middle only a little more 
than half mastoid breadth, the general out- 
line when viewed from above squarish with 
rounded occipital projection and abrupt 
g AaaawJiy anterior constriction, the surface smoothly 

rounded, with slightly indicated sagittal ridge 
anteriorly and very low, short lambdoid crest, 
the latter situated unusually close to upper 
edge of foramen magnum ; an irregular area 
in supramastoid region thickly sprinkled with 
small vacuities ; base of brain-case smooth, 
rh without special features, a narrow groove at 

Plecotus mmtus. each side of basioccipital ; auditory bulla 

Nat. stee. relatively larger than in any other European 

bat, its diameter about three times the 
space between bullae. Interorbital constriction narrow, scarcely 
hour-glass shaped, the anterior upper rim of orbit with narrow, 
slightly projecting edge ; lachrymal region scarcely half as wide 
as brain-case. Rostrum both narrow and short, the narial 
emargination, though not unusually large, extending slightly 
more than half way back to lachrymal level, its general outline 
oval, sometimes flattened posteriorly ; dorsal surface of rostrum 
with slight median crease, the sides evenly rounded ofi ; rostral 
depth at front of orbit about equal to distance from orbit to 





PLBCOTDS 259 

alveolus of inner incisor ; anteorbital foramen small, close to rim 
of orbit, its orifice over parastyle of first molar ; lachrymal fora- 
men slightly above and behind it, on inner edge of orbit. Palate 
rather broad and short, strongly concave both laterally and 
longitudinally, the anterior emargination small, scarcely extend- 
ing back to level of middle of canine ; posterior extension of 
palate squarish, slightly longer than wide, with slightly developed 
median spine ; mesopterygoid space somewhat wider than deep. 
Zygoma slightly but evidently expanded at middle. Mandible 
slender, but noticeably deeper at symphysis than behind tooth- 
row ; coronoid process moderately high, the upper margin of 
posterior section of mandible oblique-concave ; angular process 
rather heavy, scarcely or not expanded at tip. 

Teeth. — The teeth are rather small relatively to size of skull. 
Inner upper incisor about half as high as canine, with large 
secondary cusp slightly above middle of shaft, the outline of 
crown oval, its long axis in line of tooth-row ; outer upper incisor 
about half as high as inner, its apex a little below secondary cusp 
of larger tooth, its secondary cusp well developed, on inner side, the 
outline of crown ovate, with long axis perpendicular to tooth-row 
and narrower extremity directed inwards. Lower incisors closely 
crowded but slightly imbricated, the row broadly (J shaped ; the 
crown area increases regularly from first to third, by addition to 
postero-internal portion ; cutting edge obscurely trifid ; i 3 with 
well developed postero-internal tubercle. Upper canine small, 
scarcely higher than main cusps of molars, its shaft nearly terete, 
with slightly developed posterior cutting edge, on each side of 
which lies an evident groove ; cingulum complete, without cusps. 
Lower canine small, slightly exceeding molars in height, its 
shaft concave posteriorly, flattened interiorly, evenly convex 
antero^externally ; cingulum well developed, forming a prominent 
antero-internal secondary cusp. Anterior upper premolar 
perfectly in tooth-row, about as large as outer incisor, though 
more robust, its crown sub-terete, about one-third that pi canine 
in basal area, its shaft with evident posterior cutting edge ; large 
premolar with crown area about two-thirds that of first molar, 
the inner side very narrow and without crushing surface, the 
antero-internal cusp slender but well developed. Crown area of 
anterior lower premolar about half that of canine, that of 
posterior premolar nearly equal to that of canine, that of middle 
premolar a little more than half that of first ; first sub-terete, 
second oval with long diameter of crown lying across tooth-row, 
third squarish with antero-external corner rounded off ; cusp of 
first about half as high as canine, that of second a little shorter, 
that of third a little longer ; cingulum well developed but not 
forming true secondary cusps. Upper molars narrow internally, 
the protocone with rather short base, the hypocone absent or 
barely indicated ; metacone decidedly higher than paracone ; 
styles and commissures well developed, though mesostyle of m l 

s 2 



260 



CHIROPTEEA 



does not extend outward to level of parastyle and metastyle ; 
W-P a ttern normal ; m 3 with three cusps and three commissures, 
its crown area about half that of m 2 , its transverse diameter 
through metacone about half length of anterior border. Lower 
molars with no special peculiarities except that protoconids and 
inner cusps are unusually high and slender ; cingulum forming a 
barely indicated postero-internal cusp behind entbconid ; second 
triangle of ro 3 much narrower than first, but about equal to it 
in area. 

Measurements. — Adult female from Chelmsford, Essex, 
England: head and body, 42; tail, 45; tibia, 17; foot, 9'6; 
forearm, 39 ; thumb, 7 ; third finger, 69 ; fifth finger, 52 ; 
ear from meatus, 36 ; width of ear, 23 ; tragus, 19. Two adult 
males from Silos, Burgos, Spain : head and body, 50 and 51 ; 
tail, 46 and 46 ; tibia, 18 and 17 ; foot, 8 and 8 - 4; forearm, 
41 and 38; thumb, 7- 2 and 7-4; third finger, 72 and 70 ; fifth 
finger, 57 and 53 ; ear from meatus, 35 and 37 ; width of ear, 
26 and 26 ■ 4 ; tragus, 17 and 18. Adult female from La Granja, 
Segovia, Spain : head and body, 45 ; tail, 44 ; tibia, 20 ; foot, 9 ; 
forearm, 40 ; thumb, 9 ; third finger, 72 ; fifth finger, 53 ; ear 
from meatus, 37 ■ 4 ; width of ear, 23 ; tragus, 1 7 • 6. Adult female 
from Florence, Italy : head and body, 49 ; tail, 48 ; tibia, 19 • 2 ; 
foot, 9; forearm, 39-4; thumb, 7-8; third finger, 69; fifth 
finger, 54 ; ear from meatus, 34 ; width of ear, 23 • 4 ; tragus, 17. 
For cranial measurements see Table, p. 262. 

Specimens examined. — Eighty, from the following localities : — 

Scotland : Montrose, Forfar, 4. 

Ireland : Antrim, 1 ; Co. Longford, 2. 

England: Alderley, Cheshire, 1 ; Bowdon, Cheshire, 1 ; Diss, Norfolk, 1 ; 
Winfarthing, Norfolk, 1 ; Wohurn Sands, Bedfordshire, 1 ; Pembrokeshire, 
no exact locality, 1 ; Chelmsford, Essex, 1 (U.S.N.M.) ; Tring, Hertford- 
shire, 1 ; Boxmore, Hertfordshire, 1 ; Hoddesdon, Hertfordshire, 1 ; 
London, 4 ; Godalming, Surrey, 1 ; Surrey, no exact locality, 1 ; 
Bonchurch, Isle of Wight, 1 ; near Honiton, Devonshire, 2 ; Devonshire, 
no exact locality, 8 ; no exact locality, 1. 

Holland : Oosterbeek, Guelderland, 4. 

France : Near Barcelonnette, Basses-Alpes, 1 (Mottaz). 

Germany : Wernigerode, 3 ; Ummerstadt, Thiiringen, 2 ; Magdeburg, 2 ; 
Niesky, Silesia, 4 ; Damsdorf , Silesia, 1. 

Austria-Hungary: Csall6k6z-Somorja, Pressburg, 7. 

Switzerland : Geneva, 2 (Mottaz) ; Lausanne, Vaud, 1 (Mottaz) ; 
Morat, Fribourg, 1 (Mottaz) ; Boudry, Neuchatel, 2 (Mottaz) ; Teufen, 
Appenzell, 1 (U.S.N.M.) ; Thurgau, St. Gallen, 1 ; St. Gothard, Uri, 3 
(U.S.N.M.) ; no exact locality, 1. 

Italy: Borzoli, near Genoa, 1 (Genoa) ; Florence, 1 (Mottaz) ; Rimini, 1; 
Home, 1 ; no exact locality, 1 ; Sicily, 1. 

Spain : Silos, Burgos, 2 (U.S.N.M.) ; La Granja, Segovia, 1. 

Bemarhs. — Plecotus auritus is so readily distinguished from all 
other European bats by the great size of the ears that no special 
comparisons are required. 

<J, 3 ?. Montrose, Scotland. J. H. Coward (c & p). 11. 1. 3. 48-51 
c5 st. Antrim, Ireland. Hon. N. C. Rothschild 1. 9. 3. 5. 



PLECOTUS 261 

2 6 al. Longford. Dr. G. E. Dobson (o & p). 76. 2. 12. 2. 

<;. Pembroke, Wales. W.E. deWinton(o&p). 11. 1. 3. 393. 

1 st. Winfarthing, Norfolk, E. Markham (c & p). 96. 7. 27. 3. 

England. 

6. Diss, Norfolk. (W. E. C. H. B. Grant (p). 11. 1. 3. 394. 

Sherrin.) 

1 al. Woburn Sands, Bed- A. Death (c & p). 7. 9. 2. 1. 

fordshire. 

9. Boxmore, Hertford- H. Piffard (c & p). 11. 1. 3. 55. 

shire. 

1 st. Hoddesdon, Hertford- A. Chittenden (c & p). 86. 9. 27. 1. 
shire. 

i. Tring, Hertfordshire. Dr. E. Hartert (c & p). 11. 1. 3. 56. 
9 al. London. No history. 

6. London. Sergt. Brown (o & p). 11. 1. 3. 57. 
i. S. Kensington, Middle- 11. 1. 3. 54. 

sex. 
i. Godalming, Surrey. W. T. Blanford (c & p). 11. 1. 3. 58. 
1. Bonehuroh, Isle of Bev.C. A. Bury (c & p). 4*. 3. 29. 6. 
Wight. 
c5, 9. Honiton, Devonshire, G. C. Shortridge(c&p). 11.1.3.52-53. 
300 ft. 
4 <5, 4 9 al. Devonshire. No history. 

2 6, 2 9. Oosterbeek, Guelder- 0. Thomas (o & p). 98. 2. 1. 1-4. 

land, Holland. 
9. Wernigerode, Harz, W. Wolterstorff (c&p). 0.2.8.1. 

Germany. 
(5, 9. Wernigerode, Harz. Lord Lilford (p). 11. 1. 1. 45-46. 

(Wolterstorff.) 
S. Magdeburg, Saxony. Lord Lilford (p). 11. 1. 1. 47. 

(Wolterstorff.) 
(5,9. Ummerstadt, Thiirin- Lord Lilford (p). 11.1.1.43-44. 

gen. (Schuchardt.) 
£. Niesky, Silesia. (W. Lord Lilford (p). 99. 1. 9. 1. 

Baer.) 
6 al. Damsdorf, Silesia. Dr. A. Giinther (p). 66. 8. 1. 10. 

7. Csall6k6.z-Somorja, Budapest Museum (e). 94.3.1.5-11. 

Pressburg, Austria- 
Hungary. 
<5. Thurgau, St. Gallen, 0. Thomas (p). 4. 4. 5. 14. 

Switzerland. (Zolli- 
kofer.) 
1. Switzerland. Leon 0. Galliard (p). 75. 9. 20. 2. 

9 al. Bimini, Italy. Florence Museum (e). 85. 7. 6. 2. 

<5. Borne. (C. Coli.) G.Barrett-Hamilton (p). 11. 1. 2. 15. 

Skeleton Italy. (Prince Bona- Tomes Collection. 7. 1. 1. 728. 

(without parte.) 
skull). 

9 al. ' Sicily. Purchased (Parzudaki). 52. 2. 26. 18. 

1 al. La Granja, Segovia, M. de la Escalera (c). 8. 7. 30. 5. 
Spain. 
1. Europe. Leyden Museum (e). 37. 4. 28. 25. 



262 



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BARBASTELLA 



263 



Genus BARBASTELLA Gray. 

1821. Barbastella Gray, London Med. Repos., xv, p. 300, April 1, 1821. 

1825. Barbastellus Gray, Zool. Journ., n, p. 243, July, 1825. 

1839. Synotus Keyserling and Blasius, Wiegmann's Archiv fur Naturgesch., 

1839, I, p. 305. 
1857. Synotus Blasius, Saugethiere Deutschlands, p. 42. 
1878. Synotus Dobson, Catal. Chiropt. Brit. Mus., p. 175. 
1897. Barbastella Miller, Ann. and Mag. Nat. Hist., 6th ser., xx, p. 375, 

October, 1897. 
1907. Barbastella Miller, Families and Genera of Bats, p. 223, June 29, 

1907. 

Type species. — Vespertilio barbastellus Schreber. 

Geographical distribution. — Northern Africa ; central and 
southern Europe ; west-central Asia to the Himalayas. 

Characters. — Dental formula : i H, c J=?! pm H> i»n= 34. 
Skull with rather long, rounded brain-case and weak rostrum, the 
upper surface of which is occupied by a shallow, flattened-concave 
area extending from nares to faintly developed supraorbital 
ridges ; auditory bullse not specially enlarged. Ears broad and 
short (laid forward they reach slightly beyond tip of muzzle), 
joined across forehead ; nostrils opening upward and outward 
on a flat median space between two high lateral swellings and 
behind a prominent median pad. 

Bemarhs. — The short broad ears joined together across fore- 
head distinguish this genus from the other European members of 
the family Vespertilionid.se. Two species are known, one peculiar 
to the Himalayan region, the other occurring in Europe. 



BARBASTELLA BARBASTELLUS Schreber. 

1774. Vespertilio barbastellus Schreber, Saugthiere, i, pi. lv (description, i, 

p. 168, under name : Das Kurzmaul). Based primarily on " La 

Barbastelle " of Daubenton, Hist. Acad. Royale des Sci., Paris, 

1759, p. 377. 1765. 
1776. Vespertilio barbastelle P. L. S. Miiller, Natursyst. Suppl. u. Regist.- 

Band, p. 17 (Burgundy). 
1836. Barbastellus daubentonii Bell, Hist. Brit. Quadr., pt. i, p. 63 

(Burgundy ; based primarily on Daubenton). 
1838. Barbastellus communis Gray, Mag. Zool. and Bot., n, p. 495, 

February, 1838 (Renaming of barbastellus). 
1857. Synotus barbastellus Blasius, Saugethiere Deutschlands, p. 43. 
1878. Synotus barbastellus Dobson, Catal. Chiropt. Brit. Mus., p. 176. 
1897. Barbastella barbastellus Miller, Ann. and Mag. Nat. Hist., 6th ser., 

xx, p. 385, October, 1897. 
1910. Barbastella barbastellus Trouessart, Faune Mamm. d'Europe, p. 11, 

Type locality. — Burgundy, France. 

Geographical distribution. — Central and southern Europe, 



264 



CHIBOPTERA 



west to England, north to southern Norway and Sweden, east 
into Asia. 

Diagnosis. — Characters as in the genus ; forearm about 38 
to 40 mm. 

External characters. — General form slender and delicate, the 
legs long, the tail about equal to head and body. Muzzle short 
and broad, its width considerably greater than distance from tip 
of snout to joining membrane of ears, each side with a very large 
glandular mass rising above level of flat median dorsal surface 
and extending downward to involve most of upper lip behind 
nostril pad ; glandular masses densely hairy in noticeable contrast 
with the finely pubescent nostril pad and essentially naked 
median region between nostrils and inner bases of ears ; nostrils 
crescentic in outline, opening upward, somewhat crowded between 
glandular lateral masses ; nostril pad well defined, rounded off 
above, continued downward into median portion of upper lip, the 
edge of which, between glandular masses and separated from 
them by evident grooves, is distinctly convex, fitting like a 
valve, when mouth is closed, over bare median callosity of lower 
lip. Ear large and broad, though not specially elongated, the 
tip extending about 5 mm. beyond nostril when laid forward ; 
anterior basal lobe very small, appearing like a rudimentary 
Molossine keel, the portion of anterior border usually reflexed in 
Vespertilionine bats thrown forward so that its base joins its fellow 
of opposite ear, the region of juncture low but sufficiently well 
developed to form a distinct pocket on side next forehead ; out- 
line of anterior border strongly convex except for an evident 
flattening just below the rather abruptly rounded off and some- 
what recurved tip ; posterior border abruptly concave above, then 
nearly straight except for an abruptly projecting lobe near 
middle (lobe usually about 1 • 5 mm. long Toy 1 mm. wide, its upper 
border convex, its lower border concave) ; lower border of ear full 
and rounded, but with no differentiated antitragus ; tragus large, 
somewhat triangular in outline, its greatest width (slightly above 
level of anterior base) equal to about two-thirds length of anterior 
border, its upper portion rapidly narrowing to an attenuate tip ; 
meatus with well developed keel-like ridge ; outer surface of ear 
densely furred except at extreme tip and along posterior border 
to a little below level of projecting lobe ; inner surface with a 
hairy line marking juncture of anterior border of conch with 
portion usually folded backward, elsewhere irregularly sprinkled 
with fine hairs. Membranes thin and delicate, the wing broad, 
inserted at base of outer toe, the antebrachial membrane con- 
tinued outward as a very narrow fold to base of thumb ; third 
metacarpal slightly shorter than forearm and a little exceeding 
the sub-equal fourth and fifth ; fur soft and loose, the hairs on 
middle of back about 10 mm. in length; on both upper and 
under surface of wing the fur extends to a line joining middle of 
forearm with knee ; foot less than half as long as tibia ; calcar 



BARBASTELLA 265 

about as long as tibia, with small but evident terminal lobe and 
posterior keel ; tail with extreme tip projecting beyond inter- 
femoral membrane. 

Colour. — General colour a very dark brown between the seal- 
brown and vandyke-brown of Ridgway, the hairs everywhere 
dark brown to extreme base, those of upper parts tipped with 
light glossy wood-brown producing a sharply contrasted wash 
throughout region behind shoulders, those of underparts tipped 
with a paler, more drabby brown, but without producing so 
decided a contrast as on back, though the wash is usually evident 
along middle of belly ; hairs on base of under side of interfemoral 
membrane pale ecru-drab or whitish smoke-grey nearly or quite 
to base. Ears and membranes in dry specimens brown, hardly 
so dark as ground colour of body. 

Skull. — The skull, though lightly-built and small, scarcely 
exceeding that of Myotis mystacinus in length, is rather deep and 
robust, with unusually large brain-case relatively to the short 
rostrum. Dorsal profile abruptly convex over anterior half of 
brain-case, then nearly horizontal to somewhat overhanging 
occiput ; ventral profile nearly horizontal, the floor of brain-case 
scarcely elevated above level of tooth-row. Brain-case high and 
inflated anteriorly, relatively low posteriorly, its lateral outlines 
when viewed from above essentially as in the 
species of Myotis, but less rounded and globular 
posteriorly ; floor of brain-case with no special 
peculiarities; auditory bullse scarcely more in- 
flated than in the small species of Myotis. 
Interorbital region slightly constricted, its sur- 
face flattened and not well defined from that of 
rostrum ; lachrymal ridges rather prominent, 
especially at their lower extremities. Zygoma 
nearly straight, scarcely bowed outward beyond 
level of outer surface of brain-case. Ante- n , 

orbital foramen relatively large, over anterior Barbastdla barba . 
margin of m 1 . Rostrum short, its upper sur- status. Nat. size, 
face flattened-concave, deeply emarginate in 
front by the unusual backward extension of the upper margin of 
anterior nares, the posterior border of which is at level of ante- 
orbital foramina, exposing anterior extremity of vomer. Palate 
short, noticeably concave both longitudinally and laterally ; its 
anterior emargination small, extending slightly behind level of 
middle of canine ; its posterior border slightly behind level of m 3 , 
double emarginate with short median projection ; mesopterygoid 
space slightly wider than long. Mandible slender, with no special 
peculiarities. 

Teeth. — In general aspect the teeth resemble those of Plecotus 
auritus, though they are throughout smaller and more slender. 
Incisors both above and below essentially as in Plecotus auritus 
except that crown of outer upper incisor is nearly terete, and 




266 



CHIROPTERA 



space between it and cingulum of canine is very narrow ; lower 
incisors forming a less broadly U-shaped row. Canines as in 
P. auritus. Anterior upper premolar very minute, crowded 
inward completely from tooth-row, the diameter of its crown 
barely one-quarter that of outer incisor, its cusp and cingulum 
distinct ; large upper premolar much as in P. auritus but less 
narrowed on inner side and with a narrow concave crushing 
surface ; no antero-internal basal cusp. Lower premolars almost 
exactly similar to middle and posterior lower premolar of Plecotus 
auritus, but crown area of larger tooth slightly exceeding that of 
canine. Molars not appreciably different from those of Plecotus 
auritus except that they are smaller and the concave median 
region of the upper teeth is larger relatively to area of crowns. 

Measurements. — Adult male from Cheshire, England : head 
and body, 48; tail, 49; tibia, 19; foot, 7-2; forearm, 38-4; 
thumb, 6 ; third finger, 69 ; fifth finger, 52 ; ear from meatus, 
15 ; width of ear, 14. Adult female from Epping, Essex, 
England : head and body, 49 • 6 ; tail, 46 ; tibia, 18' 2 ; foot, 7 ; 
forearm, 36; thumb, 5-8; third finger, 69 ; fifth finger, 51 ; ear 
from meatus, 16 ; width of ear, 14. Average and extremes of 
five adults from Damsdorf, Silesia, Germany : head and body, 
50 (47-52); tail, 47-1 (45-49); tibia, 18-5 (18-19); foot, 6-6 
(6-7); forearm, 38 (37-4-39); thumb, 5-8 (5-4-6-2); third 
finger, 70 (68-72) ; fifth finger, 52-6 (51-54) ; ear from meatus, 
15-5 (15-16); width of ear, 14-1 (13-6-15). For cranial 
measurements see Table opposite. 

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

England : Elton, Huntingdonshire, 1 ; Ellingham, Norfolk, 1 ; Arrow 
Lodge, Warwickshire, 1 ; Alcester, Warwickshire, 1 ; Rugby, Warwick- 
shire, 1; Welford, Rugby, 1 ; Pilton, Oundle, Northamptonshire, 1; Epping, 
Essex, 1 ; Llanelwedd, Radnorshire, 1 ; Swindon, Wiltshire, 1 ; Milton, 
Hampshire, 1. 

France : Caterille, Haute-Garonne, 1, 

Germany : Biichenberg, Harz Mountains, 1 ; Niesky, Silesia, 4 ; 
Bavaria, no exact locality, 3 (U.S.N.M.) ; near Wernigerode, Saxony, 1. 

Switzerland : Grotte de Vallorbe, Vaud, 40 (Mottaz). 

Remarks. — This species is immediately recognizable among 
European bats by its short, joined ears, and by the peculiar form 
of the muzzle. 

9. Llanelwedd, Radnorshire, H. E. Forrest (c & p). 4. 7. 3. 1. 

Wales. 

9. Elton, Huntingdonshire, Lord Lilford (p). 94. 9. 5. 1. 

England. 

6. Ellingham, Norfolk. E. R. Alston (p). 81. 6. 9. 1. 

(Dr. W. M. Crowfoot.) 

S. Arrow Lodge, Warwickshire. Tomes Collection. 7.1.1.348. 

1. Alcester, Warwickshire. Tomes Collection. 7. 1. 1. 348. 

(5 st. Rugby, Warwickshire. G. Dalgleish (c & p). 4. 10. 13. 2. 

S. Oundle, Northamptonshire. Lord Lilford (c & p). 94. 4. 16. 1. 

9 al. Epping, Essex. H. Doubleday (o & p). 44. 5. 1. 1. 

<5. Milton, Hampshire. Rev. J". E. Eelsall (p). 11. 1. 3. 59. 



BAKBASTELLA 



267 



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268 



CHIKOPTERA 



S. Swindon, Wiltshire. Rev. E. A. Goddard 8. 5. 12. 1. 

(c & p). 
<5. Caterille, Haute-Garonne, 0. Thomas (p). 6. 4. 1. 7. 

1000 m. France. 

(A. Robert.) 
$, ?. Niesky, Silesia, 181 m. Ger- Dr. E. Hamilton (p). 9Y. 12. 4. 2-3. 

many. (W. Baer.) 
2 ?. Niesky, Silesia. (W. Baer.) Lord Lilford (p). 99. 1. 9. 2, 4. 

S. Wernigerode, Saxony. Lord Lilford (p) . 0. 2. 8. 2. 

(Wolterstorff.) 

Sub-Family MINIOPTERINiE. 

1878. Miniopteri Dobson, Catal. Chiropt. Brit. Mus., p. 170 (part). 
1907. Miniopterinie Miller, Families and Genera of Bats, p. 227, June 29, 
1907. 

Geographical distribution. — Africa, southern Europe and 
southern Asia, eastward to the Malay region, Japan and 
Australia. 

Characters. — Like the Vespertilioninse, but presternum with 
median lobe enormously developed and forming the greater part 
of the bone ; scapula with coracoid straight, directed con- 
spicuously inward. 

BemarJes. — The sub-family Miniopterinx, though widely dis- 
tributed in the warmer portions of the Old World, is at present 
known to contain the genus Miniopterus only. 

Genus MINIOPTERUS Bonaparte. 

1837. Miniopterus Bonaparte, Ioonogr. Faun. Ital., i, faso. xx, under 

Vespertilio emarginatus (Sub-genus of Vespertilio). 
1857. Miniopterus Blasius, Saugethiere Deutschlands, p. 45. 
1866. Miniopteris Gray, Ann. and Mag. Nat. Hist., 3rd ser., xm, p. 91, 

February, 1866. 
1878. Miniopterus Dobson, Gatal. Chiropt. Brit. Mus., p. 347. 
1892. Minyopterus Winge, Jordf undne og nulevende Flagermus (Chiroptera) 

fra Lagoa Santa, Minas Geraes, Brasilien, p. 36. 
1900. Minneopterus Lampe, Jahrb. Nassau Ver. Naturkunde, Jahrg. 53, 

Catal. Saugeth.-Samml., p. 12. 
1907. Miniopterus Miller, Families and Genera of Bats, p. 227, June 29, 

1907. 

Type species. — Vespertilio ursinii Bonaparte = V. schreibersii 
Kuhl. 

Geographical distribution. — Same as that of the sub-family. 
Characters. — Cental formula : i H, c \^, vm |4 m t? = 34. 

.i-3 1—1 ■*■ 3— .J 3—3 

Skull with unusually high brain-case, and low, flattened rostrum. 
Second phalanx of third finger nearly three times as long as first. 
Ears short, separate, the upper margin (in European species) 
appearing almost artificially truncate. 

Remarks. — The peculiar shortening of the ears together with 
the remarkable elongation of second phalanx of third finger readily 



MINIOPTERUS 



269 



distinguish this genus among the members of the European fauna. 
About a dozen forms have been described, one of which occurs 
in southern Europe. 

MINIOPTERUS SCHREIBERSII Kuhl. 

1819. Vespertilio schreibersii Kuhl, Ann. Wetterau. Gesellsoh. Naturk., iv 
(= Neue Ann., i), pt. 2, p. 185 (Hungary). 

1837. Vespertilio ursinii Bonaparte, Ioonogr. Faun. Ital., I, fasc. xxi 
(Monte Corno, Ascoli, Italy). Type in British Museum. 

1841. Vespertilio orsinii Tern mi nek, Monogr. de Mamm., n, p. 179 (modi- 
fication of ursinii). 

1857. Miniopterus schreibersii Blasius, Saugethiere Deutschlands, p. 46. 

1878. Miniopterus schreibersii Dobson, Catal. Chiropt. Brit. Mus., p. 348. 

1910. Miniopterus schreibersi Trouessart, Faune Mamm. d'Europe, p. 34. 

Type locality. — Kulmbazer Cave, mountains of southern 
Bannat, Hungary. 

Geographical distribution. — Southern Europe from the Iberian 
Peninsula eastward, north to Switzerland and Hungary. Limits 
of range not known. 

Diagnosis. — Charaters as in the genus ; forearm about 
43 mm. 

External characters. — General form rather slender, with long 
tail and legs, wing broad at base but conspicuously tapering at 
tip, and short ears with a peculiar truncate aspect. Muzzle 
rather broad, though without conspicuous glandular swellings, 
its greatest width about equal to distance from eye to nostril ; 
muzzle pad narrow, with slight median emargination, bounded 
below by a low horizontal ridge which is continuous with projecting 
inner margin of nostril. Eyelids noticeably glandular-swollen ; 
a deep horizontal groove in cheek below eye. Ear short, extend- 
ing about half way from eye to nostril when laid forward, its 
general aspect different from that of any other European bat, 
owing to the length of the anterior basal lobe, the short, straight 
anterior border, and the broadly, evenly convex posterior border 
which joins anterior border in such a manner that there is 
practically no " tip," the whole anterior border appearing like an 
obliquely, almost artificially truncate extremity ; antitragus low, 
obscurely marked off from posterior border of conch, practically 
continuous with lower lip anteriorly ; inner surface of conch 
slightly rugose, without evident cross ridges ; tragus about half 
as high as conch, a little curved forward owing to slight con- 
cavity of anterior border, the blunt tip and upper half of exterior 
border forming a uniform, rather noticeable convexity, the 
posterior margin straight below to rudimentary basal lobe ; 
greatest width of tragus about half anterior border. Wing rather 
wide basally, the fifth finger exceeding forearm by about one-fifth 
length of latter, the tip unusually slender and elongate owing to 
the great length of last bone of third finger ; third and fourth 



270 CHIEOPTEHA 

metacarpals sub-equal, about 3 mm. shorter than forearm ; fifth 
metacarpal about 4 mm. shorter than third ; membrane joining 
leg at or a little above ankle. Leg rather slender, the foot 
scarcely half as long as tibia ; calcar about as long as tibia and 
nearly equal to free border of uropatagium, its distal termination 
very obscure, its posterior border without trace of keel. Tail 
about as long as head and body, included in the membrane to 
extreme tip. 

Fur and colour. — The fur is of a very soft, silky texture, 
though rather short, the hairs at middle of back only about 
7 mm. in length, those of head abruptly much shorter (about 
4 mm.) in rather noticeable contrast. Above it scarcely extends 
on membranes except for a sparse pubescence on basal half of 
median portion of uropatagium ; below it reaches line joining 
middle of forearm with knee and continues across extreme base 
of uropatagium. General colour of upper parts drab, faintly 
lighter and more nearly hair-brown anteriorly ; underparts ecru- 
drab. The hairs of back show three evident though not strongly 
contrasted colour-bands : (a) at base (3 mm.), a slaty mouse-grey ; 
(&) at middle (tips of shorter hairs, 2 mm.), ecru-drab ; and (c) at 
tip (extremities of longer hairs, 2 mm.), drab. Muzzle, ears and 
membranes brown, scarcely darker than body. 

Skull. — The skull differs from that of all other European bats 
in the great inflation of the anterior portion of brain-case and 
the consequent very abrupt angle at which the forehead rises 
above the low, flat rostrum. In other respects the general aspect 
of skull, especially when viewed from above, is rather slender . 
and lightly built. Dorsal profile rising gradually from front 
of nares to inter-lachrymal region, then 
abruptly at an angle of nearly 45° to middle 
of anterior portion of brain-case which is 
nearly or quite as high as lambda, the region 
■Qf^iy between these two highest points occupied 

by a shallow concavity corresponding to 
anterior edge of interparietal ; below and 
behind lambda the supraoccipital bulges 
noticeably outward, but not sufficiently to 
conceal condyles when skull is viewed from 
directly above ; ventral profile nearly flat, 
fig. 47. slightly elevated posteriorly. Brain-case 

Miniopterus schrewersii. -, jS j_j_i.lt i . __ 1 

Nat. size. broadly ovate, the outline broken at each 

side posteriorly by the slightly projecting 
mastoid region ; sagittal crest low but evident anteriorly, 
obsolete posteriorly ; lambdoid crest moderately developed at 
sides, barely indicated at middle ; general outline of skull when 
viewed from behind squarish, the depth through bulla about 
equal to breadth of brain-case ; floor of brain-case with no special 
peculiarities ; between cochlea and median line a shallow groove 
with abrupt pit-like anterior termination a noticeable ridge 





MINIOPTERUS 



271 



at outer side of each pit ; auditory bulla moderately large, the • 
transverse diameter about equal to distance between bullae. 
Interorbital region moderately constricted, slightly hour-glass 
shaped, the lachrymal region scarcely more than half as wide 
as brain-case, smoothly rounded at sides ; rostrum tapering 
gradually, rounded off at sides, flattened concave along median 
line, the narial emargination small, squarish, extending backward 
about one-third of the distance to front of forehead ; rostral 
depth at front of orbit about equal to distance from orbit to 
outer incisor ; anteorbital foramen rather large, directly above 
small premolar, the anteorbital canal half as long as rostrum, 
thus much longer than in any other European bat ; lachrymal 
foramen just outside of orbit, on level with upper border of 
anteorbital foramen. Palate long and wide, distinctly concave 
both laterally and longitudinally, terminating rather abruptly a 
little behind level of last molar, a small but evident foramen at 
each side near posterior edge ; median spine large ; mesoptery- 
goid space squarish, slightly wider posteriorly than anteriorly, 
the short hamulars bent inward. Mandible slender, a little 
deeper at symphysis than behind tooth-row, the posterior seg- 
ment unusually small, with nearly horizontal, slightly concave 
upper border ; angular process relatively long, expanded at 
outer end. 

Teeth. — Relatively to size of skull the teeth are small and weak. 
Inner upper incisor low, the crown very oblique, with postero- 
external concavity and small posterointernal cusp ; outer upper 
incisor considerably larger than inner, the crown flattened in 
axis of tooth-row, the width of its flattened-concave posterior 
surface about half its height, that of its outer border scarcely 
one-fifth height, cingulum obsolete, but forming a minute postero- 
external cusp ; the two teeth lie in curve of anterior portion of 
tooth-row, and the outer is separated from canine by a space 
about equal to its greatest diameter. Lower incisors closely 
crowded but not imbricated, the crown-area increasing regularly 
from first to third, the outline of the row as a whole broadly 
V-shaped ; i t and i 2 with crown very low, the cutting edge 
obscurely tried, the crown of each tooth wider posteriorly (in line 
of booth-row) than anteriorly, but this more evident in second 
than in first ; j 3 nearly terete, the middle and posterior cusps 
enlarged and separated by a deep groove, the anterior cusp 
reduced to a mere rudiment. Upper canine slender and weak, 
sub-terete, but distinctly flattened on inner side, the shaft with 
distinct anterior and posterior longitudinal grooves but without 
well developed cutting edge, the cingulum narrow, complete, 
but without cusps ; lower canine scarcely higher than main 
cusps of molars, its inner and posterior surfaces flattened, its 
antero-external surface smoothly rounded, cingulum forming a 
low but evident antero-internal cusp. Anterior upper premolar 
with crown area about equal to that of canine, its general form 



272 



CHIKOPTEEA 



essentially as in large premolar except that the antero-internal 
basal cusp is barely indicated, the cutting edges of the main cusp 
are less developed, and the crown is narrower externally than 
internally ; large upper premolar with crown area about equal 
to that of first molar and double that of small premolar, its 
anterior and posterior borders very slightly concave, the antero- 
internal basal cusp small but evident, the main cusp with well 
developed anterior and posterior cutting edges, and antero-internal 
and postero-external longitudinal groove ; lower premolars 
essentially alike in form, the crown area of the posterior tooth 
about equal to that of canine, that of the two others successively a 
little less, the outline squarish, with antero-external angle tending 
to become rounded off, particularly in the posterior tooth ; cusp of 
first about half as high as canine, that of second slightly higher 
than first, that of third nearly as high as main cusps of molars ; 
cingulum well developed, forming in each tooth a small but 
evident antero-internal basal cusp. First and second upper 
molars with posterior border deeply and almost angularly 
emarginate near inner edge of tooth, the anterior border nearly 
straight ; protocone low but rather broad ; hypocone clearly 
indicated but not distinct from posterior commissure of protocone ; 
paracone and metacone nearer outer edge of crown than usual, 
leaving a wide, pit-like median concavity, metacone slightly the 
higher of the two cusps ; styles and commissures well developed, 
Wpattern normal ; third upper molar with crown area about 
half that of first, its width through metacone slightly less than 
half length of anterior border, its three cusps and three 
commissures normally developed. Lower molars with no special 
peculiarities, the second triangle of m 3 narrower than first, but 
of about the same area. 

Measurements. — Adult male and female from Neuchatel, 
Switzerland : head and body, 59 and 59 ; tail, 58 and 60 
tibia, 20 and 19; foot, 10 and 9 '4; forearm, 45 and 44 • 6 
thumb, 7 and 7 ; third finger, 92 and 92 ; fifth finger, 53 and 54 
ear from meatus, 12 '6 and 12 ; width of ear, 12 and 12. Adult 
male and female from Corinth, Greece : head and body, 57 and 
56; tail, 57 and 55; tibia, 18 ■ 6 'and 19; foot, 10 and 9 6; 
forearm, 44 and 45 ; thumb, 6 ■ 6 and 7 • 2 ; third finger, 92 and 
90 ; fifth finger, 54 and 53 ; ear from meatus, 12 and 12 ; width 
of ear, 12 and 12. Two adult males from Silos, Burgos, Spain 
head and body, 60 and 60 ; tail, 60 and 57 ; tibia, 19 and 19 -4 
foot, 11 and 9- 6 ; forearm, 44 and 44 ; thumb, 7 '4 and 7 -4 
third finger, 93 and 89 ; fifth finger, 55 and 56 ; ear from meatus, 
11-4 and 11-6; width of ear, 13-4 and 12-6. Two adult 
females from the same locality : head and body, 59 and 59 ; tail, 
60 and 59 ; tibia, 19 ■ 6 and 20; foot, 9'8 and 10; forearm, 
45-4 and 46 ; thumb, 8 and 7-4 ; third finger, 93 and 93; fifth 
finger, 55 and 55 ; ear from meatus, 12 and 11 • 6 ; width of ear, 
12 '6 and 12 • 6. For cranial measurements see Table, p. 274. 



MINIOPTBRUS 



273 



Specimens examined. — Three hundred and fifty-one, from the following 
localities : — 

Spain: Silos, Burgos, 9 (B.M. and U.S.N.M.); Seville, 1; Minorca, 
Balearic Islands, 3 ; Majorca, Balearic Islands, 2. 

France : Troubate, Hautes-Pyrenees, 2 ; Dions, Gard, 9 ; Marseilles, 1 
(U.S.N.M.). 

Switzerland: Geneva, 235 (Mottaz) ; Neuchatel, 2 (U.S.N.M.). 

Austria-Hungary : Ofener Mountains, Hungary, 1 ; Hungary, no oxact 
locality, 3. (U.S.N.M.). 

Montenegro : Beri, 1 ; Velgi, Czolo, 1. 

Italy : Western Liguria, 14 (Genoa) ; Finalborgo, Liguria, 5 (Genoa) ; 
Maremma, Tuscany, 1 (U.S.N. M.) ; Pisa, 1; Florence, 1 (U.S.N.M.) ; 
Livorno, 6 (U.S.N.M.); Spezia, 5 (U.S.N.M.); Monte Gorno, Ascoli, 1 
(type of ursinii Bonaparte) ; Borne, 10 ; Velletri, Bome, 2 (U.S.N.M.) ; no 
exact locality, 1 ; Elba Island, 5 (U.S.N.M.) ; Marsala, Sicily, 2. 

Sardinia: Sassari, 3 (U.S.N.M.); Cagliari, 4 (U.S.N.M.); Mount 
Gennargentu, 2 (U.S.N.M.). 

Greece : Corinth, 17 (U.S.N.M.) ; Labyrinth, Crete, 1. 

Remarhs. — The lengthened second phalanx of the third finger, 
the peculiar cropped appearance of the ears, and the short, dense, 
velvety fur of the head are the most obvious external character- 
istics of Miniopterus schreibersii as compared with other European 
bats. 

2. Silos, Burgos, Spain. Bev. S. Gonzalez (c). 88. 1. 5. 46-47. 
<J, ?. Inca, Majorca, Balearic 0. Thomas and B. I. 0.7.1.4-5. 

Islands. Pocock (c & p). 

3 <5. San Cristobal, Minorca. 0. Thomas and B. I. 0. 7. 1. 32-34. 

Pocock (c & p). 
<$, 9. Troubate, Hautes- 0. Thomas (p). 6. i. 1. 8-9. 

Pyrenees, 400 m. 
Prance. (A.Robert.) 
8 <5, 1 ? Dions, Gard, 70 m. 0. Thomas (p). 8. 8. 10. 1-9. 

Prance. (0. Mottaz.) 

1 al. Ofener Mountains, Budapest Museum (e). 94. 7. 18. 10. 

Budapest. 
9. Beri, Montenegro. 0. Thomas (p). 5. 8. 4. 6. 

(L. FUhrer.) 
9. Velgi, Czolo. 0. Thomas (p). 5. 8. 4. 5. 

(L. FiXhrer.) 
7 <5, 3 9. Rome. (C. Coli.) Lord Lilford (p). 11. 1. 2. 5-14. 

Skeleton Monte Corno, Ascoli, Tomes Collection. 7. 1. 1. 735. 

(without Italy. (Type of Vespertilio ursinii Bonaparte.) 

skull). 

2 £. Marsala, Sicily. 0. Thomas (p). 6. 8. 4. 24-25. 

(A. Robert.) 
S. Labyrinth, 200 ft. Miss D. Bate (c). 5. 12. 2. 9. 

Crete. 



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276 CHIEOPTBEA 



Family MOLOSSIT)^. 

1865. Molossi Peters, Monatsber. k. preuss. Akad. Wissensch., Berlin, p. 258. 
1872. Molossidse Gill, Arrangement of the Families of Mammals, p. 17. 
1878. Emballonuridas (part; Molossinx, part, Molossi) Dobson, Catal. 

Cbiropt. Brit. Mus., p. 402. 
1907. Molossidse Miller, Families and Genera of Bats, p. 241, June 29, 1907. 

Geographical distribution. — Warmer parts of both hemispheres ; 
in the Old World north to the Mediterranean region and 
southern Asia. 

Characters. — Essentially like the Vespertilionidae except that 
the secondary articulation of humerus with scapula is more 
perfectly developed and the fibula is robust, adding appreciably 
to strength of leg. Auditory bulla noticeably emarginate on inner 
side. Tail projecting very conspicuously beyond membrane, a 
character by which the only European member of the group may 
be immediately recognized. 

Bemarhs. — The family Molossidse is widely distributed in the 
warmer parts of both hemispheres. Eleven genera are now 
known, one of which is represented in the Mediterranean region 
of Europe. 

Genus NYCTINOMUS Geoffroy. 

1818. Nyctinomus Geoffroy, Descr. de l'Bgypte, n, p. 114 (segyptiacus). 
1821. Nyctinoma Bowdioh, Anal. Nat. Class. Mamni., p. 28 (Modification 
of Nyctinomus). 

1821. Nyctinomes Gray, London Med. Bepos., xv, p. 299, April 1, 1821 

(Modification of Nyctinomus). 

1822. Nyctinomia Fleming, Philos. of Zool., n, p. 178 (Modification of 

Nyctinomus). 
1842. Mops Lesson, Nouv. Tabl. B.egne Anim., p. 18 (Mops indicus Lesson 

= Dysopes mops F. Cuvier). 
1878. Nyctinomus Dobson, Catal. Chiropt. Brit. Mus., p. 420 (part). 
1902. Nyctinomops Miller, Proc. Acad. Nat. Sci. Philadelphia, p. 393, 

September 12, 1902 (femorosaccus). 
1907. Nyctinomus Miller, Families and Genera of Bats, p. 251, June 29, 1907. 

Type species. — Nyctinomus segyptiacus Geoffroy. 

Geographical distribution. — Warmer portions of both hemi- 
spheres, north to the southern United States and to the Mediter- 
ranean coast of Europe, east to the Philippines and Norfolk 
Island. 

Characters. — Dental formula : i tl or t?,* c t? , vm — , m s - = 30 

2-iJ 3-3' 1-1' £ 2-a' 3-3 

or 32 ; brain-case not unusually flattened, its depth at least half 
its width ; bony palate with a small median anterior emargination 
extending to behind level of roots of incisors. 

Remarks. — As thus defined the genus Nyctinomus contains 

* In the species occurring in Europe. 



NYCTINOMUS 277 

about forty species, two-thirds of which are peculiar to the Old 
World, one of them occurring in the Mediterranean region of 
Europe. 

NYCTINOMUS TENIOTIS Rafinesque. 

1814. Cephalotes teniotis Rafinesque, Pr6c. des D6couv. Somiol.,p.l2 (Sicily). 
1825. Dinops cestoni Savi, N. Giorn. de' Letterati, Pisa, x, p. 235 (Pisa, Italy). 
1840. Dysopes savii Sohinz, Europ. Fauna, i, p. 5 (Substitute for cestoni). 
1871. [Dysopes cestonii] var. nigrogriseus Schneider, Neue Denkschr. 

Schweiz. Gesellsch. Naturwiss., xxiv, p. 5 (articles separately 

paged). Basel, Switzerland. 
1877. Nyctinomus cestonii Dobson, Catal. Cbiropt. Brit. Mus., p. 423. 
1891. Nyctinomus tseniotis Thomas, Proc. Zool. Soc, London, p. 182. 
1897. [Dysopes] midas Schulze, Abh. Ges. Nat. iv, No. 10, p. 23 (Substitute 

for cestoni). Not of Sundevall, 1842. 
1910. Nyctinomus tseniotis Trouessart, Faune Mamm. d'Europe, p. 36. 

Type locality. — Sicily. 

Geographical distribution. — Mediterranean region of Europe 
and northern Africa. Accidental ? at Basel, Switzerland. 

Diagnosis. — Like the African Nyctinomus eegyptiacus but 
larger, condylobasal length of skull about 23 mm. instead of 
about 20 mm. ; lower incisor 3-3 instead of 2-2 ; small upper 
premolar with crown area more than one-half instead of less 
than one-fifth that of upper incisor. Distinguishable among 
European bats by the generic characters ; forearm about 60 mm. 

External characters. — Form heavy and robust, the legs short, the 
feet large, the wings long and narrow, the membranes thick and 
leathery, the ears very large, sub-orbicular, joined at their anterior 
bases ; these characters in connection with the thick muscular tail, 
projecting by at least one-third of its length beyond interfemoral 
membrane, immediately distinguish the animal from all other 
European bats. Muzzle projecting rather noticeably beyond 
upper lip, obliquely truncate ; nostril pad well denned, wider 
than high, the sub-circular nostrils opening forward and slightly 
outward at its outer margin, its surface with very fine reticula- 
tions and a few coarse wrinkles, glabrous except below, where it 
is sprinkled with fine hairs, this hairy area continuous with the 
brush-like fringe of recurved blunt hairs along middle of upper 
lip ; upper margin of pad broadly concave at middle, convex 
laterally over the nostrils, its edge thickly set with small, laterally 
compressed horny excrescences, about thirty-five in number, 
the outermost lying a little above level of middle of nostril ; a 
row of about ten similar excrescences crosses middle of pad 
vertically. Upper lip very large and full, marked by many deep 
oblique wrinkles, its surface both above and below rather densely 
hairy. A small wart on chin just behind level of symphysis. 
Ear very large, sub-orbicular, the margin with no very decided 
irregularities, though the anterior border usually shows some 
slight flattening and the posterior border is faintly concave above 



278 



CHIROPTERA 



and near base ; anterior basal margin of ear with a narrow 
outward-folded hem, the exposed side of which is densely 
pubescent except at extreme anterior margin, where it is glabrous 
and set with about six terete wart-like projections, widely and 
irregularly spaced ; posterior basal margin with a shorter, deeper, 
inwardly-folded lobe just above concavity limiting posterior 
base of antitragus ; keel well developed, not thickened at edge, 
its height posteriorly equal to about one-third its length, the 
margin of keel hairy, the line thus begun curving upward and 
backward under upper margin of conch ; antitragus well defined, 
its height slightly greater than that of keel, its length about 
twice height, its anterior border with ridge-like continuation 
forward to corner of mouth ; tragus squarish in outline, its 
posterior border longest and with distinct angle below middle, its 
anterior border shortest, its upper border almost horizontally 
truncate ; anterior and upper margin of tragus fringed with long 
loose hairs. Wings longer and narrower than in any other 
European bat, the membrane inserted on side of tibia just above 
ankle ; antebrachial membrane extending as a narrow fold along 
forearm to base of thumb ; the fur of body extends on both 
surfaces of wing to line joining middle of humerus with knee, 
and on to extreme base of uropatagium ; otherwise the membranes 
are essentially naked except for some fine dense pubescence on 
upper side of propatagium. Thumb short and robust, with 
moderately developed pad at distal end of metacarpal. Foot 
broad and robust, more than half as long as tibia, the sole with 
a low and rounded but evident pad at middle, four small elongated 
pads in a row at bases of toes, and some smaller, less definite 
callosities in space between this row and the large median pad. 
Outer and inner toe thickened, their outer surfaces densely 
covered with short stiffened hairs with recurved points ; sprinkled 
among these shorter hairs and also at the ends of the other toes 
are a few much longer bristles. Calcar about as long as tibia, 
its point ill-defined ; no indication of keel or of terminal lobe. 
Tail about half as long as head and body, robust and muscular, 
the terminal third or half projecting beyond membrane. 

Fur and colour. — The fur is everywhere dense and velvety 
in texture, the hairs at middle of back about 7 mm. in length, 
those on throat longer and looser. Colour a uniform light drab, 
with faint darker shading in certain lights, the hairs pale ecru- 
drab at extreme base. Ears and membranes in dry specimens 
blackish. Fringes on feet and hairy lines on ears, drab like 
body. 

Skull. — The skull is large, but rather slender, about equal to 
that of Myotis myotis in length. In general form it is distinguish- 
able among those of the bats of Europe by the depressed brain- 
case, the high, somewhat tubular rostro-interorbital portion, and 
the conspicuously emarginate inner side of auditory bullae. Dorsal 
profile essentially straight from nares to lambda, though with a 



NYCTINOMDS 



279 



slight convexity over anterior two-thirds of brain-case, and a 
more abrupt though not very conspicuous swelling posteriorly, 
the two convexities separated by a narrow concavity ; general 
direction of dorsal profile more nearly horizontal than in any 
other European bat, since the depth of occiput through condyles 
is scarcely greater than, that of rostrum through anterior portion 
of first molar. Brain-case low and wide, its depth at middle 
about half mastoid breadth, its surface smooth and evenly 
rounded, with faintly indicated sagittal crest posteriorly and 
median groove between lateral swellings anteriorly ; lambdal 
crest evident though not high ; outline of brain-case when 
viewed from above a somewhat triangular ovate owing to the 
rather squarely truncate posterior border and the strong contrast 
between the wide mastoid region and narrow interorbital con- 
striction ; floor of brain-case with median ridge and lateral 
depressions posteriorly, the basisphenoid 
with two shallow but rather well-defined 
pits about as large as glenoid surface ; 
auditory bullae rather large, deeply emar- 
ginate on inner side so that cochlea is 
conspicuously exposed, the region in 
front of middle of meatus reduced to a 
narrow ring barely more than one-third 
as wide as meatus, the extreme anterior 
border with a flange-like inward-curved 
projection. Interorbital region long, sub- 
cylindrical, a little wider at lachrymal 
level than posteriorly, but scarcely enough 
so to impart a distinctly hour-glass general 
form ; least interorbital breadth slightly 
greater than breadth of tubular narial 
region and slightly less than that across 
roots of canines. Zygoma simple, not 
bent upward, its margin with a barely indicated expansion behind 
middle. Rostrum proper short and deep, not well differentiated 
from interorbital region, the distance from orbit to front of pre- 
maxillary about equal to depth at front of m 1 ; lachrymal ridge 
short but well defined, the small, inconspicuous anteorbital fora- 
men and minute lachrymal foramen opening forward under its 
anterior edge, at level of middle of large premolar ; nares with 
distinctly tubular lateral margins separated from roots of canines 
by evident grooves, the dorsal emargination extending about half 
way to lachrymal level. Palate moderately wide, noticeably 
concave laterally, the anterior emargination small, about as deep 
as wide, its posterior border scarcely extending behind level 
of front of canine ; posterior border of palate double emargi- 
nate with well developed median projection, the emarginations 
extending forward to level of posterior margin of m 3 ; mesop- 
terygoid space large, its width anteriorly equal to that of 




Nyclinomua teniotis. 
Nat. size. 



280 



CHIEOPTERA 



temporal fossa at same level, its length about twice width ; 
hamulars small and inconspicuous. Mandible long and straight, 
its axis scarcely bent upward posteriorly ; symphysis deep, scarcely 
or not subtended by a concavity in lower border of ramus ; 
posterior portion of mandible low, the depth through coronoid 
process barely equal to distance from front of coronoid to back 
of condyle ; angular process large, straight, directed backward, 
outward and downward. 

Teeth. — Relatively to size of skull the teeth are rather large, 
the lower molars in particular. The most obvious peculiarities 
of the dentition as compared with that of other European bats are 
the single, well developed incisor in each side of upper jaw, and 
the presence of a large hypocone, quite distinct from commissure 
of protocone, in m 1 and m 2 . Upper incisor simple, a little more 

than half as high as canine, the 
shaft set at an angle so that the 
points of the two teeth are much 
nearer together than their bases ; 
cingulum faintly indicated in front, 
better developed and forming an 
incipient cusp postero-extemally ; 
posterior surface of shaft somewhat 
flattened ; space between incisor 
and canine about equal to greatest 
diameter of smaller tooth. Lower 
incisors three on each side, much 
imbricated and closely crowded in 
narrow space at front of canines, 
their cutting edge barely rising to 
level of canine cingulum. The 
inner and middle tooth sub-equal, 
rather deeply bifid, the outer 
barely equal to outer lobe of the 
others, its apex faintly notched. 
Upper canine moderately large, its crown area somewhat more 
than double that of upper incisor, the general outline of its 
base an irregular triangle with its longest side directed 
inward, and with the postero-external side slightly concave; 
cingulum narrow but well developed throughout, not tending to 
develop small cusps ; lower canine with no special peculiarities, 
its cingulum tending to form a slight anterior cusp. Small 
upper premolar in the tooth-row, usually in contact with canine 
and separated from large premolar by a narrow space, its crown 
area somewhat less than that of upper incisor, its cingulum and 
cusp well developed, the cusp about one-fourth as high as shaft 
of canine, triangular in outline when viewed from the outside, 
with well developed posterior cutting ridge ; lower premolars 
similar in form but larger, their crown areas sub-equal, the shaft of 
the second higher than that of first and about equal to larger cusps 





Fig. 49. 
Xyctinomiis leniotis. Teeth X 5. 



NYCTINOMtTS 



281 



l 

8 

I 

B 

tei 

O 

CO 
EH 

% 

cc 
<3 

H 






•Aioi-qqoo^ 
JBinqipu^K 



■Moi-q^oo? 



■9[qipuuj5 



■sippim 

%v ssro-nrejq 

jo q?dfe(r 



■9ffB0-UTBjq 

JO q^pBajg 



'qlpuoiq 
IBOl^iqoBI 



'uoi^oij^snoo 
im)!qjOJ35ni 



•qjpmiq 



•q^Susi 
lusuqopfpuoo 






o 

o 



o 

02 



o 



is 



S la 



CO 
C3 



o cq 

Oi OS 



o cq 

CO CO 



O 



a 
5 



C3 
HO 
O 



Pm 



282 



CHIROPTEEA 



of lower molars ; large upper premolar with crown area about 
two-thirds that of m 1 , its antero-internal cusp high and distinct. 
First and second upper molars alike in structure, the first slightly 
larger than second, the cusps all well developed and distinct, 
showing no special peculiarities aside from the presence of a 
conspicuous terete hypocone nearly as large and distinct as antero- 
internal cusp of large premolar, and separated from commissure 
of protocone by a deep notch ; third upper molar with crown 
area not much less than that of second, except for the absence of 
postero-internal heel and hypocone ; third commissure as long as 
in the other teeth, but metacone smaller than paracone. Lower 
molars with no special peculiarities, the cusps all well developed 
and distinct. 

Measurements. — Young adult male from Italy : head and 
body, 87 ; tail, 56 ; free portion of tail, 20 ; tibia, 19 ; foot, 
10-4; forearm, 60 - 4; third finger, 115; fifth finger, 65; ear 
from meatus, 30 • 4 ; width of ear, 32. Young adult female from 
Greece : head and body, 84 ; tail, 57 ; free portion of tail, 27 ; 
tibia, 20; foot, 114; forearm, 61; third finger, 117; fifth 
finger, 61 ; ear from meatus, 31 - 4 ; width of ear, 31. Two adult 
females from Cintra, Portugal : forearm, 60 and 61 ; third finger, 
110 and 115 ; fifth finger, 59 and 63. For cranial measurements 
see Table, p. 281. 

Specimens examined. — Six, from the following localities : — 
Portugal : Cintra, 2. 

Italy: No exact locality (probably from Pisa), 2. (B.M. and U.S.N.M.); 
Sicily, 1 (U.S.N.M.). 

Greece : No exact locality, 1. 

2 9. Cintra, Portugal. O. Thomas (c & p). 98. 2. 2. 6-7. 

S. Italy (probably Pisa). Zoological Society (p). 10. 5. 24. 1. 

{Savi.) 

?. Greece. Parreys. 46. 6. 15. 121. 



CARNIVORA 



283 



Order CARNIVORA. 

1827. Carnivora Gray, Griffith's Cuvier. Anim. Kingd., v, p. 111. 

Geographical distribution. — Continents and larger islands of 
the entire world, Australia,* New Zealand, and the Antarctic 
region excepted. 

Characters. — Terrestrial (rarely aquatic or semi-aquatic), non- 
volant, placental mammals with rather high development of 
brain, the cerebral hemispheres with distinct convolutions ; feet 
unguiculate, never modified as fins or flippers ; dentition of a 
modified tuberculo-sectorial type, the posterior upper premolar 
and anterior lower molar usually developed as special carnassial 
or flesh-cutting teeth. 

Bemarks. — The mammals of this order present much diversity 
of form and structure, though less than in the case of the 
Insectivora. Most of the living members of the group are 
carnivorous in habits, and immediately recognizable among 
placental mammals by the presence of a specially modified flesh- 
tooth in each jaw. In certain groups, however, as in the Ursidse 
among the European representatives of the order, both habits 
and dentition are of a more generalized type. The order contains 
seven recent families, five of which occur in Europe. 

KEY TO THE EUROPEAN FAMILIES AND SUB-FAMILIES 
OF CABNIVOBA. 

Larger cheek-teeth with crowns of a crushing type, the 
cusps sub-equal, low, subterete, without noticeable 
cutting edges ; upper carnassial 2-rooted, in front of 
anteorbital foramen, its inner lobe posterior; size 
very large; form heavy; feet plantigrade (Bears) .. . Ursidas, p. 284. 
Larger cheek-teeth with crown at least partly trenchant, 
the outer cusps of one or more in each jaw narrow 
and with well developed cutting edge, the inner 
cusps reduced or absent ; upper carnassial 3-rooted, 
behind anteorbital foramen, its inner lobe median 
or anterior. 
Cheek-teeth without crushing surfaces ; upper molar 
scarcely larger than outer incisor ; claws com- 
pletely retractile (Cats) Felidie, p. 455. 

Cheek-teeth, at least the hindermost, with evident 

crushing surface; upper molar (or first when more 

than one are present) much larger than outer 

incisor ; claws partly or not retractile. 

Tooth-row relatively long (more than half condylo- 

basal length of skull) ; number of teeth in 

European members of family 42 (Dogs) Canidie, p. 303. 



Represented in Australia by a species of Canis, probably introduced. 



284 



CARNIVOEA 

Tooth-row relatively short (less than half condylo- 

basal length of skull) ; number of teeth in 

European members of family not more than 40. 

Auditory bulla divided into two chambers, the 

boundary between which is marked externally 

by an oblique constriction; upper molars 

usually (always in European genera) 2-2, 

the crown of the first wider externally than 

internally (Genets and Mongoose) Viverridm, p. 440. 

Auditory bulla simple ; upper molars 1-1, the 

crown wider internally than externally MusteUdas, p. 340. 

Upper carnassial with evident crushing surface 

on inner side, the crown triangular or 

rhombic in outline ; upper molar large, the 

length of its outer portion usually equal to 

or greater than that of carnassial. 

Skull normal, the rostrum longer than broad ; 

external form not modified for aquatic 

life, the toes long-clawed, not webbed, 

the tail not conspicuously muscular ; fur 

loose and coarse (Badgers) Melinse, p. 341. 

Skull much flattened ; rostrum broader than 
long ; external form modified for aquatic 
life, the toes short-clawed, webbed, the 
tail conspicuously muscular; fur dense 

and fine (Otters) Lutrime, p. 354. 

Upper carnassial without crushing surface on 

inner side other than a small concave area 

between small inner lobe and main cusp, 

the crown not triangular or rhombic in 

outline; upper molar much reduced, the 

length of its outer portion one-third to 

one-half that of carnassial. 

Dentition highly trenchant ; small premolars 

not opposite, at least one pair capable of 

shearing action ; upper carnassial with 

posterior cusp narrow and trenchant ; 

auditory bulla longer than broad; form 

slender; feet digitigrade (Martens and 

Weasels) Mustdinx, p. 364. 

Dontition not highly trenchant ; small pre- 
molars opposite, not capable of shearing 
action, the points of all but pm? and pm i 
widely separated when jaws are closed ; 
upper carnassial with posterior cusp 
broad, almost flat-topped ; auditory bulla 
broader than long ; external form heavy ; 
feet sub-plantigrade (Glutton) Gulminte, p. 432. 



Family TJRSIDiE. 

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

Geographical distribution. — Northern hemisphere, south in the 
Old World to the Atlas Mountains and the Malay Archipelago, 
and in America to the Andes. 

Characters. — Larger cheek-teeth of a strictly crushing type, 
the crowns wide and flattened, with large terete cusps, the last 



TJKSDS -285 

upper premolar and first lower molar scarcely differentiated as 
carnassials, the former 2-rooted, its inner lobe at posterior border 
of crown, its position so far anterior to level of antborbital 
foramen as not to be at point of greatest mechanical efficiency ; 
auditory bulla flattened, without septum ; form heavy ; size 
large ; feet strictly plantigrade ; digits, 5-5. 

Remarks. — The family Ursidse, containing the bears, is at 
present represented by five or six genera, though the fossil 
remains of others are known. The members of the group are so 
easily recognizable by the peculiarities of the cheek-teeth that 
they require no special comparisons with other carnivora. Two 
genera occur in Europe. 

KEY TO THE EUROPEAN GENERA OP URSIDM. 

Cheek-teeth relatively large ; incisors and canines not 

specially enlarged and prehensive (Ordinary Bears) Ursus, p. 285. 

Cheek-teeth relatively small ; incisors and canines 
enlarged and unusually^ prehensive in character 
(Polar Bears) .' Thalarctos, p. 297. 

Genus URSUS Linnaeus. 

1758. Ursus Linnaeus, Syst. Nat., I, 10th ed., p. 47 (arctos, by tautonymy). 
1857. Ursus Blasius, Saugethiere Deutschlands, p. 196. 
1864. Euarctos Gray, Proc. Zool. Soc. London, p. 692 (americanus). 
1864. Myrmarctos Gray, Proc. Zool. Soc. London, p. 694 (eversmanni = arctos) . 
1898. Ursarctos Heude, Mem. Hist. Nat. Emp. Chinois, iv, pt. i, p. 18 
sis). 



Type species. — Ursus arctos Linnseus. 

Geographical distribution. — Northern hemisphere from northern 
limits of the great continental areas south to the Atlas Moun- 
tains, the Himalayas and Mexico. 

Characters. — Dental formula : i *"*, c J"J, pm ", m £§ = 42 ; 
inner upper incisor well developed, permanent ; first, second and 
third premolars in both jaws small, single-rooted, readily deciduous, 
especially pm 2 , pm 2 and pm 3 ; molars large and robust, the length 
of the two upper teeth together equal to width of palate. 

Remarks. — The genus Ursus as thus restricted is a very 
homogeneous group practically confined to the north temperate 
region. The species are at present so imperfectly known that no 
fair estimate can be made of their number. Recently about 
thirty forms have been recognized, only one of which is definitely 
known to occur in Europe. 

URSUS ARCTOS Linnseus. 

1758. [Ursus] arctos Linnaeus, Syst. Nat.,*i, 10th ed., p. 47 (Sweden). 
1772. [Ursus] ursus Boddaert, Kortbegrip van het zamenstel der Natuur, 

i, p. 46 (Renaming of arctos). 
1788. [Ursus arctos] a niger Gmelin, Syst. Nat., I, 13th ed., p. 100 

(Northern Europe). 



286 



CABNIVORA 



1788. [Ursus arctos] $ fuscus Gmelin, Syst. Nat., i, 13th ed., p. 100 (Alps). 

1778. [Ursus aretes] y alhus Gmelin, Syst. Nat., i, 13th ed., p. 100 
(Unknown ; based on the "ours blanc terrestre " of Buflon). 

1792. U[rsus] arctos griseus Kerr, Anim. Kingd., p. 184 (Germany; also in 
northern North America). 

1797. Ursus arctos rufus Borkhausen, Deutsche Fauna, i, p. 46 (Swiss and 

Tirolean Alps). 

1798. Ursus badius Schrank, Fauna Boica, i, p. 55 (Forests on the 

Bohemian boundary). 
1808. Ursus fuscus Tiedemann, Zoologie, I, p. 374 (Substitute for arctos). 

1814. Ursus alpinus Fischer, Zoognosia, in, p. 161 (Alps? Based on an 

individual seen alive in Paris). 
1820. Ursus arctos major Nilsson, Skand. Fauna, I, p. 112 (Wooded portions 

of southern Scandinavia). 
1820. Ursus arctos minor Nilsson, Skand. Fauna, I, p. 123 (Northernmost 

Scandinavia). 
1827. [Ursus arctos] $ brunneus Billberg, Synopsis Faunae Scandinavise, 

p. 15 (Northern Scandinavia). 
1827. [Ursus arctos] j annulatus Billberg, Synopsis Faunae Scandinavise, 

p. 15 (Northern Scandinavia). 
1827. [Ursus arctos] 8 argenteus Billberg, Synopsis Faunae Seandinaviae, 

p. 15 (Northern Scandinavia). 

1827. [Ursus] myrmephagus Billberg, Synopsis Faunae Seandinaviae, p. 16 

(Northern Scandinavia). 

1828. [Ursus] formicarius Billberg, Synopsis Faunae Seandinaviae, 2nd ed., 

p. 16 (Renaming of myrmephagus). 

1829. U[rsus] pyrenaicus Fischer, Synopsis Mamm., p. 142. Latinization 

of "Ours des Pyrenees" F. Cuvier, Hist. Nat. des Mammif., v, 

fasc. 44, 1824 (Asturias, Spain). 
1829. U[rsus] norvegicus Fischer, Synopsis Mamm., p. 142. Latinization 

of "Ours de Norwege " F. Cuvier, Hist. Nat. des Mammif., n, 

fasc. 7, 1819 (Norway). 
1836. ? Ursus falciger Reichenbach, Regn. Anim. Icon., I, p. 32 (' ' Pyrenees" ; 

afterwards supposed to be an individual of " 17. ferox." See Natur- 

gesch. des In- und Auslands, Raubsaugeth., p. 299, 1852). 
1842. Ursus pyrenieus F. Cuvier, Hist. Nat. des Mamm., Tabl. gen., p. 3 

(Described in fasc. 44, 1824) (Asturias, Spain). 
1847. ? Ursus euryrhinus Nilsson, Skand. Fauna, i, 2nd ed., p. 212 (Sweden ? 

Type an individual raised in captivity). 
1857. Ursus arctos Blasius, Saugethiere Deutschlands, p. 196. 

1861. Ursus arctos aureus Fitzinger, Wissensch.-pop. Naturgesch. der 

Saugeth., I, p. 372 (Norway). 
1864. [Ursus arctos] var. 1. normalis Gray, Proc. Zool. Soc, London, p. 682 

(Renaming of arctos). 
1864. [Ursics arctos] sub-var. a. scandinavicus Gray, Proc. Zool, Soc, 

London, p. 682 (Based on Nilsson, Ilium. Fig. Skand. Fauna, 

pi. 23). 
1864. ? [Ursus arctos] sub-var. c. rossicus Gray, Proc. Zool. Soc, London, 

p. 682 (nomen nudum). 
1864. [Ursus arctos] sub-var. f. yplonicus Gray, Proc. Zool. Soc, London, 

p. 682 (Poland ; based on Cuvier, Oss. Fossiles, iv, p. 332, pi. xxn, 

fig. 3). 
1864. [Ursus arctos] var. 2. grandis Gray, Proc. Zool. Soc, London, p. 684 

("North of Europe " ; based on " a male purchased at Hull, living 

in the Zoological Gardens from 1852 to 1863 "). 



uesds 287 

1864. [Ursus arctos] var. 4. stenorostris Gray, Proc. Zool. Soc., London, 

p. 685 (Poland ; based on Cuvier, Oss. Fossiles, iv, p. 332, 2nd var., 

pi. xxu, fig. 4). 
1864. Myrmarctos eversmanni Gray, Proc. Zool. Soc, London, p. 695 

(Norway). 
1910. Ursus arctos, U. arctos formicarius, U. arctos alpinus, and 77. arctos 

pyrenaicus Trouessart, Faune Mamm. d'Europe, pp. 67-68. 

Type locality. — Sweden. 

Geographical distribution. — Entire continent of Europe wher- 
ever sufficiently extensive forests remain ; east into Asia ; west 
formerly to Great Britain, where it became extinct about the 
eleventh century ; not certainly known to have occurred in 
Ireland. 

Diagnosis. — Size moderate, condylobasal length of skull 
ranging from about 260 to 350 mm. ; interorbital region notice- 
ably elevated, the frontal profile strongly convex ; mesopterygoid 
region not specially shortened and broadened, the width between 
pterygoids decidedly less than half distance from hamular to 
level of last molar ; colour brown or butfy, varying much in exact 
shade, the legs usually darker than body, and feet darker than 
legs. 

External characters. — General form short and heavy, this 
made more apparent by the long rather loose fur. Head 
moderately pointed, rather broad posteriorly ; ear short, narrowly 
rounded off above, nearly concealed in the fur, its tip not extend- 
ing to eye when laid forward ; muzzle squarely truncate, its pad 
naked, the upper border somewhat projecting backward, its lower 
border separated from upper lip by a broad hairy area crossed 
at middle by a nearly bare perpendicular line. Fore foot with 
digits robust, inconspicuously graduated, the third and fourth 
sub-equal and longest, the fifth and second sub-equal and slightly 
shorter, the first with anterior edge of ball extending about to 
middle of that of second, this interval greater than in the case 
of the other digits ; claws strongly curved, blunt, without evident 
cutting edges, their length at least twice that of those on hind 
foot ; balls of digits large, pad-like, their surface, like that of 
pads, coarsely rugose ; main pad wider than long, covering more 
than half surface of palm, its outer border about twice as long as 
inner, its porterior border slightly concave, its inner portion, at 
base of thumb, marked off from rest of pad by a slight furrow ; 
region between main pad and balls of digits densely furred; 
wrist pad about as large as ball of digits, near outer margin of 
palm, its long diameter transverse ; region between wrist-pad and 
main pad densely furred ; hairs along edge of palm standing out 
stiffly, especially on outer side. Hind foot longer than fore foot, 
the second and third digits sub-equal and longest, the first and 
fourth slightly shorter, the fifth with anterior edge of ball at 
middle of that of fourth ; pad like that of fore foot, but with a 
broad backward extension passing along inner side nearly or 



288 



CAKNIVOEA 



quite to heel ; region between pad and balls of toes, and at outer 
side of backward extension densely furred ; fringe along edge of 
foot conspicuous. Tail very short, concealed in the fur. 

Colour. — The colour of body is usually a light brown or dull 
buff, the head not essentially different, but feet and outer surface 
of legs darker. Many individual differences in colour have been 
described, some of which are probably characteristic of geographical 
races. 

Skull. — General form of skull rather robust, the rostrum 




KU. 50. 
Ursus arctos. 



X S. 



moderately long (distance from orbit to front of premaxillary 
contained about 2£ times in condylobasal length), the brain-case 



uksus 289 

deep (depth to level of under side of postglenoid process con- 
siderably more than distance from tip of postorbital process to 
middle of interparietal) but not unusually wide (mastoid breadth 
slightly exceeding depth to under side of postglenoid process). 
Dorsal profile usually with an evident concavity in interorbital 
region, but this character showing much variation ; highest point 
at bregma or slightly further forward, the profile nearly straight 
and sloping away at an angle of about 20° behind this point to 
slightly overhanging lambdal region ; ventral profile faintly and 



evenly concave throughout. Brain-case broadly ovate in outline, its 
greatest breadth about equal to distance from bregma to lambda,* 
its depth at middle, exclusive of sagittal crest, slightly less than 
greatest breadth ; lambdoid and sagittal crests well developed in 
adults, the lambda noticeably projecting so that occiptal condyles 
are scarcely visible when skull is viewed from above, and region 
between crest and foramen magnum is deeply concave ; sagittal 
crest dividing in region of bregma into two ridges, one of which 

* Except in very old individuals in which the lambda is greatly 
produced backward. 



'290 



CABNIVORA 



runs to extremity of each postorbital process, the hinder margin 
of which it forms ; occiput when viewed from behind moderately 
broad, the depth from lambda to lower lip of foramen magnum 
contained If to If times in mastoid breadth ; mastoid processes, 
paroccipital processes, and condyles extending to about the same 




via. 62. 
XTrsus arctos. X J. 

level. Floor of brain-case nearly flat, the basioccipital with 
raised edges applied to inner surface of bullae, and in some 
specimens marked by an evident concavity on each side of median 
line ; auditory bulla flat, not rising above edge of basioccipital, the 
greatest longitudinal diameter less than transverse diameter, the 



ursus 291 

meatus distinctly tubular, usually longer than wide ; postglenoid 
process heavy, rising to level of hamulars. Interorbital region 
broad, the width across robust, triangular, postorbital processes 
about equal to that of brain-case, the region immediately between 
orbits always a little concave and sometimes conspicuously so, 
that at base of each postorbital process usually somewhat swollen. 
Zygomata moderately expanded, the greatest zygomatic breadth 
opposite anterior glenoid edge ; orbital process well developed. 
Rostrum equal to less than half condylobasal length of skull, the 
width across alveoli of canines equal to or less than depth at 
front of orbit, the depth at front of nasal equal to about half 
distance from orbit to front of premaxillary ; nares rather large, 
their lateral margins slightly everted ; nasal bones elongate wedge- 
shaped, squarely truncate anteriorly, their posterior extremity on 
level with or extending slightly behind nasal branch of maxillary ; 
anteorbital foramen over metacone of m 1 or paracone of m 2 ; 
palate narrow, its width between posterior molars contained about 
3£ times in median length ; extension behind molars nearly 
parallel-sided, its length equal to about three-quarters breadth ; 
mesopterygoid space 1| times to twice as long as wide, its anterior 
border squarish or rounded, its lateral borders nearly parallel ; 
hamulars small but distinct, slightly hooked outward. Man- 
dible robust, the depth of ramus behind large premolar con- 
tained about five times in length, the height of posterior portion 
(measured to level of lower border) a little less than half length j 
coronoid process broad, its width at level of alveolus slightly 
greater than height, the anterior border at first straight then 
evenly convex to overhanging tip, the posterior border concave ; 
angular process short, extending slightly if at all behind level of 
articular process, its inner border nearly straight, its outer border 
convex. 

Teeth. — The teeth are moderately large relatively to size of 
skull. Upper incisors forming a continuous row, separated at 
each side from canine by a diastema about as wide as inner 
incisor ; i 1 and i 2 sub-equal, the former slightly the smaller, the 
anterior face smoothly rounded, a little more than half as wide 
as high, the posterior face abruptly concave, with shelf-like 
posterior extension, the cingulum slightly developed and forming 
a rudimentary nodule on inner and outer side of i 2 and on outer 
side of j 1 near level of middle of anterior surface ; i 3 with crown 
area nearly double that of i 2 , and height nearly half that of 
canine, its anterior surface smoothly rounded but with pronounced 
nodule on inner side, its posterior surface gradually concave 
and without shelf -like extension ; a thickened ridge along its inner 
border and a low but somewhat trenchant ridge slightly outside 
of middle. Lower incisors forming continuous row between 
canines, their crown area increasing regularly from first to third, 
their height approximately equal ; each has a high inner cusp and 
a low outer tubercle best developed in i 3 ; posterior border slightly 



292 



CARMVORA 




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Fig. 53. 
Ursus arctos. Teeth nat. size. 



uitsus 293 

concave, with inner, middle and outer ridges, the inner and middle 
low and confluent in i v Canines large and strong, oval or 
slightly ovate in cross section, the longest diameter at level of 
alveolus equal to or slightly greater than distance from alveolus 
to median line of palate ; lower canine shorter and a little more 
curved than upper ; a slightly developed posterior and antero- 
internal longitudinal ridge, most evident in upper tooth ; no 
cingulum. First and third upper premolars small, flat topped, 
with slightly indicated cusp and posterior and antero-internal 
ridge, the crown area slightly less than that of smallest incisor, 
the first close to canine, the third close to large premolar ; third 
near middle of space between first and second, much smaller than 
the others and frequently deciduous, its crown indefinitely 
rounded ; fourth upper premolar with crown area about half that 
of first molar, its general outline triangular with apex directed 
forward, the outer side longest, the posterior border shortest, the 
contrast between them sometimes noticeable, in other instances 
slight, the three cusps lying near respective angles, the anterior 
highest (reaching level of main cusps of molars) and most robust, 
the posterointernal and postero-external abruptly smaller, sub- 
equal, the inner usually lower than the outer ; a small accessory 
tubercle usually present at posterior base of postero-outer cusp ; 
cingulum obsolete but usually visible along inner base of anterior 
cusp and outer base of postero-external cusp ; first and second 
lower premolars approximately like corresponding upper teeth in 
both size, form and position ; third very early deciduous, usually 
if not always absent in adult individuals ; fourth in contact with 
first molar and with from one-third to nearly one-half its crown 
area, the outline irregularly quadrilateral with well developed 
antero-external cusp nearly as high as main cusps of molars, a 
rudimentary antero-internal cingulum cusp, and a tuberculated 
ridge extending along outer side of crown from antero-external 
cusp to posterior border ; occasionally a similar ridge is present 
on inner side of crown, its anterior tubercle forming an evident 
cusp at inner posterior base of main cusp. First upper molar 
with crown much less than twice as long as broad, its outer side 
bi-convex, its inner side evenly rounded, its two outer cusps 
sub-equal in both height and diameter, the two inner cusps 
decidedly lower than outer, and less well defined, owing to the 
presence of a low, ridge-like tubercle between them ; both outer 
and inner cusps when unworn have distinct though low anterior 
and posterior trenchant ridge ; space between outer and inner 
cusps occupied by a rather well defined longitudinal groove, the 
surface of which is marked by irregular low ridges and furrows ; 
cingulum obsolete, but indicated in the regions between the 
cusps ; second upper molar nearly twice as long as broad, the 
anterior two-thirds approximately like first molar, with the same 
four cusps and intermediate longitudinal groove, the main axis 
of which is, however, in axis of tooth-row instead of slightly 



294 



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oblique to it ; postero-external cusp slightly smaller than antero- 
external, and contrast between outer and inner cusps less marked 
than in m l ; posterior third of crown occupied by a flattened heel, 
variable in form and size, but usually narrowing off, chiefly by 
slanting inward of outer border, to about half anterior width of 
tooth, and in some specimens bearing a low but evident third 
inner tubercle ; surface of heel sculptured by irregular small 
tubercles and furrows ; cingulum obsolete but usually evident 
along anterior half of inner border. First lower molar about as 
long as second but noticeably narrower, its crown showing more 
traces of the primitive trituberculate form than any of the other 
teeth ; protoconid and hypoconid wide apart, separated by a deep 
groove, the protoconid the highest cusp in the tooth, and with 
evident antero-external commissure; paraconid forming narrow 
anterior extremity of crown and provided with a distinct com- 
missure, similar to and joining that of protoconid ; metaconid 
subterete, without commissure, near to and slightly behind inner 
base of protoconid, a minute though evident accessory tubercle 
just in front of it ; entoconid like hypoconid, at extreme posterior 
edge of crown, the deep wide groove between it and metaconid 
with small accessory tubercle at its lowest point. Second lower 
molar about 1^ times as long as wide, its outline an irregular 
parallelogram with rounded-off corners, the surface of the crown 
occupied principally by a flattened, irregularly sculptured crushing 
surface, the cusps near border ; five cusps are usually well 
developed : a rather large antero-internal and antero-external 
opposite each other, and joined by a low transverse ridge ; a 
small postero-external and two smaller postero-internal cusps ; a 
small but evident accessory tubercle at anterior base of large 
antero-internal cusp, this tubercle not infrequently dividing into 
two. Third lower molar varying from rounded-triangular to 
ovate in outline, its area about two-thirds that of m 2 , its surface 
entirely flat except for a slightly raised rim which forms a 
small antero-internal cusp. 

Measurements.' — Adult male from Sweden (mounted) : head 
and body, 1900; tail, SO; hind foot, 195 ; ear, 90. For cranial 
measurements see Table, p. 294. 

Specimens examined. — Eight, from the following localities : — 
Sweden: Kvickjock, Norbotten, 1; no exact locality, 3 (B.M. and 
U.S.N.M.). 

Switzerland : Engadine, 1. 
Austeia-Hungaby : Near Hatszeg, Hunyad, 3. 

Remarks. — As regards the existence of geographical races of 
the large European bear it is impossible to form any opinion on 
the basis of the few specimens seen. Ursus arctos is related to 
the grizzly bear of North America, U. horribilis and its local 
forms, but is readily distinguishable by the relatively greater 
height of the frontal region and the consequently more abrupt 
slope of posterior half of dorsal profile, a character suggesting 



THALARCTOS 



297 



Ursus richardsoni ; forehead in most specimens rising abruptly 
above level of rostrum so as to produce a noticeable concavity in 
dorsal profile, but this character subject to marked variations, 
the exact nature of which is not fully understood. The skulls 
of the two species are of approximately the same size, those of 
U. horribilis perhaps averaging somewhat the larger. The teeth 
of the two animals are also much alike. 

9. Kvickjock, Norbotten, Stockholm Museum 90. 8. 1. 3. 

Sweden. (b). 

16. Sweden. (Lloyd.) Purchased (Stevens) . 62.3.29.7-8. 

9 st. Engadine, Switzerland. H. Justen (p). 86. 1. 23. 1. 

3 skeletons. Hatszeg, Transylvania, C. G. Danford (c). 78. 1. 16. 1-3. 
Hungary. 

Note on the Ursus formicarws of Bieleb. 

A peculiar small bear supposed to have been taken in the 
Canton of Grisons, Switzerland, has been described by Professor 
Bieler of the Lausanne Agricultural College as Ursus formicarius 
Eversmann.* Through the author's kindness I have had the 
opportunity to examine this skull. It is that of a rather young 
individual, apparently a female, with basal suture open, but with 
teeth showing slight indications of wear. In size it is smaller 
than in a female of the same age or slightly younger from 
Sweden (see Table of cranial measurements, p. 294), the inter- 
orbital region is much depressed, so that the frontal profile is 
nearly flat, and the mesopterygoid fossa is unusually broad and 
short, the width between pterygoids equal to a, little more than 
half distance from hamular to level of last molar. The teeth on 
the other hand are slightly larger than usual in females (see 
Table, p. 295) ; in form they show no special peculiarities. Small 
bears, presumably of this type, have been reported from Spain, 
northern Italy, Russia and Scandinavia, and have formed 
the basis of such names as Ursus formicarius, U. arctos minor 
and Myrmaretos eversmanni. Until more is known of them, as 
well as of the normal variation in ordinacy Ursus arctos, their 
status must remain in doubt. 



Genus THALARCTOS Gray. 

1825. TJialarctos Gray, Ann. of Philosophy, N.S., x, p. 62, July, 1825. 
1825. Tlmlassarctos Gray, Ann. of Philosophy, N.S., x, p. 339, November, 

1825. 
1896. Thalassiarchus Kobelt, Bericht Senckenberg. naturforsch. Gesellsch. 

Frankfurt am Main, p. 93 (Substitute for Thalarctos). 

Type species. — Thalarctos polaris Gray = Ursus maritimus 
Phipps. 

* Compte-Bendu des Seances du Sixieme Congres Internationale de 
Zoologie, Berne, 1904, p. 248. 1905. 



298 



CARNIVOEA 



Geographical distribution. — North Polar region, south to 
northernmost continental coasts. 

Characters. — Similar to Ursus, but cheek-teeth much less 
robust, the combined length of the two upper molars not equal 
to width of palate, and canines and incisors enlarged and more 
prehensive in general form. 

Remarks. — The genus Thalarctos, though not very strongly 
differentiated from Ursus, is a well defined and perfectly natural 
group. 

THALARCTOS MARITIMUS PhippS. 

1774. Ursus maritimus Phipps, Voyage toward North Pole, p. 185 (Spitz- 
bergen). 
1776. Ursus marinus Pallas, Beise duroh verschiedene Provinzen des 

russisehen Beiohs, in, p. 691 (Arctic Ocean). 
1792. Ursus polaris Shaw, Museum Leverianum, l, p. 7 (Benaming of 

marinus). 
1862. Thalarctos maritimus Gray, Gatal. Bones Mamm. Brit. Mus., p. 105. 
1908. ? Thalassarctos jenaensis Knottnerus-Meyer, Sitz.-Ber. Gesellsch. 

Naturforsch. Freunde, Berlin, p. 184, July, 1908 (Jena Island, 

Spitzbergen). 
1908. ? Thalassarctos spitzbergensis Knottnerus-Meyer, Sitz.-Ber. Gesellsch. 

Naturforsch. Preunde, Berlin, p. 184, July, 1908 (Seven Island, 

Spitzbergen). 
1910. Ursus (Thalassarctos) maritimus Trouessart, Paune Mamm. d'Europe, 

p. 66. 

Type locality. — Spitzbergen. 

Geographical distribution. — Arctic Ocean, south on floating ice 
occasionally to the northern coast of Norway. Details of 
distribution unknown. 

Diagnosis. — General characters as in the genus ; size very 
large ; colour uniform whitish or buffy. 

External characters. — Form longer and less heavy than in 
Ursus arctos, the neck noticeably longer and head longer and 
more pointed ; ear actually as well as relatively shorter ; fore 
foot with palmar tubercles and balls of toes essentially as in 
U. arctos but smaller ; pad on hind foot without backward 
continuation along inner portion of sole ; claws much less 
elongated than in U. arctos, not strongly curved, but with acute 
points and well developed cutting edges. Fur very dense, its 
texture almost seal-like in the short summer coat. 

Colour. — Entire animal a uniform whitish or buffy, the winter 
pelage tending to be a creamy-white, the summer coat yellowish 
buff. 

Skull. — The skull is considerably larger than that of Ursus 
arctos, with relatively longer brain-case, deeper, wider rostrum, 
and less elevated frontal region ; lambdal region less produced 
backward than in Ursus arctos, the condyles usually visible when 
skull is viewed from above. Base of brain-case essentially as in 
U. arctos, but portion at base of condyles more narrowed and 



THALAKCTOS 



299 



elongate. Palate noticeably broader than in Ursus arclos, a 
character made more conspicuous by the relative weakness of the 
teeth. Mandible with no special peculiarities except that the 




FIS. 54. 
Thalarctos maritimus. X J. 



lower margin is nearly straight throughout, the posterior concavity 
being very slightly indicated, and lower border of angular process 
only a little elevated above general outline. 

Teeth. — While the general character of the dentition differs 



300 



CAENIVOEA 



notably from that of Ursus arctos in the reduction of the molars 
and increased size and prehensiveness of the canines and incisors, 
the details of the individual teeth present little that is specially 




Fig. 55. 
Thalarctos maritirmis. x \. 



noteworthy. General form of upper incisors as in Ursus arctos, 
but points of i l and i 2 narrower and more hooked backward ; i 3 
with cusp more slender and ridges nearly obsolete ; lower incisors 



THALAECTOS 



301 



with lobes more sharply defined. Canines both above and below 
essentially similar to those of Ursus arctos, except for their greater 
size. Owing to the greater width of palate the proportion of 




Fig. 56. 
Thalarctos maritimus. X \. 



diameter of upper canine to palatal width is about as in the 
smaller-toothed animal. Small premolars showing no special 
peculiarities. Large upper premolar with relatively higher 



302 



CAHNIVOHA 





FIG. 57. 
ThalaretoB maritimus. Teeth nat. size. 



THALARCTOS 



303 



anterior cusp than in TI. arctos, its inner side more flattened, 
giving the tooth a more carnassial appearance ; postero-internal 
cusp relatively less developed. Large lower premolar essentially 
as in U. arctos but with somewhat more slender cusp. Molars 
differing from those of Ursus arctos in their smoother, less sculp- 
tured crushing surface, and slightly more trenchant cusps. Form 
of m 1 not peculiar, though outer cusps are higher and narrower 
and inner cusps relatively lower ; m 2 with inner cusps obsolete 
and heel relatively narrower and less developed. Anterior lower 
molar with metaconid and its accessory tubercle reduced to a low 
irregularly tuberculate ridge ; commissure of protoconid and 
paraconid obsolete ; hypoconid and entoconid smaller and much 
nearer together than in Ursus arctos, though separated from 
anterior cusps by a normally wide interval, in which, however, 
there are no definitely formed accessory tubercles. Second lower 
molar with the same elements as in U. arctos except for the 
absence of all trace of an intermediary tubercle on inner side of 
crown. Third lower molar with crown nearly flat, its margin 
showing only the faintest trace of antero-internal and antero- 
external elevations. 

Measurements. — Adult male from Behring Strait (mounted) : 
head and body, 2670 ; tail, 90 ; hind foot, 370 ; ear, 80. For 
cranial measurements see Table, p. 294. 

Specimens examined. — Nine, from the following localities : — Spitzbergen, 
1 (U.S.N.M.); Griffin Bay, Wellington Channel, 1; Melville Island, 1; 
Arctic Ocean, 1 ; no history, 5. 

[The Museum specimens appear all to have come from the 
American side of the Atlantic] 



Family CANID^E. 
1821. Canidm Gray, London Medical Eepository, xv, p. 301, April 1, 1821. 

Geographical distribution. — Essentially cosmopolitan ; in E arope 
west to Ireland. 

Characters. — Larger cheek-teeth of a combined trenchan t and 
crushing type, the last upper premolar and first lower molar 
strongly differentiated as carnassials, the former 3-rooted, its 
inner lobe in front of middle of crown, its position, somewhat 
posterior to level of anteorbital foramen, at point of greatest 
mechanical efficiency ; auditory bulla moderately or considerably 
inflated, without septum ; form rather light, the legs long ; size 
moderate ; feet digitigrade ; toes, 5-4 or 4-4. 

Remarks. — Notwithstanding its wide distribution the family 
Canidse is not rich in genera. About a dozen are now recognized, 
three of which occur in Europe. 



304 



CAENIVOEA 



KEY TO THE EUROPEAN GENERA OF CANIDJE. 



Interorbital region distinctly elevated ; postorbital processes 

convex above ; pupil of eye round Canis, p. 304. 

Interorbital region not elevated ; postorbital processes not 
convex above ; pupil of eye elliptical. 
Postorbital processes flat or very slightly concave above ; 
forehead rising abruptly above level of rostrum ; ear 

rounded Alopex, p. 318. 

Postorbital processes distinctly concave above ; forehead 

rising gradually above level of rostrum ; ear pointed Vulpes, p. 325. 



Genus CANIS Linnseus. 

1758. Canis Linnseus, Syst. Nat., i, 10th ed., p. 38 (type by tautonymy 

G. familiaris). 
1816. Lupus Oken, Lenrb. d. Naturgesch., in, pt. 2, p. 1039 (Canis lupus, 

by tautonymy). 
1837. Vulpicanis Blainville, Ann. Sci. Nat., Paris, 2nd ser., Zool., vni, 

p. 279, November, 1837 (Canis aureus Linnseus). 
1839. Lyciscus H. Smith, Jardine's Naturalists' Library, Mammals, ix, 

p. 160 (Canis latrans Say). 
1839. Thous H. Smith, Jardine's Naturalists' Library, Mammals, ix, 

p. 193 (Canis anthus P. Cuvier). 
1839. Sacalius H. Smith, Jardine's Naturalists' Library, Mammals, ix, 

p. 213 (Canis aureus Linnseus). 
1841. Oxygous Hodgson, Calcutta Journ. Nat. Hist., n, p. 213 (Canis 

aureus Linnseus). 
1857. Canis Blasius, Saugethiere Deutschlands, p. 177. 

1868. Neocyon Gray, Proc. Zool. Soc, London, p. 506 (Canis latrans Say). 

1869. Dieba Gray, Catal. Carn. Pachyd. and Edentate Mamm. Brit. Mus., 

p. 180 (Canis anthus P. Cuvier). 

Type species. — Canis familiaris Linnseus. 

Geographical distribution. — Nearly as in the family, but 
absent from the Malay Archipelago and South America ; in 
Europe west within historic times to Great Britain, but now 
restricted to the continent. 

Characters. — Skull heavy and deep (depth of brain-case more 
than one-third condylobasal length) ; interorbital region thickened 
and elevated, the frontal sinuses rather large, the postorbital 
processes thick, convex above, their edges rounded off; dorsal 
profile of forehead rising rather abruptly and noticeably above 
level of rostrum ; dental formula ; * g, e 1=5, pm tl in || = 42 ; 
teeth heavy and large, the length of carnassial and upper molars 
together contained about 2 J times in palatal length ; canines 
robust and not specially elongated, the point of upper tooth 
extending scarcely beyond middle of mandibular ramus when 
jaws are closed (fig. 65). 

Remarks. — Much uncertainty exists at present with regard 
to the limits of the genus Canis. As here defined the group 
includes the domestic dogs, the true wolves, the American prairie 



CANIS 305 

wolves, and the Old World jackals. Two species are known 
to occur in Europe, one of which is represented by several 
geographical races. 

KEY TO THE EUROPEAN FORMS OP CANIS. 

Condylobasal length of skull less than 200 mm. ; teeth 
not so large as in the largest domestic dogs (length 
of upper carnassial 17 to 18 mm.) ; oingulum on 
outer margin of m l broad and conspicuous (South- 
eastern Europe ; Jackal) C. aureus, p. 315. 

Condylobasal length of skull more than 200 mm. ; 
teeth larger than in the largest domestic dogs 
(length of upper carnassial 25 to 27 mm.) ; cingu- 
lum on outer border of m 1 narrow, tending to be 
incomplete at middle (Distribution general; true 

Wolves) C. lupus, p. 305. 

Size rather small (exact dimensions unknown) 

(Southern Spain) C. I. deitanus, p. 315. 

Size large. 
White of throat not extending uninterruptedly 

on to cheek (Central and northern Europe) ... C. I. lupw, p. 313. 
White of throat extending uninterruptedly on to 

cheek (Spain except extreme south) C. I. signatus, p. 314. 

CANIS LUPUS Linnaeus. 
(Synonymy under subspecies.) 

Geographical distribution. — Originally throughout Europe from 
Ireland eastward and across Asia, now exterminated in the 
British Islands, Holland and Denmark. 

Diagnosis. — Condylobasal length of skull more than 200 mm. 
(220 to 255 mm.) ; cheek-teeth larger than in the largest races 
of domestic dogs, the upper carnassial 25 to 27 mm. in length, 
but structure not peculiar, the upper molars with narrow, incon- 
spicuous cingulum on outer side (fig. 61). 

External characters. — General form essentially as in domestic 
dogs of the " collie " type. Ear moderately long, erect, somewhat 
pointed, extending about to eye when laid forward. Muzzle pad 
completely bare. Fore foot with third and fourth digits sub- 
equal and longest, second and fifth shorter, the large pad-like 
balls fitting closely between those of third and fourth and the 
slightly trilobed, heart-shaped main pad, the combined area of 
balls of digits greater than that of pad ; thumb much shorter 
than other digits, the nail smaller, but not peculiar in form, its 
extremity not reaching level of posterior border of main pad, its 
ball scarcely indicated, no pad at its base ; wrist pad single, near 
outer side, its area somewhat more than half that of ball of toes. 
Hind foot essentially like fore foot, but hallux and posterior pad 
absent. Claws robust, slightly curved, sub-equal throughout. 
Pads and balls narrowly edged with short hair. 

Colour. — General colour of upper parts, tail, and outer surface 
of legs yellowish brown or buff, darker along median region of 



306 



CAENIVOKA 



back, on posterior portion of head and outer surface of ears, 
lighter and more inclined toward greyish at sides of shoulders 
and between ears and eyes ; longer hairs of back and sides black- 
tipped, producing an evident dark shading over middle of back, 




FI0. 58. 
Camis lupus. X *. 



especially behind middle and at base of tail ; pencil narrowly 
clear black, rest of tail essentially like back. TJnderparts and 
inner surface of legs pale buff or buffy white, not strongly 
contrasted with sides, the chin and interramia usually grizzled, 



CANIS 307 

frequently margined with blackish ; upper lip to muzzle pad and 
including lower half of cheek dull whitish, usually not very 
different from throat ; inner surface of ear light buff. 

Skull. — In general aspect the skull differs slightly if at all 




Via. I 
Canis lupus, x i. 



from that of some of the larger races of domestic dogs, though 
often attaining a greater size. The rostrum, however, appears 
to be relatively less robust than in dog skulls of approximately 
the same length. Dorsal profile rising gradually from nares to 

x 2 



308 



CARNIVORA. 



just in front of orbit, then abruptly to a little in front of bregma, 
behind which it is nearly flat to strongly overhanging lambdal 
region. Depth of brain-case through auditory bulla about i\ 




Fig. 60. 
Canis lupus. X +. 



times that of rostrum behind canine, and about equal to mastoid 
breadth. Brain-case rather elongate ovate in outline when 
viewed from above, its breadth above roots of zygomata about 
li times that of rostrum over canines and approximately equal 



CANIS 309 

to distance from bregma to most posterior point of occiput. 
Posterior portion of occiput strongly concave when viewed from 
the side, the condyles nearly hidden beneath the projecting 
lambdal region. Floor of brain-case with no specially noteworthy 
features, the auditory bullre sub-circular in outline, with short but 
evident meatal tube, their surface evenly inflated (more so than 
usual in domestic dogs) except for an evident flattening on antero- 
external aspect. Sagittal and lambdoid crests well developed, 
the former dividing just in front of bregma into two ridges 
curving outward to form posterior border of postorbital processes. 
Interorbital region moderately elevated, well defined, with distinct 
longitudinal concavity between raised and thickened postorbital 
processes. Zygomata widely spreading, the greatest zygomatic 
breadth (at level of anterior glenoid edge) a little more than half 
greatest length of skull ; orbital process well developed, the 
orbit surrounded by bone through about four-fifths of its 
circumference. Rostral breadth at canine about equal to depth 
at front of carnassial ; premaxillary extending posteriorly to 
about middle of nasal ; mnxillary extending back nearly to middle 
of orbit, slightly exceeded by nasal ; anteorbital foramen about 
9x5 mm. in diameter, over posterior root of third premolar. 
Palate moderately wide, nearly flat, not extending posteriorly 
beyond level of last molar, terminating in an obscure median 
spine ; incisive foramina between canines, 11 to 17 mm. in length, 
their combined breadth usually a little less ; mesopterygoid fossa 
rather more than one-third as long as palate, considerably 
narrower posteriorly than anteriorly. Mandible strong, but not 
remarkably robust, the depth at posterior edge of carnassial 
contained about six times in length ; symphysis rather long ; 
coronoid process high, the depth of mandible through its middle 
noticeably greater than distance from last molar to back of 
condyle ; angular process heavy, nearly horizontal, distinctly 
raised above level of under margin of ramus. 

Teeth. — The teeth are relatively larger than in any of the 
races of domestic dogs, though in form they show no tangible 
features by which they may be distinguished. Upper incisors form- 
ing a continuous, slightly convex row, the outer tooth separated 
from canine by a distinct space ; size, when viewed from in front, 
increasing regularly from first to third, but third abruptly much 
larger than the others in cross-section and nearly half the size of 
canine ; anterior surface of i 1 and i 2 slightly more than half as 
wide as high, smoothly rounded off, the cutting edge narrow but 
not acute ; a small but distinct secondary lobe at each side of 
front aspect, that of inner side a little below middle, that of 
outer side about equally above ; posterior surface of crown con- 
cave longitudinally though without backward-projecting basal 
shelf ; a well developed median longitudinal rib, and a low but 
noticeable cingulum, the latter terminating abruptly and forming 
the lobes seen in front view ; outer incisor with no secondary 



310 



CAEKIVOEA 



lobes, its general form intermediate between tbat of canine and 
of inner incisors. Lower incisors forming continuous row between 
canines, the three teeth essentially alike in form, but increasing 





Fia. 61. 
Canis lupus. Teeth nat. size. 



regularly in size from first to third though less conspicuously 
than in the case of the upper incisors ; crowns (viewed from in 
front) about twice as high as wide, distinctly bilobed, the outer 



CANIS 311 

lobe scarcely half as wide as inner ; on inner tooth the outer lobe 
is nearly level with cutting edge, on second it lies slightly above 
middle of crown, and on third slightly below middle of crown ; 
posterior surface oblique, slightly concave, with noticeable longi- 
tudinal furrow extending back from notch between lobes. Canines 
large, usually 15 mm. or more in diameter at alveolus and about 
twice as long, a size rarely if ever attained in domestic dogs, 
their surface smooth except for a low antero-internal and 
posterior-median logitudinal ridge, the upper teeth slightly longer 
and less recurved than the lower. Premolars moderately spaced 
except that pm 3 is nearly or quite in contact with the 
carnassial ; first, second and third teeth essentially alike in the 
two jaws, those of the mandible, however, slightly the less robust ; 
first premolar both above and below single-rooted, the crown 
simple, that of pm l subterete, that of pm 1 nearly twice as long as 
broad, the height in both slightly less than length, the crown area 
approximately the same as that of corresponding inner incisor, the 
small cusp a little in front of middle and with slightly developed 
anterior and posterior ridge. Second and third premolars two- 
rooted, the crown about twice as long as wide, sub-elliptical in 
outline, the inner margin sometimes (especially in pm 3 and pm 2 ) 
slightly concave, the long axis nearly parallel with sagittal plane 
except in pm 3 , which is obliquely set ; main cusp a little in front 
of middle of crown, its height distinctly more than half length of 
crown, its anterior and posterior cutting ridge well developed, the 
posterior bearing a distinct secondary cusp situated over middle 
of posterior root and relatively larger in lower than in upper 
teeth ; a slight shelf-like projection behind secondary cusp ; pm i 
similar to pm 3 but considerably larger, its secondary cusp 
better developed and succeeded by a small but evident postero- 
basal cusp springing from the posterior edge of crown ; cingulum 
of all the smaller premolars complete though low and incon- 
spicuous. Upper carnassial large and robust, the length of crown 
along middle slightly more than twice greatest breadth exclusive 
of antero-internal lobe, the main axis of the tooth extending 
evidently through middle of crown, so that the small, cuspless 
inner lobe stands as an offset, slightly breaking the symmetry of 
the outline ; main cusp slightly behind middle of crown, its 
height more than half length of tooth, its axis slanting distinctly 
backward, its anterior and outer surfaces evenly convex except 
for the rudimentary longitudinal ridge on basal two-thirds of 
front, its inner surface, together with that of posterior cusp, 
flattened ; posterior cusp low and robust, obscured by its very 
high nearly horizontal commissure which meets the somewhat 
shorter but equally trenchant commissure of main cusp at an 
angle of about 75° ; cingulum complete, though low and incon- 
spicuous. Lower carnassial narrower than upper but with equally 
high crown, the most elevated portion in front of middle instead 
of behind it ; protoconid large and robust, resembling main cusp 



312 



CARNIVORA 



of upper carnassial, but with well developed cutting edge both, 
in front and behind ; paraconid near middle of anterior portion 
of crown ; its general form like posterior cusp of upper carnassial, 
its commissure bearing essentially the same relation to that of 
protoconid as in the case of the two cusps of the upper tooth, 
except that the relative lengths of the cutting edges is reversed ; 
metaconid small but evident, at postero-internal base of proto- 
conid ; hypoconid and entoconid low, occupying the posterior edge 
of a well developed though relatively small heel (area of heel 
scarcely more than one-third that of anterior portion of tooth) 
separated from the cusps of the main triangle by a wide transverse 
groove ; crown area of entoconid equal to about half that of 
hypoconid, its cusp approximately the same size as that of meta- 
conid. Second lower molar essentially like heel of carnassial 
but larger, its two anterior cusps corresponding in size and form 
with hypoconid and entoconid of the large tooth, the posterior 
edge of its crown with a small outer cusp resembling the antero- 
inner, and sometimes with a slightly developed inner ridge or 
rudimentary fourth cusp. Third lower molar single-rooted, the 
crown subterete, about as large as that of first premolar, with low 
central cusp and rudimentary longitudinal ridge. First upper 
molar large, with high outer two-cusped sectorial portion and low 
inner crushing portion, the two areas sharply differentiated, the 
antero-posterior diameter of the outer decidedly greater than that 
of inner ; paracone and metacone conical, terete, with slightly 
developed anterior and posterior cutting ridges, the area and 
height of metacone about two-thirds those of paracone, the width 
of base of which is at least equal to width of inner portion of 
tooth ; protocone very low, with low but distinct anterior and 
posterior commissures, each of which joins cingulum at base of 
corresponding large outer cusps, and each of which bears an 
intermediate cusp soon disappearing with wear, the posterior 
intermediate cusp larger and more definite in form than anterior 
cusp ; hypocone ridge-like, at postero-inner border of crown, 
separated from protocone and its posterior commissure by a deep 
groove. Second upper molar with about half the crown area of 
first, its elements essentially the same, though so reduced that 
the paracone is scarcely larger than protocone of large tooth, 
intermediate cusps on commissures of protocone are barely 
indicated, and hypocone is not distinguishable as a cusp distinct 
from the cingulum. In both molars the cingulum on outer border 
is narrow and inconspicuous relatively to the broad cusps ; in 
region between paracone and metacone of m 1 it is usually 
obsolete (compare figs. 61 and 62). 

Remarks. — The material available for study has been so poor 
that I have found it impossible to come to any conclusion with 
regard to the existence of local forms of the European Wolf. 
The following races have been distinguished by Mr. Cabrera. 
There seems to be no good reason to doubt their validity. 



CANIS 



313 



The only known characters by which the skull of Canis lupus 
can be distinguished from that of the larger domestic dogs is the 
greater average general size and the relatively larger teeth. In 
a dog's skull with condylobasal length of 230 mm. the length of 
upper and lower carnassials is respectively 21 • 6 and 25 - mm. 
In ten skulls with condylobasal length of more than 200 mm. 
the average and extremes for these teeth are : upper, 20 • 5 
(19-22); lower, 24 -0(22 -8-26 -0).* In all the dog skulls which 
I have examined, representing such different breeds as the pug, 
fox-terrier, bloodhdund, mastiff, ancient Egyptian, ancient 
Peruvian, Eskimo (Greenland and Alaska) and American Indian, 
the teeth are strictly of the wolf type, never showing any 
approach to that of the jackal (fig. 62). 



Canis lupus lupus Linnaeus. 

1758. [Canis] lupus Linnaaus, Syst. Nat., I, 10th ed., p. 39 (Sweden). 
1792. C[anis] lupus flavus Kerr, Anim. Kingd., p. 137 (France and 

Germany). 
1804. Gcmis lupus niger Hermann, Observ. Zool., p. 32. Not of Kerr, 

1792 (Forest of Hagenau, Alsace, Germany). 
1839. ? [Canis lupus] var. canus de Selys-Longchamps, Etudes de Micro- 

mamm., p. 144 (nomen nudum). 
1839. 1 [Canis lupus] var. fulvus de Selys-Longchamps, Etudes do Micro- 

mamm., p. 144 (nomen nudum). 
1841. Lupus orientalis Wagner, Schreber's Saugthiere, Suppl., n, p. 367 

(Europe). 
1857. Canis lupus Blasius, Saugethiere Deutschlands, p. 180. 
1863. [Canis lupus] var. major Ogerien, Hist. Nat. du Jura, in, p. 59 

(Lower slopes of the Jura). 
1863. [Canis lupus] var. minor Ogerien, Hist. Nat. du Jura, in, p. 69 

(Higher portions of the Jura). 
1897. Canis lupus minor Mojsisovics von Mojsvar, Thierleben der osterr.- 

hung. Tiefebenen, p. 241 (Southern Hungary). Based on tbe 

"Rohrwolf," an animal supposed to be smaller and greyer than 

true lupus. 
1910. Canis lupus and C. lupus lycaonf Trouessart, Faune Mamm. 

d'Europe, p. 90. 

Type locality. — Sweden. 

Geographical distribution. — Northern and central Europe, 
exact limits of range unknown ; formerly west to Ireland. 

Characters. — Size maximum for the species; general colour 

* Winge (Danmarks Fauna, Pattedyr, p. 123, 1908) states that in the 
skull of a dog from a prehistoric grave (Iron Age) in Denmark, the length 
is 209, and that of the two carnassials 20 and 22-5 respectively, while in a 
rather large modern "great Dane " the corresponding measurements are 
255, 22 and 28. This author (p. 124) regards the domestic dogs as derived 
from Canis aureus. 

t Applied to the wolf of the Pyrenees; but Schreber's plate lxxxix, 
the basis of the name, is a copy of Bufion's plate xli, representing an 
animal brought alive to Paris from Canada. 



314 



CARNIVORA 



not markedly tawny ; white of throat not extending to cheeks. 
The few skulls examined agree with Asiatic specimens in having 
the outer cusps of m 1 moderately large, the paracone with trans- 
verse diameter of base about equal to width of large flattened 
portion of crown. 

Measurements. — For cranial measurements see Table, p. 316. 

Specimens examined. — Pour skulls, from the following localities : — 
Sweden: No exact locality, 2 (U.S.N.M.). 
Russia: No exact locality, 1 (U.S.N.M.). 
Italy : Near Sassello, Liguria, 1 (Genoa). 

Canis lupus signatus Cabrera. 

1907. Canis lupus signatus Cabrera, Bol. Real Soc. Espan. Hist. Nat., 

Madrid, vn, p. 195. 
1910. Canis lupus signatus Trouessart, Faune Mamm. d'Europe, p. 91. 

Type locality. — Escorial, Madrid, Spain. 

Geographical distribution. — Central Spain. 

Characters. — Size and general appearance as in Canis lupus 
lupus ; colour a more tawny brown than in the northern animal, 
particularly on muzzle ; white of throat extending uninterruptedly 
to cheeks.* 

Measurements. — Type (adult male), from Cabrera : head and 
body, 1230; tail, 400; hind foot, 265; ear, 125. Young adult 
male and female from Province of Burgos, Spain : head and 
body, 1130 and 1180 ; tail, 350 and 380 ; hind foot, 225 and 220 ; 
ear 120 and 115. For cranial measurements see Table, p. 316. 

Specimens examined. — One from Seville, Spain, and two from Province 
of Burgos, Spain. 

Remarks. — In dentition the Seville specimen differs from all 
the other Old World wolves with which I have compared it in 
the unusual development of the outer cusps of the upper molars. 
The transverse diameter of paracone in m 1 conspicuously exceeds 
width of the small inner portion of tooth. Mr. Cabrera informs 
me that the type shows much the same peculiarities. This 
character is also present, though less pronounced, in the two 
skulls from Burgos, which further differ from northern specimens 
in the smaller size and more globular form of the auditory 
bullse. 

<J. Seville, Spain. (A. Ruiz.) Lord Lilford (p). 95. 3. 3. 6. 
<S, ?. Riocabado, Burgos. Hon. N. C. Roths- 11. 10. 5. 1-2. 

(Bev. S. Gonzales.) child (p). 



* In the Burgos specimens the colour is not unusually tawny : back 
and sides a coarse mixture of black, whitish, ochraceous-buff , and drab grey 
(underfur), the black and whitish most conspicuous along back, the 
ochraceous-buff on legs and feet (clear and unmixed on latter) ; ear 
ochraceous-rufous on outer side (darker and duller at tip), pallid 
ochraceous-buff on inner surface ; throat and lower half of cheeks the same 
pallid ochraceous-buff ; chin and interramia blackish. 



315 



Canis lupus deitanus Cabrera. 



1907. Canis lupus deitanus Cabrera, Bol. Real Soc. Espan. Hist. Nat., 

Madrid, vn, p. 197. 
1910. Canis lupus deitanus Trouessart, Faune Mamm. d'Europe, p. 91. 

Type locality. — Moratalla, Murcia, Spain. 

Geographical distribution. — Now known from the type locality 
only. 

Diagnosis. — Smaller than Canis lupus lupus and brighter in 
colour, the general appearance much as in C. aureus. 

Measurements. — Unknown . 

Specimens examined. — I have seen the two living examples in Madrid 
on which the form was based. 

Remarks. — Tn general appearance the two Moratalla wolves 
are strikingly different from Can in lupus. Unfortunately their 
true characters are not yet known. 

CANIS AUREUS Linnseus. 

1758. [Cams'] aureus Linnseus, Syst. Nat., i, 10th ed., p. 40. 

1835. Canis aureus var. moreotica I. Geoffroy, Exped. Sci. de Mor6e. Zool., 

pi. i (Morea, Greece). 
1841. C[anis] dalmatinus Wagner, Sohreber's Siiugthiere, SuppL, n, p. 383 

(Dalmatia). 
1841. Clams'] griecus Wagner, Sohreber's Siiugthiere, Suppl., n, p. 383 

(Peloponesus). 
1892. Canis aureus balcanicus Brusina, Glasnik Hrvatskoga Naravoslovnoga 

Drustva, Zagreb, vn, p. 317 (Drava River, Croatia). 

Type locality. — Province of Lar, Persia. 

Geographical distribution. — India and westward through Asia 
Minor to the Balkan Peninsula, north to Heves Comitat, 
Hungary. 

Diagnosis. — Smaller than Canis lupus (condylobasal length of 
skull less than 200 mm.) ; teeth not equal to those of the larger 
domestic dogs in size, the upper molars with wide, conspicuous 
cingulum on outer side (fig. 62). 

Colour. — Upper parts buffy cinnamon, clouded by black hair 
tips along dorsal region, nearly clear on sides, and becoming clear 
bright cinnamon on outer surface of legs and ear and on area 
ai'ound and behind base of ear ; muzzle more heavily washed 
with black y from middle of back to base of tail the cinnamon is 
replaced by whitish, causing a rather noticeable contrast between 
this region and the surrounding parts when fur is disarranged ; 
tail essentially like back, the basal half above greyish, the 
terminal half and underside buffy much overlaid with black ; 
pencil blackish ; underparts and inner side of legs dull buffy 
grey ; chin blackish ; feet dull buffy. 



316 



CAKNIVOEA 





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318 



CAENIVORA 




Skull. — The skull is much smaller than that of Ganis lupus. 
In form it differs slightly in the less elevated frontal region and 
somewhat more inflated auditory bullae. 

Teeth. — While agreeing with those of Ganis lupus in general 

form and in the position of the cusps, the teeth 

are on the whole more trenchant in character, as 

shown by the general tendency toward narrowness 

of crown and prominence of ridges. This is 

particularly noticeable in the upper molars, in 

which the large cusps are relatively higher, more 

slender, and less terete than in Ganis lupus, their 

cutting ridges much more developed ; transverse 

diameter of metacone in m 1 noticeably less than 

width of inner portion of crown ; cingulum on 

outer border of both upper molars wide and 

conspicuous in contrast with narrow cusps, 

Fig. 62. showing no tendency to become obsolete in 

Canis aureus, region between paracone and metacone. Lower 

La clee ™teeth ary carna ssial with metaconid actually as well as 

Nat. size. relatively larger than in Canis lupus, and posterior 

heel with area equal to nearly half that of anterior 

portion of tooth, its cusps strongly developed. 

Measurements. — For cranial measurements see Table, p. 316. 

Specimens examined* — One from Greece (Piraeus) ; numerous others 
from Asia Minor and India. 

Remarks. — The single specimen from Greece agrees sufficiently 
with a series of five from Khotz, near Trebizond, Asia Minor, 
to make it appear unwise, in the absence of more satisfactory 
material, to use one of the Balkan names. 

1. Piraeus, Greece. (0. Mottaz.) Hon. N. 0. Roths- 8. 10. 2. 49-50. 

child (p). 



Genus ALOPEX Kaup. 

1829. Alopex Kaup, Entw.-Gesch. und Natiirl. Syst. Europ. Thierw., i, 

p. 83. 
1857. Leucocyon Gray, Proc. Zool. Soc, London, p. 512. 

Type species. — Ganis lagopus Linnaeus. 

Geographical distribution. — Arctic region of both Old and New 
Worlds ; in Europe south to southern Norway and Sweden. 

Characters. — Skull intermediate in general form between that 
of Canis and Vulpes ; occipital depth about one-third condylo- 
basal length ; interorbital region more elevated than in Vulpes 
owing to greater inflation of the frontal sinuses ; postorbital 
processes thin, flat or slightly concave above, with bead-like, 
overhanging edges ; dorsal profile of forehead rising abruptly 



ALOPEX 319 

above rostrum as in Canis ; teeth moderately heavy and large, 
the length of carnassial and upper molars together contained 
about 2£ times in palatal length ; canines and incisors inter- 
mediate between those of Canis and Vulpes (see fig. 65) ; external 
form fox-like, but ear short and rounded, not conspicuously 
overtopping the surrounding fur. 

Remarks. — Although in most respects intermediate between 
Canis and Vulpes the Arctic foxes form such a natural and 
circumscribed group that it seems desirable to set them apart as 
a distinct genus.* Half a dozen species have been described, two 
of which come within the scope of the present work. 

KEY TO THE EUROPEAN SPECIES OP ALOPEX. 

Condylobasal length of skull about 130 in males, 

124 in females (Scandinavia and Finland)... A. lagopus, p. 319. 
Condylobasal length of skull about 120 in males, 

114 in females (Spitzbergen) A. spitzbergenensis, p. 324. 

ALOPEX LAGOPUS Linnaeus. 

1758. [Canis] lagopus Linnaeus, Syst. Nat., i, 10th ed., p. 40 (Lapland). 
1816. V[ulpes] arctica Oken, Lehrb. d. Naturgeseh., in, pt. 2, p. 1033 

(Renaming of Canis lagopus). 
1820. C[anis] vulpes cosrulea Nilsson, Skand. Fauna, I, p. 88 (Lapland). 
182Y. [Canis lagopus] & argentms Billberg, Synopsis Faunae Scandinaviae, 

p. 14 (Lapland). 
1910. Vulpes lagopus Trouessart, Faune Mamm. d'Europe, p. 96. 

Type locality. — Lapland. 

Geographical distribution. — Arctic portions of the mainland 
of Europe and Asia ; in Europe south along the mountains of 
Scandinavia to south-western Norway, and as an occasional 
visitant as far as southern Sweden. 

Diagnosis. — General characters as in the genus ; condylobasal 
length of skull about 130 mm. in males, about 124 mm. in 
females. 

External characters. — In general external characters Alopex 
lagopus resembles Vulpes vulpes, though the muzzle is less 
elongated, and the low, rounded ears (not extending to eye when 
laid forward) impart a somewhat un-fox-like appearance to the 
head. Fur very dense, the underfur in summer about 12 mm. 
deep on back, nearly twice as deep and somewhat looser in 
texture on sides (in winter longer throughout) ; longer hairs 
rather sparse, not concealing underfur. Tail bushy, with 
abundant underfur. Feet as in Canis, but claws longer and 
more slender, and entire palm and sole covered with a dense' 
woolly growth of hair, 15-17 mm. deep in winter, shorter in 

* See Collett, Norges Pattedyr, p. 275, 1911, for account of apparently 
complete sterility of Arctic fox male with Red fox female, a fact which 
indicates a fundamental physiological difference between the two animals. 



320 



CAENIVOEA 



summer when it sometimes wears away sufficiently to expose 
balls of toes, and parts of pads. 

Colour. — Summer pelage : ground colour of back, shoulders, 
and outer side of legs drab, darkening to about prout-brown or 




FIG. 63. 
Alopex lagopus. X $. 



dark bister on feet, head, chin, interramial region and outer 
surface of ears, the face thickly sprinkled with whitish hairs, 
especially on cheeks and between eyes, the interramial region 
tinged with slaty grey ; each of the longer hairs of back with 



321 



one cream-buff sub-terminal annulation, producing a noticeably 
speckled appearance throughout dark area ; flanks with a few 
long, entirely buff hairs ; sides of body and of neck light cream- 
buff tinged with clay-colour, in striking contrast with dark areas, 
the buff lateral area divided at shoulder by band about 60 mm. 
wide where drab of back crosses to leg ; anteriorly the buff 
lateral area extends to about level of ears where it abruptly gives 
place to dark brown of head ; inner surface of ear light buffy 
grey ; underparts and inner surface of legs buffy greyish, slightly 
contrasted with sides ; under surface of tail like sides of body, 
upper surface essentially like back at base (where line of 
demarcation is well defined), but becoming tinged with buffy 
toward tip. Winter pelage : entirely white, tinged with yellowish 
on throat, neck and face ; underf ur and posterior surface of ear 
light drabby grey. In the " blue " phase the entire animal is at 
all seasons a bluish drab, usually washed with sepia on head and 
feet, and sprinkled with pure white hairs on face, chin and throat. 

Skull. — The skull is shorter and narrower than that of Vulpes 
vulpes, but of nearly equal depth, a difference in form due in part 
to the greater elevation of the interorbital region and in part to 
the relatively greater depth of brain-case (depth equal to one- 
third condylobasal length instead of noticeably less as in 
V. vulpes) ; muzzle less produced 
than in the common fox and rela- 
tively wider proximally ; zygomata 
less abruptly spreading anteriorly, 
so that the region of greatest 
zygomatic breadth is noticeably at 
glenoid level. In other respects 
there is essential agreement with 
the skull of V. vulpes. Anteorbital 
foramen over space between pm 3 
and pn& ; auditory bulla relatively 
as large as in the larger animal ; 
depth of brain-case through bulla 
equal to greatest breadth above 
roots of zygomata ; postorbital pro- 
cesses slightly less concave on 
underside and somewhat more 
flattened above ; mandible with 
ramus relatively a little deeper 
and more compressed than in the 
common fox. 

Teeth. — In general the teeth 
closely resemble those of Vulpes 
vulpes except for their slightly 
smaller size. Incisors both above 
and below with relatively wider crowns than in the common fox, 
the cingulum of i 1 and i 2 much more developed and forming a 

T 




no. 64. 
Jlopex lagopus. Teeth. Nat. size. 



322 



OARNIVOEA 



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324 



CARNIVORA 



noticeable inner and outer secondary cusp (the former soon 
wearing away in i 1 ) much as in Canis ; along posterior border of 
crown the cingulum shows a strong tendency to develop irregular 
tubercles, a condition not observed in Vulpes vulpes. Canines 
and premolars with no special peculiarities, the form of the upper 
carnassial quite as in V. vulpes. Lower carnassial with posterior 
heel narrower than main portion of tooth but not otherwise 
peculiar. First upper molar with metaconule relatively less 
developed than in Vulpes vulpes. 

Measurements. — For cranial measurements see Table, p. 322. 

Specimens examined. — Ten, from the following localities : — 
Norway : Roros, Trondhjem, 1 ; Tolgen, Hedemarken, 2 ; Dovre, 2 ; 
Egersund, Stavanger, 1. 

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

Lapland : No exact locality, 3 skulls (B.M. and U.S.N.M.). 

?. Roros, Trondhjem, Nor- Christiania Museum 95. 11. 14. 1. 
way. (e). 

skull. Egersund, Stavanger. K.H. Schaanning(c). 11.6.3.12. 

2 9. Dovre. Christiania Museum 95. 11. 14. 2-3. 

(E). 

2 imm. Tolgen, Hedemarken. Christiania Museum 93. 3. 1. 1. 

(e). 0. 5. 2. 1. 

2 skulls. Lapland. Wheelwright (c). 64. 3. 8. 3-4. 



ALOPEX SPITZBERGENENSIS Barrett-Hamilton and Bonhote. 

1799. ? Canis fuliginosus Beohstein, Thomas Pennant's allgem. Uebersicht 
d. vierfuss. Thiere, I, p. 270 (Iceland). 

1799. ? Canis groenlandicus Bechstein, Thomas Pennant's allgem. Ueber- 
sicht d. vierfuss. Thiere, i, p. 270 (Greenland). 

1898. Canis lagopus spitzbergenensis Barrett-Hamilton and Bonhote, Ann. 
and Mag. Nat. Hist., 7th ser., I, p. 287, April, 1898 (Spitzbergen). 
Type in British Museum. 

1910. Vulpes lagopus spitzbergensis and ? V. lagopus fuliginosus Trouessart, 
Paune Mamm. d'Europe, p. 97. 

Type locality. — Spitzbergen. 

Geographical distribution. — Spitzbergen ; also Iceland and 
Greenland ? 

Diagnosis. — Like Alopex lagopus but smaller, condylobasal 
length of skull about 120 in males, about 114 in females. 

Colour. — Type (summer pelage) : colour pattern well defined 
and exactly as in A. lagopus, but dark areas wood-brown against 
which the cream-buff annulations of longer hairs make no marked 
contrast. Another skin, also in summer pelage, is a uniform 
dark slaty drab throughout, the hind feet darker and more 
brown ; sides and underparts with a few long white hairs 
(50 mm.) ; lips with slight grizzling due to presence of short 
white hairs. 

Measurements. — For cranial and dental measurements see 
Tables, pp. 322, 323. 



VULPES 



325 



Specimens examined. — Six, all from Spitzbergen(B.M.and U.S.N.M.): — 

i skull. Spitzbergen. Stockholm Museum (e). 90. 8. 1. 2. 

6, 9. Spitzbergen. Dr. J. W. Gregory (c & p). 96. 9. 23. 2-3. 

(96. 9. 23. 3. Type of species.) 
S, ¥ skulls. Spitzbergen. Dr. J. W. Gregory (c & p). 96. 9. 23. 4-5. 

Genus VULPES Oken. 

1816. V[ulpes] Oken, Lehrb. d. Naturgesch., in, pt. 2, p. 1033, in full on 
p. 1034 and in index, p. 1268 (Vulpes communis Oken = Canis 
vulpes Linnseus). 

1821. Vulpes Bowdich, Anal. Nat. Olassif. Mamm., p. 40 (Canis vulpes). 

1857. Vulpes Blasius, Saugethiere Deutschlands, p. 178 (Sub-genus of 
Canis). 

Type species. — Canis vulpes Linnseus. 

Geographical distribution. — Northern portion of the northern 
hemisphere from about the limit of tree growth south to Morocco, 
India and Mexico ; in Europe west to Ireland. 

Characters. — Skull slender and low (depth of brain-case less 
than one- third condylobasal length) ; interorbital region nearly 
flat, the frontal sinuses scarcely inflated, the postorbital processes 
thin, slightly concave above, their edges overhanging and bead- 
like ; dorsal profile of forehead rising very slightly and gradually 
above level of rostrum ; dental formula as in Canis ; teeth 
relatively light and small, the length of upper carnassial and 
molars together contained about 2| to 3 times in palatal length, 
the general character of cheek-teeth somewhat more trenchant 
than in Canis, the canines slender and elongated, the point of 
upper tooth extending to about level of lower margin of mandi- 
bular ramus when jaws are closed (fig. 65). 

Remarks. — As thus restricted the genus Vulpes contains about 
thirty-five forms, all peculiar to the northern hemisphere. Five 
of these occur in Europe. 

KEY TO THE EUROPEAN FORMS OP VULPES. 

Size small, hind foot in adult male about 125 mm., 
condylobasal length of skull in both sexes less 

than 130 mm. (Sardinia and Corsica) V. ichnusse, p. 336. 

Size large, hind foot in adult male 135 to 165 mm., 
condylobasal length of skull in adult male 135 to 

165 mm., in adult female 127 to 155 mm V. vulpes, p. 326. 

Teeth larger and more robust, the premolars tend- 
ing to be slightly spaced or in contact, their 
secondary cusps well developed (Scandinavia)... V. v. wipes, p. 330. 
Teeth smaller and less robust, the premolars tend- 
ing to be widely spaced, their secondary cusps 
usually obsolete or absent. 
Yellowish and reddish tints bright ; posterior 
half of back seldom much frosted with whitish ; 

tail never clear grey (Central Europe) V. v. crurigera, p. 331. 

Yellowish and reddish tints pale and dull; pos- 
terior half of back usually much frosted with 
whitish ; tail often clear grey (Iberian Penin- 
sula) V. v. silacea, p. 333. 



326 



CARNIVOEA 




FIG. 65. 
Incisors and canines from front, of Cards (a), Alopex (6), and Vulpes (c). 



VULPES VULPES Linnaeus. 

(Synonymy under subspecies.) 

Type locality. — TTpsala, Sweden. 

Geographical distribution. — Europe from the Arctic coast to 
the Mediterranean, and from Ireland eastward into Asia. 

Diagnosis. — Size large: hind foot, in adult male 135 to 165 
mm.; condylobasal length of skull in adult male 135 to 165 mm., 
in adult female 127 to 155 mm. 

External characters. — Form more slender and legs relatively 
shorter than in Canis ; muzzle long and pointed ; ear high, 
pointed, rising conspicuously above surrounding fur ; tail long, 



v0lpes 327 

thick and bushy, with abundant underfur ; longer hairs of back 
normally concealing the underfur ; feet as in Ganis, but with 
the soles hairy between the pads, the pads themselves sometimes 
furred. 

Colour. — General colour *a yellowish brown brighter and more 
inclined toward reddish along median dorsal region and on face, 
duller and more yellowish or greyish on sides of body to shoulder 
and on sides of neck to base of ear, the flanks and usually the 
sides sprinkled with white hairs which may produce a decided 
effect of frosting ; underparts very variable, ranging from 
whitish to slaty black, rarely almost concolor with sides ; feet 
dusky or blackish ; ear tawny or buff at base and on inner 
surface, the terminal half of outer side black or very dark brown 
in strong contrast with surrounding parts ; upper lip dull white. 
Blackish and greyish variations not uncommon, especially at the 
north. 

Skull. — General form of skull slender and somewhat flattened, 
with widely spreading, nearly parallel zygomata. Dorsal profile 
almost flat from nares to slightly beyond middle of nasal bones, 
then rising at a slight angle (10° or less) to or a little beyond 
bregma, behind which it slopes away by an evenly convex curve 
(distorted in old individuals by the development of the sagittal 
crest) to lambda, which lies a little above level of middle of orbit ; 
ventral profile essentially straight. Brain-case distinctly broader 
than high, its outline ovate when viewed from above, the 
lambdoid and sagittal crests well developed, the latter extending 
forward about to bregma, where it divides, sending a branch to 
form posterior border of each postorbital process. Occiput 
obliquely truncate, so that condyles are not visible from above, 
but region between lambda and foramen magnum slightly if at 
all concave. Floor of brain-case with no special peculiarities ; 
auditory bullae moderately and evenly inflated, slightly flattened 
antero-externally, meatal tube short but distinctly indicated, 
especially its hinder wall. Interorbital region flattened, with 
median longitudinal groove, the postorbital processes prominent, 
flattened, triangular, much shorter along posterior edge than 
along antero-external edge, the margin slightly raised so that 
the upper surface is somewhat concave, the under surface so 
abruptly concave that the process is much less thick than in 
Alopex and Ganis. Rostrum moderately long (the distance from 
orbit to gnathion about equal to that from postorbital process to 
lambda), rather abruptly narrowed proximally, so that the sides 
are nearly parallel through a noticeable portion of their extent 
(occasionally the sides diverge from region of greatest narrowing 
to bases of canines) ; nasal slender, narrowing gradually back- 
ward and extending nearly to level of middle of orbit ; nasal 
branch of premaxillary extending to about middle of nasal and 
usually not in contact with frontal ; posterior extremity of 
maxillary extending slightly behind that of nasal ; anteorbital 



328 



CARNIVORA 



foramen over posterior root of pm 3 or over space between pm 3 
and pm i . Palate relatively narrower than in the European 






Fig. 66. 
Vulpes vulpes. X h 



species of Alopex and Canis, its median posterior termination 
about at level of middle of last molar ; mesopterygoid fossa 



329 



about twice as long as wide, the sides gradually converging 
posteriorly ; hamulars slender, straight ; incisive foramina about 
four times as long as wide, the median septum usually wider 
than the foramen, the posterior margin a little in front of 
posterior margin of alveolus of canine. Mandible slender, but 
with no special peculiarities of form. 

Teeth. — In general form and structure the teeth do not differ 
very widely from those of Canis lupus. The canines are, however, 
relatively much longer and more slender (fig. 65), the incisors are 
weaker and narrower, and the premolars show a more decided 
tendency to develop secondary 



S 9 




cusps. Upper incisors slender, 
not closely crowded, their 
crowns relatively narrower than 
in either Canis lupus or Alopex 
lagopus, the secondary cusps 
obsolete, early disappearing 
with wear, the cingulum barely 
indicated ; lower incisors with 
crowns less simplified than in 
the upper teeth, the usual lon- 
gitudinal groove present on 
posterior surface, and i 3 with 
well developed outer basal lobe. 
Upper canine slender, its dia- 
meter at alveolus about 7 mm., 
its height when unworn usually 
about three times as great ; 
lower canine with diameter 
contained about 2£ times in 
height. Premolars with no 
special peculiarities, their crowns 
relatively narrower than in 
Canis lupus, and secondary cusps 
tending to be more developed, 
the latter character varying in 
different geographical forms. 
Upper carnassial with inner 
lobe better developed than in 
Canis, and bearing a distinct terete cusp, its position a little 
more forward and outward than in the dogs, so that it appears 
to he in or nearly in the main axis of the tooth. Lower carnassial 
with posterior heel essentially as broad as main portion of tooth, 
the cusps essentially as in Canis, but general aspect of tooth more 
trenchant. Upper molars differing from those of Canis in the 
relatively smaller size of paracone and metacone, these cusps 
appearing to stand in from border of crown, leaving a noticeable 
cingulum beyond them ; paraconule obsolete. Second and third 
lower molars with no marked peculiarities. 







Fig. 67. 
Vulpes vulpes. Teeth. 



Nat. size. 



330 



CABNIVOEA 



Remarks. — With the material at hand it has been impossible 
to reach wholly satisfactory conclusions with regard to the 
number of local forms represented by the common foxes of various 
parts of Europe. The existence of three such races, one in the 
Scandinavian Peninsula, a second in Central Europe, and a third 
in the Iberian Peninsula, seems well established ; but the status 
of the forms inhabiting Italy and Greece is still in doubt. 



Vulpes vulpes vdlpes Linnseus. 

1758. [Canis] vulpes Linnseus, Syst. Nat., i, 10th ed., p. 40 (Sweden). 
1758. [Canis] alopex Linnseus, Syst. Nat., I, 10th ed., p. 40 (Sweden). 
1798. C[anis] vulpus Thunberg, Beskrifning pa Svenske Djur, Mamm., p. 7 

(Variant of vulpes). 
1816. V[ulpes] vulgaris Oken, Lehrb. d. Naturgesoh., in, pt. 2, p. 1034 

(Renaming of vulpes). 
1820. Canis nigro-argenteus Nilsson, Skand. Fauna, i, p. 91 (Lofoten 

Islands, Norway). 
1827. [Canis vulpus] y nigrocaudatus Billberg, Synopsis Faunse Scandi- 

navise, p. 12 (Uppland, Sweden). 
1827. [Canis vulpus] £ variegatus Billberg, Synopsis Faunse Soandinavise, 

p. 13 (Uppland, Sweden). 
1827. [Canis vulpus] 17 Uneatus Billberg, Synopsis Faunse Soandinavise, 

p. 13 (Skane, Sweden). 
1830. ? [Vulpes] communis Burnett, Quart. Journ. Sci. Lit. Art, xxviii, 

1829, p. 349 (Substitute for vulpes). Nomen nudum. 
1898. Vulpes vulpes Thomas, The Zoologist, 4th ser., n, p. 100, March, 

1898 (part). 
1910. Vulpes vulpes Trouessart, Faune Mamm. d'Europe, p. 93 (part). 

Type locality. — Upsala, Sweden. 

Geographical distribution. — Scandinavian Peninsula. 

Diagnosis. — Teeth larger and more robust than in the central 
and southern races, the premolars tending to be slightly spaced 
or in contact ; skull attaining maximum size for European 
foxes. 

Measurements. — For cranial and dental measurements see 
Tables, pp. 334, 335. 

Specimens examined. — Seven skulls from Sweden (for exact localities 
see Table of cranial measurements), and eleven from Egersund, Stavanger, 
Norway. 

Remarks. — While I have seen no skins of this fox, the 
distinctness of the race from those occurring in central and 
southern Europe seems well established by the characters of 
the teeth. 

skull. Sweden. Wheelwright (c). 64. 3. 8. 2. 

11 skulls. Egersund, Stavanger, K. H. Schaanning (c). 11. 6. 3. 1-11. 
Norway. 



VULPES 331 

Vulpes vulpbs crucigera Bechstein. 

1789. [Cams] crucigera Bechstein, Gemeinn. Naturgesoh. Deutsohlands, 
I, p. 250 (Thiiringen, Germany). 

1792. C[anis] Vulpes alopex europaeus Kerr, Anim. Kingd., p. 142 (Bur- 
gundy, France). 

1797. Cams vulpes alba Borkhausen, Deutsche Fauna, I, p. 33 (Vogelsberg, 
near Budigshain, Hessen, Germany). 

1797. Cams vulpes nigra Borkhausen, Deutsche Fauna, I, p. 33 (Hessen 
and Thiiringen, Germany). 

1801. Clams'] v[ulpes] lutea Bechstein, Gemeinn. Naturgesch. Deutsoh- 
lands, i, 2nd ed., p. 628 (Thiiringen, Germany). 

1801. C[anis] v[ulpes] cinerea Bechstein, Gemeinn. Naturgesch. Deutsoh- 
lands, I, 2nd ed., p. 628 (Thiiringen, Germany). 

1832. Canis melanogaster Bonaparte, Ioonogr. Fauna Ital., i, faso. 1 
(Neighbourhood of Borne, Italy). 

1841. Vulpes hypomelas Wagner, Schreber's Saugthiere, Suppl., n, p. 405 
(Oberbayern, Germany). 

1857. Cams vulpes Blasius, Saugethiere Deutsohlands, p. 191. 

1861. ? V[ulpes] vulgaris meridionatis Fitzinger, Wissensch.-pop. Natur- 
gesch. der Saugeth., I, p. 194 (Dalmatia). 

1910. Vulpes vulpes (part) and V. vulpes melanogaster Trouessart, Faune 
Mamm. d'Europe, pp. 93-94. 

Type locality. — Thiiringen, Germany. 

Geographical distribution. — Central and southern Europe from 
Ireland eastward and from the coast of the Baltic to the 
Pyrenees, Italy and Greece. 

Diagnosis. — Maximum size rather less than in V. v. vulpes, 
and teeth distinctly smaller, the premolars rather widely spaced 
and seldom if ever in contact ; general colour a bright yellowish 
or reddish brown, the posterior half of back not conspicuously 
frosted with whitish, and tail never clear greyish. 

Colour. — In seventeen skins the general colour ranges from 
nearly cinnamon-rufous to a light ochraceous-rufous, the sides 
of neck and region immediately behind shoulder lighter than 
median dorsal area (in extreme instances clear buff with a 
decided rufous tinge) ; posterior half of back with evident 
white frosting in some specimens, scarcely any in others, but 
this character never so pronounced as in average Spanish skins ; 
underparts dull slaty overlaid with white, the slaty nearly always 
predominating, except on throat, and not infrequently giving the 
effect of an almost blackish tinge throughout, this apparently 
not in the least dependent on regional or local climatic con- 
ditions. In a flat skin from Cephalonia, Greece, the characteristic 
slaty and white is confined to the throat and chin, all the rest 
of the ventral region being a dull tawny-ochraceous like sides. 
Two specimens from Tatoi, near Athens, taken in July, have 
shed all the longer hairs of the back, leaving only the velvety 
underfur. This is of the usual colour, a dull umber brown, in 
one specimen with a slaty cast. 

Skull and teeth. — The skull and teeth do not attain so great 



332 



CARNIVOKA 



size as in true Vulpes vulpes, and the small premolars are seldom 
in contact, the spaces between them usually conspicuous. 

Measurements. — Adult female from Cappagh House, Water- 
ford, Ireland : head and body, 613 ; tail, 340 ; hind foot, 146 ; 
ear from meatus, 90. Adult male and female from Tunbridge 
Wells, Sussex, England : head and body, 690 and 630 ; tail, 
343 and 370 ; hind foot, 160 and 141 ; ear from meatus, 95 and 
82. Adult male from Ingelheim, Rheinhessen, Germany : head 
and body, 578 ; tail, 440 ; hind foot, 160 ; ear from meatus, 98. 
Adult male and female from Haute-Garonne, France : head and 
body, 610 and 610; tail, 370 and 350; hind foot, 150 and 135 ; 
ear from meatus, 94 and 89. Two adult males from Porlezza, 
Como, Italy (Ghidini) : hind foot, 150 and 148-6. Adult male 
from Borghetto S. Spirito, Italy : head and body, 745 ; tail, 380 ; 
hind foot, 157. Adult male from Zinnigas, Siliqua, Sardinia 
(measured from mounted specimen, Genoa) : head and body, 700 ; 
tail, 300 ; hind foot, 144± ; ear from meatus, 83. 

Specimens examined. — Fifty, from the following localities : — 

Ireland : Kilmanock, Wexford, 2 ; Cappagh House, Waterford, 1. 

Scotland : Ben Nie, Sutherland, 1 ; Inversanda, Ardgour, 1. 

England : Northumberland, 1 ; Thame, Oxford, 2 ; Hassocks, Sussex, 1 ; 
Tunbridge Wells, Sussex, 3; Ditchling, Sussex, 1; Mayfield, Sussex, 1 
(Grant). 

France : Ax-les-Thermes, Ariege, 3 ; Caterille, Haute-Garonne, 1 ; Pic 
Sessire, Haute-Garonne, 1 ; St. Aventin, Haute-Garonne, 1. 

Germany : Ingelheim, Rheinhessen, 1 ; Nuremberg, Bavaria, 1 
(U.S.N.M.) ; Grossgraben, Silesia, 1 (Breslau) ; Biesengiberge, Silesia, 1 
(Breslau) ; southern Germany, 3. 

Austria-Hungary : Trentino, 1 (Genoa) ; Gazza, Trentino, 1 (Genoa) ; 
Vigolo Vattaro, Trentino, 1 (Genoa). 

Switzerland : Geneva, 1 (Ghidini) ; Valais, 1 (Ghidini). 

Italy : Porlezza, Como, i (Ghidini) ; Garbagna, Piedmont, 1 (Genoa) ; 
Borghetto S. Spirito, 1 (Genoa) ; Vargo, Liguria, 1 (Genoa) ; Torriglia, 
Liguria, 1 (Genoa) ; Cornigliano, Liguria, 2 (Genoa) ; near Genoa, 1 ; 
Molasana, 1 (Genoa) ; Tuscany, 1 ; Pisa, 1. 

Sardinia : Zinnigas, Siliqua, 1 (Genoa). 

Greece : Cephalonia, 1 ; Tatoi, near Athens, 2. 

Remarks. — The fox of Italy appears to be the same as that of 
Central Europe, though further material from the southern 
portion of the peninsula may show that it should be distinguished. 
The three specimens from Greece are in such unsatisfactory 
condition of pelage that their status cannot be determined with 
any degree of certainty. If they represent a peculiar local race 
this should probably take the name meridionalis Fitzinger. 

9. Ben Nie, Sutherlandshire, E. B. Alston (p). 79. 9. 25. 80. 

Scotland. (H. Brown.) 

6. Inversanda, Ardgour. H. Leigh (c & p). 1. 16. 12. 1. 

2. Kilmanock, Wexford, G. Barrett-Hamilton 6. 5. 20. 1. 

Ireland. (c & p). 9. 12. 15. i. 

?. Cappagh House, Waterford. R. J. Ussher (c & p). 96. 12. 28. 1. 

1. Northumberland, England. Rev. H. H. Slater 0. 2. 24. 5. 

(c & p). 



volpbs 333 

2 6. Thame, Oxfordshire. Hon. N. C. Roths- 0. 10. 31. 1-2. 

child (p). 

2i5. Ditchling, Sussex. Guy Do 11m an (9. 11. 3. 1. 



child (p). 

Dollman (9. 11. 
(c & p). \10. 9. 13. 1. 



3. Tunbridge Wells, Sussex. C. H. B. Grant (c). 1. 2. 15. 1-3. 

2 6,9. Ax-les-Thermes, Ariege, V. Builles (a & p). 8.3.27.6-7. 

Prance. 8. 3. 27. 14. 

9. Caterille, Haute-Garonne, 0. Thomas (p). 8. 7. 15. 2. 

900 m. (A. Robert.) 

9. Pic Sessire, Haute-Garonne. O. Thomas (p). 8. 7. 15. 3. 

(A. Robert.) 

6. St. Aventin, Haute-Ga- 0. Thomas (p). 8. 7. 15. 4. 

ronne, 900 m. (A. Robert.) 

6. Ingelheim, Rheinhessen, C. Hilgert (c). 8. 11. 2. 15. 
Germany. 

3 skulls. South Germany. Dr. A. Giinther (c). 59. 9. 6. 83, 88. 

175. k. 

6. Genoa, Liguria, Italy. Marquis G. Doria(p). 88. 12. 1. 3. 

skull. Pisa. Zool. Soc. Mus. 58. 5. 4. 126. 

1 yg. S. Italy. (Prof. Savi.) Zool. Soc. Mus. 55. 12. 24. 240. 

9. Tuscany. Purchased (Dr. 45.7.22.15. 

Riippell). 

6, 9. Tatoi, Athens, Greece. Hon. N. 0. Roths- 8. 10. 2. 22-23. 
(C. Mottaz.) child (p). 

Vulpbs VULPES silacea Miller. 

1907. Vulpes vulpes silaceus Miller, Ann. and Mag. Nat. Hist., 7th ser., 

xx, p. 393, November, 1907. Type in British Museum. 
1910. Vulpes vulpes silaceus Trouessart, Faune Mamm. d'Burope, p. 45. 

Type locality. — Near Silos, Province of Burgos, Spain. 

Geographical distribution — Iberian Peninsula. 

Characters. — Size about equal to that of V. v. vulpes, but 
teeth noticeably less enlarged, the wider spacing of the premolars 
especially evident ; general colour a dull buff without bright 
yellowish or reddish tints, the posterior half of back conspicuously 
frosted with whitish (except in abraded pelage) the tail often a 
clear greyish. 

Measurements. — Type (adult male) : head and body, 750 ; 
tail, 370 ; hind foot, 150. Adult female from near the type 
locality : head and body, 670 ; tail, 330 ; hind foot, 125. Old 
male from Elche, Alicante: head and body, 770; tail, 480; 
hind foot, 160; ear from meatus, 113. 

Specimens examined. — Eighteen, from the following localities in 
Spain : Olot, Gerona, 1 (probably not typical) ; vicinity of Silos, Burgos, 5 ■ 
near Burgos, 1; Arrechavaleta, Vitoria, 1; Torres del Alio, Coruna, 1 
Madrid, 1 ; near Seville, 3 ; Coto Donana, Huelva, 3 (B.M. and U.S.N.M.) 
Elche, Alioante, 1 ; Barracas, Castellon, 1. 

Remarks. — The Spanish fox is well characterized by its light 
colour, large size and rather small, widely-spaced premolars as 
compared with those of the Scandinavian form. 

2 6. Silos, Burgos, Spain. G. S. Miller (c). 8. 8. 4. 46-47. 

(8. 8. 4. 46. Type of subspecies.) 
6, 2 9, 1. Burgos. Rev. Saturio Gon- 8. 7. 7. 12-15. 

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CARNIVORA 



9. Arrechavaleta, Vitoria. 0. Thomas (p). 8. 2. 9. 49. 

(N. Gonzalez.) 

S. Torres del Alio, Corufia. Dr. V. L. Seoane 94. 5. 29. 1. 

(o & p). 

5. Madrid. M. de la Esoalera (c). 8. 9. 24. 7. 
<$, ?. Seville. (A. Buiz.) Lord Lilford (p). 95. 9. 4. 7-8. 
2 c5. Coto Donana, Huelva. A. Chapman (c & p). 8. 3. 8. 4-5. 

6. Elehe, Alicante. G. S. Miller (c). 8. 8. 4. 48. 
i juv. Barraoas, Castellon. 0. Thomas (p). 8. 2. 9. 50. 

(N. Gonzalez.) 



VULPES ICHNUS.E Miller. 

1907. Vulpes ichnusee Miller, Ann. and Mag. Nat. Hist., 7th ser., xx, p. 391, 

November, 1907. Type in British Museum. 
1910. Vulpes vulpes ichnusse Trouessart, Paune Mamm. d'Burope, p. 94. 

Type locality. — Sarrabus, Sardinia. 

Geographical distribution. — Sardinia and Corsica. 

Diagnosis. — Smaller than any of the races of Vulpes vulpes ; 
both hind foot and condylobasal length of skull in adult male 
less than 130 mm., ear from crown 60 to 70 mm. 

Colour. — Face and head dark rufous becoming lighter and 
more dull on base of ears and on neck, and fading to ochraceous- 
ruf ous on shoulders and back ; sides of neck, outer surface of 
upper arm and region just behind axilla still lighter, a tawny 
buff, hairs of underfur on back drab grey at base, tawny clay 
colour at tip ; longer hairs of head, sides, and back (behind 
shoulders) much speckled by buffy white subterminal areas about 
5 mm. in length, the extreme tips reddish ; feet and legs 
ochraceous-rufous, slightly clouded with blackish and a little 
speckled with buffy white ; tail like back above, the tawny 
gradually fading out through a buffy grey to the whitish buff 
tip, the longer hairs except in pencil black tipped (30-40 mm.) ; 
underparts to front legs buffy whitish tinged with hair-brown, 
this nearly clear on middle of throat ; rest of underparts a 
mixture of hair-brown and dull tawny, the latter predominating 
along sides. 

Measurements. — Type (adult male) : hind foot, 123 j ear from 
meatus, 70. Adult male and female from the type locality : 
head and body, 640 and 590 ■ tail, 280 and 290 ; hind foot, 
127 and 125 ; ear from meatus, 74 and 71. Adult female from 
Siliqua : head and body, 600 ; tail, 350 ; hind foot, 1 27 ; ear, 74. 
Adult from near St. Florent, Corsica; hind foot, 123 ; ear from 
meatus, 73. 

Specimens examined. — Eight, from the following localities : — 
Corsica : Grotto Campu Cousule, near St. Florent, N.W. Corsica, 1 

(Major). 

Sardinia : No exact locality, 1 (Turin) ; Lamisei, 1 ; Sarrabus, 3 (B.M. 

and Genoa) ; Zinnigas Siliqua, 2. 



337 



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Lanusei .... 
Corsica : near St. Plorent 


Germany : Southern Germany 

Ingelheim, Rheinh 
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Italy : near Genoa . 
Pisa 
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Greece : Tatoi, near Athens 
ii ii 


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340 



CARNIVORA 



Bemarks. — This small, short-eared fox is probably a native 
on the two islands where it occurs. The presence of the larger 
animal, similar to that of Italy, on Sardinia may be due to recent 
introduction. The Corsican specimen, though a mummy and 
without fur, appears to be a typical example of Vulpes ichnusse. 

6. Sarrabus, Sardinia. Marquis G. Doria (p). 88. 12. 1. 2. 

(Type of species.) 
6. Lanusei, Sardinia. 0. Thomas (p). 0. 2. 21. 1. 

(W. Wolterstorff.) 

Family MUSTELIDvE. 
835. Mustelidss Swainson, Nat. Hist, and Classif. of Quadrupeds, p. vn, 361. 

Geographical distribution. — Essentially cosmopolitan ; absent 
from Madagascar and Australia ; in Europe west to Ireland. 

Characters. — Larger cheek-teeth of a combined trenchant and 
crushing type, the last upper premolar and first lower molar 
strongly differentiated as carnassials, the former 3-rooted, its 
inner lobe in front of middle of crown ; upper molars, 1-1 ; upper 
carnassial with not more than two outer cusps ; auditory bulla flat 
or moderately inflated, without septum ; form usually slender, 
the legs always short ; size moderate or small (including the 
smallest known carnivores) ; feet digitigrade or sub- plantigrade ; 
toes. 5-5. 

Bemarks. — The family Mustelidse is, next to the Ganidte, the 
most generally distributed group of carnivores. It is divisible 
into four sub-families, all of which occur in Europe, where they 
are represented by six of the two dozen or more known genera. 

KEY TO THE EUROPEAN GENERA OP MTJSTELIDM. 

Grown of upper carnassial triangular or rhombic in 
outline, its length and width sub-equal ; lower car- 
nassial with anterior triangle distinct, the mefca- 
conid nearly as large as the outer cusps. 
Upper molar much larger than carnassial ; skull 
narrow and high (normal), the rostrum much 
longer than broad; tail short, bushy, not mus- 
cular ; habits fossorial (Badgers, sub-family 



Meles, p. 341. 



Upper molar about equal to carnassial ; skull broad 
and flat, the rostrum broader than long ; tail long, 
densely furred, very muscular ; habits aquatic 

(Otters, sub-family Lutrime) Lutra, p. 354. 

Crown of upper carnassial not triangular or rhombic in 
outline, its length much greater than its width ; 
lower carnassial with anterior triangle obsolete or 
absent (represented by the two outer cusps only), 
the metaconid when present much smaller than the 
other cusps. 



341 



Premolars t-.. 

4-4 



Lower carnassial without metaconid ; skull robust 
(in European species considerably more than 
100 mm. in condylobasal length) ; form robust ; 
tail short (Wolverenes, sub-family Ouloninm)... Oulo, p. 433. 

Lower carnassial with evident though small meta- 
conid ; skull slender (in European species con- 
siderably less than 100 mm. in condylobasal 
length) ; form slender ; tail long (Martens, sub- 
family Mustelinie) Maries, p. 365. 

Premolars ||. 

Lower carnassial with evident though small meta- 
conid ; hamular in contact with bulla ; back and 
sides spotted and striped (Tiger Polecats, sub- 
family Mustelinae) Vormela, p. 428. 

Lower carnassial without metaconid ; tip of hamu- 
lar widely separated from bulla ; back and sides 
never spotted, rarely (in certain Asiatic species) 
with median dorsal stripe (Weasels, Polecats, 
&c, sub-family Mustelime) Mustela, p. 381. 

Sub-Family MELINiE. 

1857. Melinss Baird, Mamm. North Amer., p. 148. 

Geographical distribution. — Temperate and tropical portions of 
both hemispheres ; in Europe west to Ireland and north to 
central Scandinavia. 

Characters. — Upper carnassial with evident crushing surface, 
its crown triangular or rhombic in outline ; upper molar large, 
the length of its outer portion usually equal to or greater than 
that of carnassial ; skull rather high and long, the rostrum 
longer than broad ; external form short and heavy, the fur long 
and loose ; toes not webbed, the claws large, fossorial ; tail 
variable in length (short in European members of the group), 
never unusually muscular. 

Bemarks. — About a dozen genera, or nearly one-half of the 
family, are now placed in the sub-family Melinse. Only one 
occurs in Europe. 

Genus MELES Brisson. 

1762. Meles Brisson, Kegn. Anim. in Classis ix, distrib., 2nd ed., p. 13 

(Meles Brisson = Ursus meles Linneeus). 
1780. Meles Storr, Prodr. Meth. Mamm., p. 34. First use of name by an 

author following the Linnsean system ( Ursus meles) . 
1795. Taxus Geoffroy and Cuvier, Magasin Encyclopedique, II, p. 184 

(Ursus meles Linnaeus). 
1815. Melesium Rafmesque, Analyse de la Nature, p. 59 (Modification of 



1857. Meles Blasius, Saugethiere Deutschlands, p. 202. 

Type species. — Meles * Brisson = Ursus meles Linnseus. 

* Not "Metes meles," the form in which this and other Brissonian 
monomial specific names are often cited, apparently with the intention of 
palliating the absurdity of recognising in nomenclature the terms applied 
to genera by an author who did not follow the Linnsean system. 



342 



CARNIVORA 



Geographical distribution. — Northern temperate portions of 
Old World ; in Europe from central Scandinavia to the 
Mediterranean, and from Ireland eastward. 

Characters. — Skull narrow and high (depth of brain-case 
much more than half mastoid breadth), the zygomatic arches not 
widely expanded, and postorbital region not specially narrowed 
(distance from point of greatest narrowing to zygoma less than 
breadth of postorbital constriction) ; rostrum elongate, the 
distance from orbit to gnathion about three-fourths length of 
brain-case ; auditory bulla elevated along inner margin, but else- 
where somewhat flattened, the meatal tube distinct ; paroccipital 
process robust, standing out conspicuously behind bulla ; dental 
formula: ijgj, eg, pm^* mt? = 38; teeth not specially enlarged 
as compared with width of rostrum and palate (greatest trans- 
verse diameter of upper carnassial equal to a little more than 
one-third distance between carnassials) ; three small premolars 
(one upper and two lower) capable of trenchant action ; upper 
carnassial with crown triangular in outline, the cutting portion 
consisting of a single large anterior cusp with its slightly developed 
anterior commissure and more distinct posterior commissure, the 
crushing portion represented by the obliquely sloping inner base 
of this cusp and ridge ; upper molar rhombic in outline, its crown 
area about three times that of carnassial, its greatest diameter 
in axis of tooth- row, the four primitive cusps present ; lower 
carnassial with the three anterior cusps well developed, sub-equal, 
the posterior heel decidedly larger than anterior triangle ; 
external form heavy, the head pointed, the ears short but plainly 
visible, the body thick, the tail short, not muscular ; feet sub- 
plantigrade, the toes with long fossorial claws and without webs ; 
fur coarse and loose. 

Memarhs. — The genus Meles contains about half a dozen 
named forms, the status of several of which is at present not 
clearly understood. Two species occur in Europe. 

KEY TO THE EUROPEAN POEMS OP MELES. 

Maxillary tooth-row, exclusive of incisors, about 
85 mm. ; auditory bullee rather strongly in- 
flated, the inner border not ridge-like (Crete) M. arcalus, p. 352. 
Maxillary tooth-row, exclusive of incisors, about 
40 mm. ; auditory bullae slightly inflated, the 
inner border ridge-like (distribution general) M. meles, p. 343. 
General colour moderately pale ; teeth averaging 
smaller, less frequently attaining maximum 

size (Central and southern Europe) M. m. meles, p. 348. 

General colour slightly paler ; teeth averaging 
larger, and more frequently attaining maxi- 
mum size (Iberian Peninsula) M. in. marianensis, p. 352. 



* In adults usually — ,, owing to the early disappearance of the small 
pm l . 



343 



meles MELES Linnaeus. 
(Synonymy under subspecies.) 

Type locality. — Upsala, Sweden. 

Geographical distribution. — Europe, west to Ireland, south to 
the Mediterranean and north to central Scandinavia. Eastern 
limits of range not known. 

Diagnosis. — Size large (upper length of skull in adult more 
than 120 mm. ; maxillary tooth-row, exclusive of incisors, about 
40 mm. or more ; hind foot about 90 to 110 mm.) ; auditory bulla 
moderately inflated, the highest region close to inner margin and 
forming an evident longitudinal ridge, the region between ridge 
and meatus noticeably concave ; teeth large, the two lower molars 
together 22 mm. or more in length ; postero-external border of 
upper molar usually convex, though occasionally straight or 
slightly concave. 

External characters.- — Pelage coarse and loose, practically 
without underfur, the hairs at middle of back about 60 mm. in 
length, those on sides longer ; underparts scantily haired, the 
skin usually visible ; palm bare, usually with a slight bristly 
pubescence at middle in area between tubercles ; a large 
tubercular mass at base of digits, convex in front, concave behind, 
wider on outer border than on inner border, showing no tendency 
to trilobation ; a roundish tubercle about half as large at postero- 
external border of palm, separated from anterior mass by a wide 
space ; a small, ill-defined pad at base of thumb ; sole densely 
haired from heel to a little beyond middle, then completely naked ; 
plantar tubercles essentially like those on palm except that 
small pad at base of hallux is absent, and the two large masses 
tend to coalesce, owing to absence of the wide intervening space ; 
surface of pads on both palms and sole finely rugose, this 
especially noticeable in dried specimens ; muzzle pad entirely 
naked, but separated from upper lip by a narrow hairy band. 
Mamma; : a 2-2, i 1-1 = 6. 

Colour. — Back and sides a coarse grizzle of black and bufly 
■white, the black usually predominating on back, the lighter 
colour on sides. Throat, median ventral area and all four legs 
and feet black or blackish ; face, chin and entire neck (except 
underside) clear whitish except for a broad dark brown or black 
band beginning on each side about 15 mm. behind nostril pad 
and extending back, including eye and ear, to middle of neck, 
where it fades insensibly into colour of back ; width of white 
median area on face usually greater than that of the dark lateral 
stripe, and about equal to that of the light area between lateral 
stripe and dark ventral area ; ear black, its anterior border 
white in strong contrast ; eye usually a little below middle of 
dark band ; tail like back at base, soon fading to soiled white. 

Skull. — Except for the greatly developed sagittal crest, the 



344 



CAENIVOKA 



height of which in old individuals is equal to nearly half inter- 
orbital breadth, the skull of Meles nieles is rather smooth and 
without marked angularity. Depth of brain-case (without crest) 
and depth through posterior portion of orbit nearly equal, and 
about two-thirds mastoid breadth • depth at front of nasal about 
two-thirds that at orbit. Dorsal profile (without crest) nearly 
flat from middle of brain-case to interorbital region, convex over 
posterior half of brain-case ; rostrum sloping downward at angle 
of about 30° with surface of brain-case. When crest is fully 




Fig. 88. 
Meles meles. X 4. 



developed it reduces this angle to 20° or less. It is slightly 
convex anteriorly, rather abruptly convex posteriorly, the hinder- 
most portion slightly overhanging. Postorbital processes short 
but well developed, sharply outlining the small orbit (greatest 
diameter of orbit slightly less than half that of interorbital 
region) which, though widely open posteriorly, is more than half 
encircled with bone. Rostrum moderately long, the distance 
from gnathion to front of zygoma about equal to width of palate 
including molars, and continued about 3£ times in condylobasal 
length of skull. Zygomata widely spreading posteriorly, narrow 



MELES 



345 



anteriorly, the widest portion opposite front of glenoid fossa, the 
arch very slightly bowed upward. Anteorbital foramina rather 
large, sub-circular in outline, their greatest diameter about half 
that of orbit. Ventral profile of skull slightly concave through- 
out. Palate moderately wide, the distance between molars 
contained about 2 J times in distance from gnathion to level of 
posterior edge of molar. Incisive foramina small, at level of 
space between canine and i 3 , elongate pyriform in outline, the 
minute median foramen a little behind middle. Posterior exten- 
sion of palate about equal to distance between molars, and slightly 
more than half that from molar to hamular. Mesopterygoid 




x h 



space short and wide, its length only a little more than distance 
between tips of hamulars. Auditory bull* slightly inflated, 
irregularly triangular in outline, the meatus lying in angle formed 
by the large, forward-projecting mastoid process. The surface is 
irregular, with evident ridge near inner margin along crest of 
most highly inflated region, this ridge often terminating anteriorly 
in a bluntly pointed projection. Length of flattened portion 
extending inward from meatus about equal to transverse diameter 
of inflated portion of bulla, but the two regions not sharply 
defined ; least distance between bullae about 1£ times diameter of 
inflated portion ; paroccipital process short, triangular (not ridge- 
like), its extreme base applied to posterior border of bulla. 



346 



CAENIVOKA 



Mandible heavily built, the ramus nearly straight, the lower 
border with the usual upward curve posteriorly, its depth at 
middle about one-third that through coronoid process ; angular 




fig. 70. 

Meles males. X f. 

process short, thick, and ill-defined, lying close to base of articular 
process ; coronoid process squarely truncate above, its height 
above articular process about equal to width at articular level. 

Teeth. — The teeth are moderately large relatively to size of 
skull, the incisors and canines rather short, the crown area of the 
upper molar fully double that of any of the other maxillary teeth, 
this last peculiarity unique among the European Mustelidse. Upper 
incisors robust, the crowns somewhat higher than wide when 
viewed from in front, the teeth closely crowded in a slightly 
convex row, the longitudinal diameter of crown a little greater 
than transverse diameter ; i 1 and P sub-equal, the latter slightly 
the larger, their anterior surface convex with two faint longi- 
tudinal grooves soon disappearing with wear, the cutting edge 
entire, the posterior surface concave with narrow but distinct 
heel ; i 3 with crown area about double that of i 2 and nearly half 



347 



that of canine, its crown higher than in the other incisors, 
owing to lower insertion of root, bujt extremity of cutting edge 
not extending beyond level of smaller teeth ; laterally the cutting 
edge is continued down outer-posterior margin of shaft where it 
functions against lower canine. Lower incisors not so large as 
upper, forming a continuous, nearly straight row between 
canines, the shafts straight, sloping obliquely forward, the root 
of « 2 implanted behind the others ; cutting edge irregularly 
2-lobed, that of i 3 longer than the others. Canines rather short 
and weak, the shaft simple, that of upper tooth nearly straight, 
that of lower tooth strongly curved backward, the enamel surface 
of both essentially smooth, though that of lower canine shows 
indications of a rudimentary cingulum in front. Anterior 
premolar both above and below a minute or spicular tooth closely 
crowded between canine and 
second premolar, pm 1 early 
deciduous, pm^ usually more 
persistent. Second premolars 
similar to each other, the crown 
area about equal to that of i 2 , 
the outline irregularly oval when 
viewed from above, the cusp 
about as high as long, its apex 
slightly in front of middle of 
crown ; root of each tooth single, 
that of pm 2 showing a tendency 
to become divided longitudi- 
nally. Other small premolars 
(pm 3 , pm 3 and pm^j alike in form, 
the crowns laterally compressed, 
triangular when viewed from 
the side, with apex slightly in 
front of middle, pin 3 and pm 3 
nearly alike in size, pm i some- 
what longer ; cingulum very slightly developed, not forming 
secondary cusps. Upper carnassial triangular in outline, the 
postero-internal border longest ; outer, trenchant portion of teeth 
consisting of a main anterior cusp resembling pm i in size and form, 
joined by a nearly horizontal commissure to a moderately high 
posterior cusp ; cingulum low but bearing three distinct secondary 
cusps, one at anterior base of main cusp, one (sometimes obsolete) 
near middle of antero-internal border, and the third and largest 
at middle of postero-internal border. Lower carnassial with 
anterior triangle well developed, the three cusps sub-equal in 
height, the metacouid subterete, the protoconid more compressed 
than paraconid, its commissure slightly longer than that of the 
anterior cusp ; crushing portion of crown longer and wider than 
anterior triangle, its inner portion occupied by a large basin-like 
concavity, its border with two large cusps on outer edge (of 




no. 71. 

Meles meles. Teeth. Nat. size. 



348 



CAKNIVORA 



which the anterior is the larger), a large cusp similar to postero- 
external cusp at middle of inner margin ; posterior margin 
occupied by three or four minute cusps soon disappearing with 
wear, and a similar minute cusp sometimes .in angle behind 
protoconid and metacooid. Second lower molar flat, terete, its 
area about half that of crushing portion of carnassial ; middle of 
crown with basin-like concavity ; margin with small but evident 
antero-external and postero-external cusps, and a smaller 
elevation at middle of inner edge. Upper molar rhomboidal 
in outline, the inner and outer margins parallel, the former 
nearly It 1 , times as long as latter; outer border occupied by two 
rather large, subterete, conical cusps, the anterior (paracone) 
slightly larger than posterior (metacone), so that greatest trans- 
verse diameter of crown is slightly behind anterior border ; 
remainder of crown a shallow basin-like concavity with rugose 
surface and raised, irregularly nodulate margin, the middle of 
concavity crossed by a ridge, convex internally, extending from 
anterior base of paracone to posterior base of metacone and 
bearing from three to five small cusps soon obscured by wear, 
the three anterior separated from two posterior by a deep angle ; 
the anterior group probably representing the protocone, the 
posterior group the hypocone. 

JRemarks. — Two moderately well differentiated local races are 
represented by the material that I have examined, one confined 
to the Iberian Peninsula, the other general in distribution. 
British specimens do not differ appreciably from Swedish 
examples of the typical form. 

Meles meles meles Linnseus. 

1758. [Ursus] meles Linnseus, Syst. Nat., i, 10th ed., p. 48 (Sweden). 

1785. [Meles] taxus Boddaert, Elenchus Animalium, i, p. 80 (Europe). 

1789. [Ursus meles} £ alba Gmelin, Syst. Nat., i, 13th ed., p. 102. 

1789. [Ursus meles'] y maculata Gmelin, Syst. Nat., I, 13th ed., p. 102. 

1808. Taxus vulgaris Tiedemann, Zoologie, i, p. 376 (Renaming of Ursus 



1816. Meles europmus Desmarest, Nouv. Diet. d'Hist. Nat., in, p. 465 

(Eenaming of XIrsus meles). 
1822. Taxus meles F. Cuvier, Hist. Nat. des Mamm., in, faso. 36, January, 

1822. 
1827. [Meles] communis Billberg, Synopsis Faunae Soandinaviae, p. 16 

(Renaming of Ursus meles). 

1827. [Meles communis] caninus Billberg, Synopsis Faunae Scandinavia, 
p. 17 (Scandinavia). 

1857. Meles taxus Blasius, Saugethiere Deutschlands, p. 204. 

1894. Meles meles Dahl, Die Heimat, iv, p. 125, June, 1894. 

1899. M[eles] m[eles] typicus Barrett-Hamilton, Ann. and Mag. Nat. Hist., 
7th ser., iv, p. 384, November, 1899. 



MELES 349 

1906. Meles meles britannicus Satunin, Mitteilungen des Kaukasisohen 
Museums, n, p. 115 (Based on the cranial measurements of 
English specimens recorded by Barrett-Hamilton, Ann. and Mag. 
Nat. Hist., 7th ser., iv, p. 384). Co-types in British Museum. 

1910. Meles meles and M. meles britannicus Trouessart, Faune Mamm. 
d'Europe, p. 69. 

Type locality. — Upsala, Sweden. 

Geographical distribution. — Central and southern Europe from 
southern Sweden to the Pyrenees and Italy, and from Ireland 
eastward ; eastern limit of range not known. 

Characters. — Skull and teeth moderately large, the teeth 
rarely attaining maximum size for the species ; colour moderately 
light, the sides not often conspicuously whitish. 

Measurements. — Adult male and female from Woodpark, 
Galway, Ireland: head and body, 686 and 618; tail, 153 and 
150 ; hind foot, 108 and 102 ; ear from meatus, 48 and 45. 
Adult female from Boueonne, Gers, France : head and body, 670 ; 
tail, 170; hind foot, 100; ear from meatus, 45. Adult female 
from Ingelheim, Rheinhessen, Germany : head and body, 672 ; 
tail, 180; hind foot, 104; ear from meatus, 50. For cranial 
measurements see Table, p. 350. 

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

Ibeland : Woodpark, Galway, 2. 

England: Burnley, Lancashire, 1; Ross, Hereford, 1; Kentchurch, 
Herefordshire, 4 ; Tetworth, Oxfordshire, 1 ; Ibstone, Buckinghamshire, 1 ; 
Horsham, Sussex, 1 ; Bicton, Devonshire, 1 ; Oornwall,l; no exact locality, 4. 

Norway : Holme, Mandal, 7 ; Egersund, Stavanger, 2. 

Sweden : Middle Sweden, no exact locality, 1 (Stockholm) ; Takern, 
Ostergotland, 1 (Stockholm) ; Smaland, 1 (Stockholm) ; Skane, 2 (Stock- 
holm) ; no exact locality, 1. 

Ekance : Eor£t de Boueonne, Gers, 1. 

Gebmany : Ingelheim, Bheinhessen, 1 ; Strass, near Burgheim, 
Bavaria, 1. 

Italy : Liguria, 1 (Genoa). 

<5, ?. Woodpark, Galway, Ire- R. E. Hibbert (p). 95. 6. 2. 1. 
land. 95. 12. 3. 1. 

(Co-types of M. m. britannicus, Satunin.) 
?. Burnley, Lancashire, Hon. N. C. Roths- 0. 10. 31. 4. 

England. child (p). 

i. Ross, Herefordshire. Hon. N. C. Roths- 0. 10. 31. 3. 

child (p). 
9, 3 juv. st. Kentchurch, Hereford- W. E. de Winton 
shire. (c & p). 

6. Tetworth, Oxfordshire. Hon. • H. Parker 

(c & p). 
i. Ibstone, Buckingham- A. H. Cocks (c & p). 4. 1. 25. 1. 

shire. 
A. Horsham, Sussex. E.C. Hawes (c & p). 7.4.13.1. 

9. Bicton, Devonshire. J.C.Stagdon(c&p). 97.2.23.1. 

1 (albino) Cornwall. Lord Lilford (p). 89. 10. 28. 1. 
skull. Sweden. Purchased (Wheel- 64. 3. 8. 1. 

wright). 
4 (J, 2 9. Holme, Mandal, Norway. R. J. Cuninghame 8. 8. 9. 7-12. 

(c & p). 

2 skulls. Egersund, Stavanger. K. H. Schaanning /ll. 6. 3. 13. 

(o), ill. 10. 23. 2. 



350 



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352 



CARNIVORA 



9. Foret de Bouconne, Gers, 0. Thomas (p). 8.7.15.5. 

250 m. Prance. (A. 

Robert.) 
9. Ingelheim, Rheinhessen, C. Hilgert (o). 8. 11. 2. 16. 

Germany. 



Meles meles marianensis Graells. 

1897. [Meles taxus] var. marianensis Graells, Mem. Real Acad. Cien., 

Madrid, xvn, p. 170 (Central Spain). 
1899. Meles meles mediterraneus Barrett-Hamilton, Ann. and Mag. Nat. 

Hist., 7th ser., iv, p. 384, November, 1899 (Seville, Spain). Type 

in British Museum. 
1910. Meles meles mediterraneus Trouessart, Faune Mamm. d'Europe, p. 70. 

Type locality. — Central Spain. 

Geographical distribution. — Iberian Peninsula. 

Diagnosis. — Skull and teeth large, the teeth frequently attain- 
ing maximum size for species ; colour light, the sides often 
conspicuously whitish, especially in region bordering dark ventral 
area. 

Measurements. — For cranial and dental measurements see 
Table, p. 351. 

Specimens examined. — Seven, from the following localities in Spain: 
Arrechavaleta, Vitoria, 1 ; Quintanar de la Sierra, Burgos, 2 ; near 
Seville, 2 ; Coto Dofiana, Huelva, 2. 

Remarks. — Although not very strikingly differentiated, the 
large-toothed, pallid, Iberian badger seems worthy of recognition 
as a geographical race. 

6. Arrechavaleta, Vitoria, N. Gonzalez (c). 8.7.7.18. 

Spain. 
6, 9. Quintanar, Burgos. Bev. Saturio Gon- 8. 7. 7. 19-20. 

6,9. Seville. (A. Ruts.) Lord Lilford (p). 95.3.3.7-8.1 

(95. 3. 3. 7. Type of mediterraneus Barrett-Hamilton.) 
<S, 9. Coto Donana, Huelva. A. Chapman (p). 8. 3. 8. 6-7. 

MELES ARCALUS Miller. 

1899. Meles meles mediterraneus Barrett-Hamilton, Ann. and Mag. Nat. 
Hist., 7th ser., iv, p. 131, November, 1899 (part). 

1906. Meles meles mediterraneus Bate, Proc. Zool. Soc, London, 1905, ir, 

p. 318, April 5, 1906. 

1907. Meles arcalus Miller, Ann. and Mag. Nat. Hist., 7th ser., xx, p. 394, 

November, 1907. Type in British Museum. 
1910. Meles arcalus Trouessart, Faune Mamm. d'Europe, p. 70. 

Type locality. — Lassethe Plain, Crete. 
Geographical distribution. — Island of Crete. 
Characters. — Size small (upper length of skull in adult less than 
110 mm. ; maxillary tooth-row, exclusive of incisors, about 35 mm. ; 



353 



DENTAL MEASUREMENTS OP MELES MELES AND M. ABC ALUS. 



Looality. 


Number. 


Sex. 


Tr , ' Lower molar 
Upper molar. , (flrst) 


M. meles meles. 




i 




Sweden : no exact locality 


64. 3. 8. 1 


9 


13-0 X 11-4 


15-4 x 7-0 


Wegelin 


Stockholm 


£? 


15-0 x 130 


16-2 x 7-8 


Skane 


,, 


6 1 


15-2 x 13-0 


18-0 X 8-2 


Ulriksdal . 


,, 


<5juv. 


13-4 x 11-6 


15-8 x 7-8 


Smaland . 


,, 


9? 


14-2 x 11-8 


16-2 x 7-4 


Orebro . . . . ! 


, 


5 juv. 


14-2 x 11-8 


15-8 X 7-0 


Takern .... 


„ 


6 ? 


13-6 X 11-2 


15-6 X 7-2 


Norway : Holme, Mandal 


152622 


6 


11-6 x 10-4 


14-0 X 6-4 


• 


8. 8. 9. 9 


i 


12-8 x 11-8 


14-6 x 7-0 


» „ 


8. 8. 9. 10 


6 


12-6 x 11-2 


14-6 X 7-0 


„ „ 


8. 8. 9. 11 


9 


11-2 X 10-8 


14-0 x 6-8 


)i 


8. 8. 9. 12 


9 


12-2 x 10-4 


14-0 x 7-0 


Egersund, Stavanger 


11. 6. 3. 13 




15-0 X 11-4 


14-8 x 7-0 


England : Boss, Hereford 


0. 10. 31. 3 


i 


13-2 x 11-4 


16-0 x 8-0 


Cornwall 


98. 10. 28. 1 


9 ? 


14-0 x 12-0 


16-6 x 7-8 


no exact locality 


2116 


SI 


13-4 x 12-0 


16-4 x 8-0 


,> 


211c 


9? 


14-0 X 12-4 


16-2 x 8-2 


» .. 


— 




14-2 x 12-2 


— 




— 




13-6 X 11-8 


— 


Burnley, Lancashire 


0. 10. 31. 4 


9 


14-4 x 12-2 


16-2 x 8 


2 


Cornwall 


— 




15-0 x 12-2 


18-0 X 8 


2 


Ireland : Woodparb, Co. Galway 


95. 12. 3. 1 




15-6 x 12-8 


17-0 x 7 


8 


,, *> 


95. 6. 2. 1 




14-8 X 12-2 


16-0 x 7 


8 


Denmark: Zealand. 


Andersen 


6 


14-0 X 13-2 


17-0 X 7 


8 


>> 


Copenhagen* 


S 


15-0 X 12-6 


18-5 x 8 


2 




„ 


6 


15-0 X 13-7 


18-3 x 8 


3 






S 


15-5 x 13-6 


18-0 x 9 









9 


14-6 X 125 


17-5 x 7-7 


various localities, (Maxi-1 
13 specimens : \ mum J 






16-3 X 13-6 













fMini-1 
\mumj 






13-0 X 12-0 


— 










various localities, f Maxi-1 
17 specimens : \ mum | 









18-7 x 8-5 


" 








jMini-1 
\ mum / 









150 x 7-0 










France: Bouconne, Gers. 


8. 7. 15. 5 


9 


14-0 X 120 


16-2 X 7-2 


Germany : Ingelheim, Rheinhessen . 


8. 11. 2. 16 


9 


14-8 x 12-8 


1^-2 X 7"8 


Italy: Borghetto S. Spirito, Liguria 


33 Genoa 


9 


13-6 X 110 


15-6 X 72 


M. meles marianensis. 










Spain : Quintanar, Burgos 


8. 7. 7. 19 


S 


15-2 X 12-6 


17-4 x 8-6 


Seville 


95. 3. 3. 7t 


i 


15-0 X 12-0 


16-6 x 7-8 




95. 3. 3. 8 


9 


14-6 X 12-6 


16-8 x 8-2 


Coto Dofiana, Huelva . 


8. 3. 8. 


6 


16-4 X 13-2 


17-8 X 8-0 


» >) 


8. 3. 8. 7 


9 


14-8 X 12-0 


17-6 X 8-2 


M. arcalus. 










Crete .... 


5. 12. 2. 17t 


?juv 


! 13-6 x 11-2 


15-4 X 7-2 


>>...... 


5. 12. 2. 38 


ad. 


13-8 X 11-6 





* Measured by H. Winge. 



+ Type of mediUiTanevx Barrett-Hamilton. 

■2 



t Type. 



354 



CAENIVOEA 



hind foot about 25 mm.) ; auditory bulla strongly inflated, the 
highest region near middle of bulla proper (exclusive of meatal 
tube) and so broadly rounded as not to form a longitudinal 
ridge, the region between highest portion and meatus not notice- 
ably concave ; teeth smaller than in Meles meles (lower molars 
together about 20 mm. in length), with smaller cusps tending to 
be more strongly developed ; metacone of m 1 relatively larger 
than M. meles, so that breadth of crown through this cusp is as 
great as or greater than that through paracone ; colour as in 
Meles meles marianensis or slightly paler. 

Measurements. — For cranial and dental measurements see 
Tables, pp. 351, 353. 

Specimens examined. — Four, all from Crete. 

(5,9. Lassethe, 2,820 ft. Crete. Miss D. Bate (c)'. 5.12.2.16-17. 

(5. 12. 2. 17. Type of species.) 

skull. Katharo, Crete. Miss D. Bate (c). 5. 12. 2. 38. 

9. Crete. H. 0. Jones, R.N. (c). 99. 6. 13. 1. 

Sub-Family LUTEINS. 
1857. Lutrinse Baird, Mamm. North Amer., p. 148. 

Geographical distribution. — Nearly cosmopolitan ; absent only 
in the Antarctic and high Arctic regions, Madagascar, Australia 
and the Pacific Islands. 

Characters. — Teeth of the same general type as in the 
Melinse ; skull much flattened, and rostrum so shortened that its 
length is less than its width ; external form long and slender, 
the fur very dense, the legs unusually short ; toes webbed, the 
claws short or absent ; tail long and highly muscular. 

'Remarks. — The family Lutrinse, the members of which appear 
to be essentially badgers modified for semi-aquatic life, contains 
four genera, one of which occurs throughout Europe from 
Ireland eastward. 

Genus LUTRA Brisson. 

1762. Lutra Brisson, Regnum Animale in Classis rx distrib., 2nd ed., p. 13 

(Lutra Brisson = Mustela lutra Linnaeus). 
1780. Lutra Briinnich, Zoologiae Fundamenta, p. 34. First use of name 

by an author following the Linnaean system (Mustela lutra). 
1806. Lutris Dumeril, Zoologie Analytique, p. 12 (Modification of Lutra). 
1817. Lutrix Rafinesque, Analyse de la Nature, p. 59 (Substitute for Lutra). 
1843. Lontra Gray, Ann. and Mag. Nat. Hist., xi, p. 118, February, 1843 

(Lutra canadensis Schreber). 
1843. Latax Gray, Ann. and Mag. Nat. Hist., xi, p. 119, February, 1843 

(Lutra lataxina F. Cuvier). Not Latax Glager, 1827. 
1843. Lataxina Gray, List Spec. Mamm. Brit. Mus., p. 70 (L. mollis Gray 

= Lutra lataxina F. Cuvier). 
1857. Lutra Blasius, Saugethiere Deutschlands, p. 236. 
1865. Barangia Gray, Proc. Zool. Soc. London, p. 123 (B. sumatrana Gray 

= Lutra barang F. Cuvier). 



LDTRA 355 

1865. Lutrogale Gray, Proc. Zool. Soc. London, p. 127 (Lutra monticola 

Hodgson). 
1865. Nutria Gray, Proo. Zool. Soo. London, p. 128 (Lutra felina Molina). 
1867. Lutronectes Gray, Proo. Zool. Soc. London, p. 180 (L. whiteleyi 

Gray = L. lutra Linnseus ?). 

Type species. — Lutra * Brisson = Mustela lutra Linnaeus. 

Geographical distribution. — Same as that of the sub-family. 

Characters. — Skull broad and low (depth of brain-case only 
about half mastoid breadth), the entire dorsal profile nearly 
straight, the zygomatic arches so widely expanded and post- 
orbital region so much narrowed that distance from point of 
greatest narrowing to zygoma is greater than postorbital con- 
striction ; rostrum so shortened that posterior border of narial 
opening approaches level of anterior zygomatic root, and distance 
from orbit to gnathion is much less than half length of brain- 
case ; auditory bulla flattened, with conspicuous meatal tube ; 
paroccipital process low and ridge-like but distinct from bulla ; 
dental formula : i ||, c £?, pm g, m £? = 36 ; teeth large as com- 
pared with width of rostrum and palate (greatest transverse 
diameter of upper carnassial equal to half distance between 
carnassials) ; all of the small premolars opposed and capable of 
trenchant action with those of opposite jaw ; upper carnassial 
with crown triangular in outline, the outer side occupied by a 
moderately high cutting edge formed by two cusps and a con- 
necting ridge, the inner side (about half the total crown area) 
by a flat crushing surface ; upper molar rhombic in outline, its 
crown area about equal to that of carnassial (usually somewhat 
smaller), its greatest diameter transverse to axis of tooth-row, 
the four primitive cusps present ; lower carnassial with three 
anterior cusps well developed, sub-equal, the posterior heel 
slightly larger than anterior triangle ; external form highly 
modified for aquatic life, the body long and of approximately the 
same width as neck and head, the tail long, very muscular, broad 
at base, tapering distally, the legs short, feet broad, toes webbed, 
short-clawed, the fur soft, dense and impervious to water. 

Bemarks. — Although more widely distributed than any other 
living genus of land mammals, Eptesicus and Myotis perhaps 
excepted, Lutra is not rich in species. A dozen or fifteen forms 
are currently recognized, only one of which occurs in Europe. 

LUTRA LUTRA Linnaeus. 

1758. [Mustela] lutra Linnseus, Syst. Nat., i, 10th ed., p. 45 (Sweden). 
1777. [Lutra'] vulgaris Erxleben, Syst. Regni Anim.,i, p. 448 (Renaming of 

lutra). 
1792. M[ustela] Lutra giscatoria Kerr, Anim. Kingd., p. 172 (Renaming of 

lutra). \i 



Not " Lutra lutra" (see footnote under Meles, p. 341). 

2 A 2 



356 



OARNIVOEA 



1816. ? Lutra fluviatilis Leach, Syst. Gatal. Spec. Indig. Mamm. and 

Birds Brit. Mus., p. 6 (nomen nudum : " Kiver Otter "). 
1827. [Lutra vulgaris] j8 marinus Billberg, Synopsis Faunae ScandinaviEe, 

p. 28 (Coasts of Scandinavia). 
1830. ? [Lutra] fluviatilis Burnett, Quart. Journ. Sci. Lit. Art. xxviii, 

1829, p. 349 (Substitute for lutra), nomen nudum. 
1834. [Lutra] nudijoes Melchior, Den Danske Stats og Norges Pattedyr, 

p. 50 (Coast of northern Norway). 
1834. Lutra roensis Ogilby, Proo. Zool. Soc. London, p. Ill (Roe Mills, 

near Newton Lemavaddy, Londonderry, Ireland). Type in British 

Museum. 
1857. Lutra vulgaris Blasius, Saugethiere Deutschlands, p. 237. 

1884. [Lutra] lutra Lataste, Actes Soc. Linn, de Bordeaux, xxxvni, p. 34. 

1885. [Lutra] angustifrons Lataste, Actes Soc. Linn, de Bordeaux, xxxix, 

p. 168, August, 1885 (Bone, Algeria). Perhaps in part only : 
specimen from Liguria, Italy, referred to this form on p. 239, 
September, 1885. Type in Lataste collection. 
1910. Lutra lutra Trouessart, Paune Mamm. d'Europe, p. 86. 

Type locality. — Upsala, Sweden. 

Geographical distribution. — Europe and northern Africa, east- 
ward into Asia ; limits of range not known. In Europe west to 
Ireland and north to the Arctic coast. 

Diagnosis. — Size medium (head and body in adult male about 
700 mm., condylobasal length of skull, 105 to 123); tail about 
three-quarters as long as head and body ; naked muzzle pad with 
upper border strongly convex at middle ; skull much flattened, 
the depth of brain-case not conspicuously more than half mastoid 
breadth; interorbital region narrow, its least width less than 
distance from front of zygoma to anterior extremity of pre- 
maxillary ; teeth not specially enlarged, the greatest diameter of 
upper carnassial not greater than width of palate between 
carnassials. 

External characters. — General form long and slender, the 
limbs very short, the feet broad, with conspicuously webbed toes, 
the head short flat and ill-defined from neck, the ears incon- 
spicuous, the tail long, broad at base, tapering toward tip, 
covered with the same short waterproof fur as body. Head 
rounded and flattened, not well defined externally from the 
muscular neck ; ear rounded, densely haired on both surfaces, 
scarcely rising above level of fur, the antitragal lobe valve-like ; 
a second and third valve-like lobe above and behind meatus ; 
muzzle short and wide, the nostril pad entirely naked, its 
surface reticulate, its upper margin strongly convex at middle, 
the lower slightly so, its lower border separated from mouth by 
the densely haired upper lip, the width of which at middle is 
about equal to height of pad ; whiskers stiff and bristly, the 
longest extending about to ear when laid back. Legs short, feet 
broad and rounded, with short toes joined by a naked membrane 
extending to base of terminal phalanges ; claws short but strong, 
non-retractile, those on fore-feet best developed (about 8 mm. in 



LUTKA 



357 



length) ; palm bare, a large, heart-shaped, obscurely trilobed 
tubercular mass behind base of digits, and a round posterior 
tubercle separated from the larger pad by a deep groove ; sole 
with a bare area and heart-shaped pad essentially as on palm, 
but with posterior tubercle represented by an ill-defined prolonga- 
tion of the main pad ; posterior portion of sole densely furred. 
Fur very dense and waterproof, alike in texture throughout the 
body and tail, the hairs of underfur 10 to 15 mm. in length, the 
longer overlying hairs, which almost completely conceal the 
underfur, about 25 mm. in length. Mammae : 6* (probably i 3-3). 
Colour. — Winter pelage : upper parts, legs, feet and tail a 
rich dark brown (about the prout-brown of Ridgway or some- 




RG. 72. 
Lutra lutra. Nat. size. 



what darker), with a drabby cast more evident in some lights 
than in others, the hairs with a conspicuous metallic gloss ; 
underfur light grey, the extreme tips of its hairs changing 
abruptly to prout-brown ; on underparts the drab becomes more 
conspicuous as well as paler, usually assuming a tinge of cream- 
buff, the throat and cheeks fading to buffy white ; interramial 
region and upper lip with irregular white mottlings, the hairs of 
which are white to base ; whiskers and claws light horn-colour. 

The exact colour is subject to considerable variation, but the 
material examined is not sufficient to show whether such 
differences as occur are correlated with locality or season. 

* Southwell, Field, en, p. 1043, December 19, 1903. 



358 



CAKNIVOKA 



Sometimes the brown is darker and richer than usual, or the 
drab may be especially pronounced. Occasionally the long hairs 
of the back are a light dull buff, imparting to the animal a 
peculiar faded appearance. 

Skull. — General form of skull broad and flattened, more so 
than in any other European carnivore, the depth of brain-case at 
middle scarcely more than half mastoid width ; the brain-case exces- 
sively narrowed anteriorly, broad posteriorly and with greatly 




Fie. 
Lutra lutra. Nat. size. 



developed lambdoid crest, but low though evident sagittal crest ; 
widely spreading zygomata forming a marked contrast with narrow 
postorbital region ; rostrum short, deep and robust. Dorsal profile 
nearly flat throughout, though region in front of middle of post- 
orbital constriction slopes gradually downward. Owing to depth 
of rostrum and shallowness of brain-case the dorsal and ventral 
profiles are approximately parallel. Postorbital processes short 
but evident, though not forming any considerable portion of border 
of rather large, somewhat upturned orbit, the greatest diameter 



LUTRA 



359 



of which Dearly equals width of .flattened inter-orbital region. 
Rostrum short, broad and deep, the distance from front of 
zygoma to gnathion barely equal to greatest breadth across 
canines, about equal to depth at front of orbit, and contained 
about 4J times in condylobasal length of skull. Zygoma 
widely spreading throughout, the arch heavy, not much bowed 
upward, the anterior root conspicuously perforated by the large 
anteorbital foramen the upper margin with low but evident 




na. 74. 
iMtra lutva. Nat. size. 



angular projection marking posterior border of orbit. Palate 
rather narrow, the distance between molars contained about 
three times in that from gnathion to level of posterior border of 
molar : incisive foramina moderate, about half as wide as long, 
lying entirely between canines ; posterior extension of palate 
about equal to distance between molars and about one-third that 
from molar to hamular ; mesopterygoid space narrow anteriorly, 
wider posteriorly, its length about double the width between 
hamulars. Auditory bulla small, flattened, though a little 



360 



CAENIVOBA 



inflated along inner margin ; outline triangular-flask shaped, the 
tubular portion of meatus not well defined from rest of bulla ; 
least distance between bullae about equal to greatest diameter of 
bulla including meatus. Paroccipital process low and ridge- 
like, widely removed from inflated portion of bulla. Mandible 
very robust, the ramus slightly bowed outward posteriorly, its 
lower margin essentially straight except for a slight upward 
curve behind level of tooth-row, the depth of ramus at middle 
about one-third that through coronoid process ; angular process 
very short ; coronoid process narrowly rounded off above, its height 
above articular process about equal to width at articular level. 

Teeth. — The teeth are large and strong, with well developed 
cusps and commissures and relatively small crushing areas, the 
small premolars of upper and lower mandible fitting closely 
between each other when jaws are shut; crown area of upper 

molar about equal to that of car- 
nassial, sometimes less. Upper 
incisors forming a nearly straight 
transverse row, the anterior faces 
of the smaller teeth exactly in line, 
those of the two larger teeth slightly 
more posterior ; i 1 and i 2 small, 
sub-equal, the shafts compressed, 
the cutting edges rounded, simple ; 
i 3 higher than the others, somewhat 
resembling the lower canine in form, 
its apex curved outward and back- 
ward beyond level of smaller teeth, 
its postero-external surface with two 
concavities separated by a trenchant 
ridge ; lower incisors subterete, the 
crown of « 3 obscurely and unsym- 
metrically bilobed, that of the others 
simple; cross section of i x about. one-third that of i 3 . Canines 
robust though not greatly elongated ; shaft simple, without 
evident cingulum, that of upper tooth slightly curved backward, 
that of lower tooth abruptly recurved, its axis set obliquely 
outward. Anterior premolar both above and below well 
developed and functional, the point of the upper tooth lying 
internal to that of lower when jaws are closed ; pm 1 small, single 
rooted, its crown area about equal to that i 2 , its form essentially 
like that of succeeding tooth, though with cusp relatively less 
developed ; pm 2 and pm 3 successively larger, two-rooted, the 
crown area of pm 3 fully three-fourths that of canine, each tooth 
with a well-developed cusp lying in front of middle of crown, 
and a distinct anterior and posterior cutting ridge ; cingulum 
moderately developed ; outline of crown of pm 2 elliptical, that of 
pm 3 with inner margin bulging inward behind middle, though 
without developing a definite inner lobe ; pm 2 and pm 3 much like 





FIG. 75. 
Lutra lutra. Teeth. Nat. size. 



LUTRA 361 

pm 2 in both size and form, the three teeth cutting against each 
other when jaws are closed ; pm i larger than the other small 
premolars, its cusp less anterior in position, its crown decidedly 
in wider posteriorly than anteriorly. Upper carnassial triangular 
general outline, though this is somewhat obscured by the broadly 
rounded inner portion and the somewhat projecting postero- 
external lobe ; outer and posterior borders sub-equal and longest ; 
trenchant portion of tooth with high anterior cusp connected 
with well developed posterior cusp by a high, abruptly angled 
commissure ; crushing portion nearly as wide as long, its area 
about as great as that of trenchant portion, its inner border 
noticeably raised at middle though without secondary cusps ; 
cingulum moderately developed along outer border of crown, and 
forming a distinct though small antero-basal cusp. Lower 
carnassial with anterior triangle well developed, the protoconid 
and paraconid sub-equal, the metaconid somewhat smaller ; 
crushing portion of crown wider than anterior triangle though 
not so long, the areas of the two portions of the tooth 
approximately equal ; hypoconid evident though not high, the 
outer surface of its base in line with that of protoconid, the 
inner surface continuous with the concave though scarcely 
basin-shaped main portion of crushing area ; cingulum narrow 
but evident throughout, not specially developed in region 
bordering outer base of hypoconid. Second lower molar flat, 
subterete, slightly wider than long, the inner and outer margins 
each with a small cusp somewhat behind middle, the cusps joined 
by a low transverse ridge. Upper molar about equal to carnassial 
in crown area, but length of its outer border decidedly less than 
that of preceding tooth ; crown slightly constricted near middle, 
the outer portion bilobed, the anterior lobe bearing a low paracone 
and a broad outer projection representing the parastyle, the 
posterior lobe bearing a low but robust metacone ; inner portion 
of crown with a large protocone and low hypocone, the former 
sending forward a conspicuous commissure to base of paracone, 
the latter connected with cingulum that extends around base of 
protocone. 

Measurements. — Adult male from Warwickshire, England : 
head and body, 712; tail, 495; hind foot, 134; ear, 28. Adult 
female from Csehtelek, Hungary : head and body, 640 ; tail, 380 ; 
hind foot, 115; ear, 22. Por cranial and dental measurements 
see Tables, pp. 362, 363. 

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

Ireland : Londonderry, 1 (type of roensis) ; Ahascragh, Oo. Galway, 1. 

England : Pembrokeshire, 1 ; Eiver Stour, Dorsetshire, 1 ; Rugby, 
Warwickshire, 1 ; Norfolk, 1 ; no exact locality, 1. 

Nobwat : Egersund, Stavanger, 5. 

Sweden : Jockmock, Lappmark, 1 (Stockholm) ; Gnesta, Soderman- 
land, 1 (Stockholm) ; Skane, 1 (Stockholm) ; no exact locality, 2. 

France : St. Gilles, Gard, 1 (Lataste) ; Etupes, Doubs, 1 (Mottaz). 

Germany : Southern Germany, 2. 




Observations. 


eth moderately worn. 
, not worn. 

much worn. 

slightly worn. 
, , not worn. 

slightly worn. 

> 

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, slightly worn. g 
, not worn. 


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, not worn. 


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CAKNIVOEA 



Austbia-Hongary : Karlsbad, Bohemia, 1 ; Csehtelek, Bihar, Hun- 
gary, 1. 

Italy: Torriglia, Liguria, 1 (Lataste). 

Spain: Castrillo de la Beina, Burgos, 3(B.M.andU.S.N.M.); Galioia, 1; 
Seville, 5. 

Remarks. — Although the specimens show some rather note- 
worthy variations in colour, and in cranial and dental measure- 
ments, the material examined is not sufficient to indicate the 
existence of geographical races in Europe ; while the north 
African animal to which the name augustifrons has been applied 
appears to be not separable from true Intra. 



1 . Newton Lemavady , 

Londonderry, Ireland. 
(Miss A. Moody.) 
6 st. Ahascragh, Galway. 

9, 3 juv. st. Pembrokeshire, Wales. 

5. Bugby, Warwickshire, 

England. 
9 st. B. Stour, Dorset. 

1. England. 

4 6, 1 9, Egersund, Stavanger, 

skulls. Norway, 

ad., juv. South Germany. 

6. CsShtelek, Bihar, Hun- 

gary. 
1. Karlsbad, Bohemia. 

6, 9. Castrillo de la Beina, 

Burgos, Spain. 
1 juv. Galicia. 

skull. Seville. 

9, juv. Seville. (Dr. A. Ruiz.) 
skull. Seville. 



Zoological Society's 57. 12. 14. 4. 
Museum. (Type of L. roensis 
Ogilby). 



Purchased (Bow- 
land Ward). 

Purchased (Bow- 
land Ward). 

Dr.T. S.Townsend 
(c & p). 

J. 0. Mansel Pley- 
dell (c & p). 

K. H. Schaanning 
(c). 

Dr. A. Giinther (c). 

Hon. Mrs. N. 0. 
Bothschild (p). 

Lord 0. Bussell 
(c & p). 

Bev. Saturio Gon- 
zalez (c). 

Dr. V. L. Seoane 
(c & p). 

Lord Lilford (p). 

Lord Lilford (r). 
Col. L. H. Irby 
(c & p). 



5. 7. 10. 1. 
5. 5. 21. 1-4. 

9. 12. 16. 1. 

98. 5. 13. 1. 

99. A. 
11.6.3.14-18. 

59. 9. 6. 62-63. 

10. 9. 14. 1. 

57. 2. 14. 1. 

8. 7. 7. 16-17. 

94. 6. 18. 1. 

74. 10. 7. 2. 
76. 3. 4. 1. 
95.9.4.10-11. 
2. 11. 16. 1. 



Sub-Family MUSTELINE. 
1835. Mustelinn: Swainson, Nat. Hist, and Classif. Quadr., p. VII. 

Geographical distribution. — Northern hemisphere, south into 
northern Africa, the Malay Archipelago, and northern and 
western South America ; in Europe west to Ireland. 

Characters. — Dentition highly trenchant, upper carnassial 
without crushing surface other than a small concave area between 
inner lobe and main cusp, the crown much longer than broad, 
not triangular or rhombic in outline, the posterior cusp compressed, 
trenchant, barely half as high as main cusp ; upper molar much 
reduced, the length of its outer portion one-third to one-half that 
of carnassial ; small premolars alternating when jaws are closed, 
at least one pair capable of shearing action : skull varying in 



MARTBS 365 

form but never greatly flattened and never very robust, the 
rostrum always at least as long as broad ; external form slender, 
the legs usually rather short ; feet digitigrade ; toes partly 
webbed ; tail varying in length, slender or bushy, never con- 
spicuously muscular. 

Remarks. — As here understood the sub-family Mustelinm 
contains the three genera, Martes, Mustela and Vormela, all of 
which occur in Europe. 



Genus MARTES Pinel. 

1792. Martes Pinel, Actes Soo. d'Hist. Nat., Paris, i, p. 55 (M. domestica 

Pinel = M. foina Erxleben). 
1820. Martes Nilsson, Skand. Fauna, I, p. 38 (M. foina and M. sylvatica = 

martes). 
1829. Zibellina Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierw., i, 

p. 31 (M. zibellina). 
1857. Mustela Blasius, Saugethiere Deutschlands, p. 211. 
1911. Martes Thomas, Proo. Zool. Soo., London, p. 139, March, 1911. 

Type species. — Mustela martes Linnaeus. 

Geographical distribution. — Northern hemisphere from the 
limits of tree growth south to the Mediterranean, the Malay 
Archipelago, and the eentral United States ; in Europe west to 
Ireland. 

Characters. — Skull narrow, moderately high (depth of brain- 
case much more than half mastoid breadth), the dorsal profile 
moderately curved, the zygomatic arches not specially wide- 
spreading, and postorbital region not unusually narrowed (distance 
between region of greatest narrowing and zygoma normally less 
than breadth of postorbital constriction) ; rostrum narrow and 
somewhat elongate, its width noticeably less than that of inter- 
orbital region, the distance from anterior rim of orbit to gnathion 
exceeding width of rostrum between anteorbital foramina; auditory 
bullse moderately inflated, the meatal tube evident though short, 
the longitudinal diameter of bulla greatest ; paroccipital process 
small, slightly projecting, partly distinct from bulla ; dental 
formula : i t5, c t- 1 , prn ££, m t| = 38 ; cutting edges of five small 
premolars (2 upper and 3 lower) capable of trenchant action ; 
upper carnassial long and narrow, not triangular in outline and 
without crushing surface, the small inner lobe standing as an 
offset to antero -internal extremity of crown, the sectorial portion 
consisting of a high anterior and low posterior cusp with some- 
what concave connecting ridge ; upper molar pyriform or pan- 
durate in outline, its long axis nearly perpendicular to that of 
tooth-row, its crown mainly flat, but with a small paracone, still 
smaller, sometimes obsolete metacone, and crescentic ridge-like 
protocone ; lower carnassial wider posteriorly than anteriorly, 
the anterior triangle much distorted, the metaconid reduced to 



366 



CAKN1V0RA 



a well defined postero-internal process on base of protoconid, the 
posterior crushing heel slightly more than half as large as 
trenchant portion of tooth ; external form slender, somewhat 
catlike or squirrel-like, the head moderately elongated, the 
muzzle pointed, the ears high and conspicuous, the tail long, 
bushy ; feet digitigrade, the moderately long claws partly 
retractile ; fur long, dense and soft. 

Remarks. — The genus Martes contains about ten species, some 
of which are represented by numerous geographical races. Three 
are known from western and southern Europe. 

KEY TO THE EUROPEAN FORMS OP MABTES. 

Third .upper premolar with crown strongly 
convex on inner side, slightly concave on 
outer side ; width of inner lobe of upper 
carnassial nearly equal to that of tren- 
chant portion of crown ; greatest diameter 
of upper molar about equal to length of 
outer border of carnassial ; fur usually 
finer and softer, and throat-patch more 

yellow (Pine Martens) M. martes, p. 366. 

Throat-patch cream-buff, general colour 

darker (Central and northern Europe)... M. m. martes, p. 372. 
Throat-patch buff-yellow, general colour 

lighter (Mediterranean region) M.m. latvrurrum, p. 373. 

Third upper premolar with crown evenly bicon- 
vex ; width of inner lobe of upper car- 
nassial barely half that of trenchant 
portion of crown ; greatest diameter of 
upper molar noticeably less than length 
of outer border of carnassial ; fur usually 
coarser and less soft, and throat-patch more 
whitish (Beech Martens). 
Condylobasal length of skull in adult male, 
76 to 79 mm. ; pale throat patch always 
much encroached on by brown of sur- 
rounding parts, occasionally obliterated 

(Crete) M. buwites, p. 380. 

Condylobasal length of skull in adult male, 
79 to 84 mm. ; pale throat-patch seldom 
much encroached on by brown of sur- 
rounding parts and never obliterated 
(Central and southern Continental 

Europe) M. foina, p. 374. 

General hue of upper parts drab M.f. foma, p. 375. 

General hue of upper parts wood-brown .. M.f. mediterranea, p. 380. 

MARTES MARTES Linnaeus. 
(Synonymy under subspecies.) 

Geographical distribution. — Entire wooded region of Europe, 
from Ireland eastward into Asia, and from the Mediterranean 
coast and islands northward to the limits of tree growth. 

Diagnosis. — Third upper premolar with outline of crown 



MAETES 367 

concave on outer side, irregularly convex on inner side, the region 
of greatest width behind middle ; upper carnassial with inner 
projection robust, its diameter in line of tooth-row nearly or 
quite equal to greatest width of trenchant portion of tooth behind 
middle of crown ; upper molar large, its greatest diameter about 
equal to outer length of oarnassial, the metacone more than half 
as large as paracone, the outer border of the crown usually 
notched; general colour yellowish brov^n, the throat patch 




FIG. 76. 
Ma/rtes martes. Nat. size. 



always distinctly tinged with yellow ; underfur moderately long 
and dense. 

External characters. — Form slender and graceful, suggesting 
that of both cat and squirrel ; legs moderately long ; tail about half 
head and body, densely furred and bushy, the hairs at tip about 
one-third as long as vertebrae. Head pointed ; eyes of medium 
size ; ear rising distinctly above fur, its height from crown about 
25 mm., the outline rounded but forming a slightly indicated 
point above, the entire surface densely furred ; muzzle pad 



368 



CARNIVOKA 



well defined, completely naked, communicating with upper lip 
by narrow median line. Feet densely furred throughout in 
winter, the pads bare in summer ; palm with deeply divided, 
trilobed, heart-shaped tubercular mass at bases of median digits, 
a small round pad at base of thumb, and a larger round pad at 
outer posterior border near wrist ; sole with similar trilobed 
mass, but elements more distinct, almost forming three tubercles ; 
small pad at base of hallux like that at base of thumb ; no 
posterior pad. Fur very dense and soft, the hairs of underfur 
about 25 mm. long at middle of back in winter, shorter in 
summer, the longer hairs reaching about 40 mm. Mammae : 4. 

Colour. — The colour varies considerably with season and 
climate, but is usually characterized by rich yellowish brown and 
blackish tints, the legs, tail and underparts darker than the 
back, the head noticeably lighter than the body, the throat and 
fore part of chest yellowish, and interramia, chin and muzzle 
dusky. Claws light horn-colo'ur. 

Skull. — As compared with that of the European species of 
Mustela the skull of Maries mattes is characterized by a general 




Fig. 77. 
Maries martes. Nat. size. 



smoothness, lightness and lack of angularity. Dorsal profile 
strongly convex posteriorly, flattish behind orbits, the rostrum 
falling away at an angle of about 20° ; a slight concavity usually 
present near middle of nasals. Brain-case elongate-ovate in 
outline when viewed from above, nearly as deep as wide, the 



MARTBS 369 

sagittal crest moderately developed in old individuals and slightly 
overhanging occiput. Floor of brain-case with no* special 
features ; a slightly developed median ridge. Auditory bullse 
moderately inflated, the surface smooth, the general outline 
broadly flask-shaped (longitudinal diameter greatest), the meatal 
tube well defined, rather wider than long ; space between bullse 
about equal to diameter of bulla without tubular portion. 
Postorbital region rather abruptly constricted, much as in 
Mustela, its width about half that of brain-case. Interorbital 



Fig. 78. 
Marten martes. Nat. size. 

region wider than rostrum, the discrepancy in breadth greater 
than in Mustela erminea. Postorbital processes well developed 
though short. Rostrum relatively longer than in Mustela 
erminea or M. putorius, the distance from orbit to gnathion about 
equal to width between outer margins of anteorbital foramina, 
the width across canines much less than distance from foramen 
to gnathion. Orifice of anteorbital foramen over anterior root of 
carnassial, and under anterior border of rather large orbit. 
Zygamata rather widely but gradually spreading, strongly bowed 

2 b 



370 



CAKNIVOEA 



upward behind middle. Palate rather narrow, its width 
between molars equal to about 1£ times greatest diameter of 
molar, incisive foramina small, ovate, slightly oblique, at level of 
front of canine, the minute median foramen slightly behind 
middle ; posterior extension of palate broader than long, reaching 
about half way from level of molar to hamular ; mesopterygoid 
space about a quarter longer than broad, the hamulars short, 
slightly everted. 

Teeth. — In proportion to size of skull the teeth are larger 
than in the members of the other genera of Mustelinse occurring 
in Europe ; transverse diameter of upper molar much more than 
half width of palate between molars. Upper incisors forming a 
straight row separated at each side from canine by a space about 
equal to transverse diameter of i 1 and i 2 together ; crowns strongly 
compressed, the antero-poscerior diameter fully twice transverse 
diameter, the anterior face convex, the posterior concave with 

faintly indicated cingulum ; i' 2 slightly 
larger than i 1 , i 3 abruptly much larger 
than the other two teeth together, its 
crown more than one-third as high as 
that of canine and with the posterior 
concavity extending to outer basal 
portion. Mandibular incisors smaller 
and lower than the upper teeth, their 
crowns projecting obliquely forward, 
the root of i 2 implanted in jaw behind 
level of the other two; crowns ob- 
scurely bifid, the outer lobe smaller 
than the inner ; crown area of i t 
j. IG j 9 about one-third that of the com- 

Martesmartes. Teeth. Nat. size, pressed i 2 or the subterete i 3 . Canines 

with no special peculiarities ; diameter 
of the upper tooth along alveolus equal to one-half width of 
palate between canines, that of lower somewhat more; shaft 
of upper tooth with posterior longitudinal ridge, and an antero- 
internal ridge which near base curves backward across inner 
side of shaft nearly to base of posterior ridge ; cingulum barely 
indicated ; shaft of lower canine directed a little forward at 
base, a little backward beyond middle, the anterior profile 
strongly convex, the posterior equally concave ; surface of 
enamel, especially on basal half of tooth, much roughened by 
longitudinal wrinkles. Anterior premolar both above and below 
single-rooted, small (crown area about equal to that of i 1 and i 2 
together), subterete, the height of the single ill-developed cusp 
scarcely equal to diameter of crown in pm 1 , much less in pm v 
Other premolars, except upper carnassial, two-rooted, compressed, 
the outline of crown triangular when viewed from the side, with 
apex slightly in front of middle of tooth. Crown area of pm 2 
about three times that of pm 1 ; outline of crown flattened-elliptical 





MARTES 371 

or slightly concavo-convex, the concavity, when present, on outer 
side ; cusp simple, its height a little less than length of crown 
along base, its posterior surface with slightly developed longi- 
tudinal ridge. Crown of pm? somewhat longer than that of pm 2 , 
but its area much greater owing to widening of strongly convex 
inner side ; cusp essentially as in pm 2 though somewhat higher. 
Second lower premolar about equal in size to pm 2 , but axis of 
shaft more anterior in position ; pm 3 essentially like pm 2 , but 
cusp with faintly indicated anterior longitudinal ridge, and 
posterior ridge with a slight nodule or rudimentary basal cusp 
(sometimes absent) ; pm i noticeably larger than pm 3 and with a 
well developed secondary cusp at middle of posterior border of 
main cusp. Upper carnassial (pin 4 ) long and narrow, the width 
of crown just behind internal lobe less than half length along 
outer border, the internal lobe robust, its diameter in line of 
tooth-row nearly or quite equal to width of trenchant portion of 
crown, its axis nearly perpendicular to main axis of tooth, its cusp 
well developed, about as high as that of pm 1 , and separated from 
main cusp by deep concavity ; main cusp robust, its height 
contained about 1| times in length of crown, its axis slanting 
a little backward, its anterior border with well developed 
longitudinal ridge ; posterior cusp about half as high as main 
cusp, its outer surface sloping obliquely to well developed 
cingulum, the two cusps connected by a high, sharply trenchant 
and obtusely angled commissure. Lower carnassial with crown 
nearly 2| times as long as broad, the sectorial portion of the 
tooth consisting of a much distorted triangle, the paraconid 
forming anterior extremity of tooth, its anterior border nearly 
perpendicular, its posterior commissure meeting anterior commis- 
sure of the higher protoconid at an abrupt angle, the two together 
acting in opposition to the angled commissure of upper carnassial ; 
metaconid reduced to a subterete posterointernal process on 
base of protoconid, its area scarcely one-sixth that of larger 
cusp ; crushing portion of crown slightly broader than trenchant 
portion, its outline sub-circular with slightly raised edge, this 
edge forming a noticeable though low postero-external cusp 
separated from base of protoconid by a shallow but distinctly 
angled notch. Second lower molar about equal to heel of 
carnassial in size and essentially like it in form, the crown flat, 
with faintly indicated outer ridge and low postero-internal cusp. 
Upper molar large, its crown area at least equal to that of 
carnassial, its greatest diameter about equal to outer length 
of carnassial (see measurements, p. 378), the diameter of inner 
portion of crown usually much greater than that of outer 
portion, the median constriction well marked ; outer portion of 
crown with two small cusps, probably the paraoone and metacone, 
the latter more than half as large as former, the outer margin of 
tooth often though not invariably notched between them ; inner 
portion of crown with a. slightly curved, ridge-like antero- 

2 b 2 



372 



CARNIVORA 



external cusp (usually notched or partly divided into two), the 
surface elsewhere finely wrinkled ; in some specimens a minute 
tubercle, perhaps representing the hypocone, occurs near posterior 
border of crown in region of constriction between outer and 
inner portions ; cingulum low but evident, especially around 
inner margin of crown. 

Remarks. — The well - known pine marten is so strongly 
characterized as to require no special comparisons with any other 
European species except Martes foina (see account of latter). 
Two geographical races are currently recognized, though their 
status is by no means clear. 

Martes martes martes Linnseus. 

1758. [Mustela] martes Linnaeus, Syst. Nat., i, 10th ed., p. 46 (Sweden). 
1816. M[ustela] sylvesiris Oken. Lehrb. d. Naturgesch., hi, pt. 2, p. 1029 

(Renaming of martes). 
1827. M[artes] vulgaris Griffith, Guvier's Anim. Kingd., v, p. 123 

(Renaming of martes). 
1847. Martes sylvatica Nilsson, Skand. Fauna, i, 2nd ed., p. 41 (Renaming 

of martes). 
1857. Mustela martes Blasius, Saugethiere Deutschlands, p. 213. 

1910. Mustela martes Trouessart, Paune Mamm. d'Europe, p. 72. 

1911. [Martes] martes Thomas, Proo. Zool. Soc. London, p. 139, March, 

1911. 

Type locality. — Vicinity of Upsala, Sweden. 

Geographical distribution. — Europe north of the Mediterranean 
region, from Ireland eastward into Asia. 

Diagnosis. — Throat patch cream-buff or slightly more yellow. 

Colour. — Upper parts a rich dark brown, usually rather near 
the bister of Ridgway, the tips of the longer hairs blackish ; 
underfur light grey, the tips of the hairs tinged with drab or 
with wood-brown ; face essentially like back or not so dark, the 
muzzle and chin usually not darker than forehead ; ear edged 
with bufi'y drab ; tail very dark brown (nearly the seal-brown of 
Ridgway), its underfur raw-umber or somewhat darker ; feet 
and lower portion of legs blackish ; throat-patch varying 
considerably, but as a rule rather pale, nearly cream-buff or 
slightly more yellow. 

Measurements. — For cranial and dental measurements see 
Tables, pp. 376, 378. 

Specimens examined. — Twenty, from the following localities : — 

Ireland : Kenmare, Kerry, 1 ; Co. Kerry, no exact locality, 1. 

Scotland : No exact locality, 1. 

England : Keswick, Cumberland, 1 ; Cockermouth, Cumberland, 1 ; 
Pontrilas, Herefordshire, 1 ; Ludford Park, Herefordshire, 1 ; no exact 
locality, 3 (B.M. and U.S.N.M.). 

Norway : Egersund, Stavanger, 3. 

Sweden : No exact locality, 3 (U.S.N.M.). 

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

Germany : Southern Germany, 2. 

Austria-Hungary : Bohemia, 1. 



MARTES 373 

9. Kenmare, Kerry, Ire- E. Dodson (o). 94. 3. 27. 1. 

land. 

1. Kerry. J. H. Gurney (p). 72. 1. 10. 1. 

1. Scotland. (McLeay.) Hargitt Collection. 86. 9. 9. 1. 
st. Keswick, Cumberland, W. E. Wilson (c). 96. 11. 6. 1. 

England. 
4. Cockermouth, Cum- W. R. Wilson (c). 97. 1. 15. 1. 

befland. 
<$. Pontrilas, Hereford- Mrs. St. John A. 85. 6. 10. 1. 

shire. Matthews (p). 

4. Ludford Park, Here- H. J. Bailey (c & p). 0. 2. 23. 1. 

fordshire. 

2. Wales (1828). Earl Cawdor (p). 61. c. d. 

2 4, 1 9, Egersund, Stavanger, K. H. Schaanning 11. 6. 3. 19-21. 
skulls. Norway. (o). 

2 skulls. Southern Germany. Dr. A. Gunther (c). 188i, 1299f. 

i. Bohemia. Lord O. Russell 50. 12. 23. 1. 

(c&p). 

Martes maetes latinorum Barrett-Hamilton. 

1904. Mustela martes latinorum Barrett-Hamilton, Ann. and Mag. Nat. 

Hist., 7th ser., xm, p. 389, May, 1904 (Nurri Mountains, Sardinia). 
1910. Mustela martes latinorum Trouessart, Paune Mamm. d'Europe, 

p. 72. 

Type locality. — Nurri Mountains, Sardinia. 

Geographical distribution. — Mediterranean region (Italy, Sar- 
dinia, Balearic Islands). 

Diagnosis. — Throat patch buff-yellow ; general colour lighter 
than in true martes. 

Colour. — Upper parts a rather light brown, the longer hairs 
between the raw-umber and mars-brown of Bidgway, the 
general effect darker in certain lights, especially on posterior 
half of body ; underfur a pale bluish grey, essentially as in true 
martes, the tips of the hairs ranging from pale wood-brown to 
clear buff; face isabella-colour streaked with whitish hairs, the 
muzzle and lips between mars-brown and prout-brown ; ear 
edged with buffy drab ; tail like back at base, darker at tip ; 
feet and legs washed with dark brown ; throat-patch varying 
considerably, but as a rule approaching buff-yellow. 

Measurements. — Adult male from Porlezza, Como, Italy : 
head and body, 470; tail, 235; hind foot, 94" 6; ear from 
meatus, 42. Adult female from San Cristobal, Minorca, Balearic 
Islands ; head and body, 430 ; tail, 230 ; hind foot, 87 ; ear 
from meatus, 45. For cranial and dental measurements see 
Tables, pp. 376, 378. 

Specimens examined. — Nineteen, from the following localities : — 
Italy : Porlezza, Como, 11 (Ghidini) ; near Rome, 4 ; Milan, 1. 
Sardinia : Nurri Mountains, 1 (type). 
Spain : Mancor, Majorca, Balearic Islands, 1 ; San Cristobal, Minorca, 1. 

Remarks. — The status of this form is very unsatisfactory. So 
far as can be judged from the few specimens examined there is 



374 



CAENIVORA 



an average difference, however, between the Mediterranean 
animal and that occurring north of the Alps. Two of the 
specimens from Porlezza are darker than the others, in this 
respect agreeing with the northern animal. 



1. Italy. (C. Coli.) 

1. Milan. 

1. Nurri Mountains, Sar- 
dinia. 
?. Mancor, Majorca, Bale- 
aric Islands. (Biutort.) 
9. San Cristobal, Minorca. 



G. Barrett-Hamilton 11. 1. 2. 29. 

B. Cavendish Taylor 5. 5. 6. 8. 

E. N. Buxton (p). 95. i. 16. 1. 

(Type of subspecies.) 
0. Thomas (p). 1. 3. 6. 1. 



0. Thomas & B. I. 
Pocock (c & p). 



0. 7. 1. 43. 




MARTES FOINA Erxleben. 

(Synonymy under subspecies.) 

Geographical distribution. — Central and southern Continental 
Europe, from the Atlantic coast eastward, and from the 
Mediterranean to the Baltic. 

Diagnosis. — Third upper premolar with outline of crown 
nearly biconvex, the outer side occasionally 
flattened, the greatest transverse diameter 
at middle ; upper carnassial with inner 
lobe slender, its diameter in line of tooth- 
row equal to only about half greatest 
width of trenchant portion of tooth behind 
middle of crown ; upper molar not so large 
„, f, as in M. martes, its greatest diameter less 

Fiq. 80. than outer length of carnassial, the meta- 

Larger cheek-teeth i of Martes cone less than half as large as paracone; 

martes (re) and M.fmnaOi). .„ . ,.- s . r , . j. ' 

Nat. size. external form as in M. martes, but iur 

of less fine quality ; colour usually more 

greyish or drab than in M. martes and seldom with the rich 

brown tints of the related species, the throat-patch never strongly 

tinged with yellow. Mammse : 4. 

Skull. — The skull resembles that of Martes martes, but may 
usually be distinguished by its greater breadth and less depth. 
Brain-case noticeably wider than high, the general outline when 
viewed from behind nearly as in Mustela erminea and distinctly 
less elevated than in Martes martes. Interorbital region and 
rostrum wider than in Martes martes, and concavity of dorsal 
profile in nasal region much more pronounced. Anteorbital 
foramen usually smaller than in the related species, though in 
the same position. In other respects the skulls of the two 
animals show no tangible differences. 

Teeth. — Except as already pointed out the teeth agree with 
those of Martes martes. 



MARTES 375 

Remarks. — Though readily distinguishable from Martes martes 
by the character of the skull and teeth M. foina is sometimes 
difficult to recognize by external peculiarities alone. Usually 
the colour has a slaty or drab cast that is highly characteristic, 
and the quality of the fur is inferior to that of the pine marten ; 
but I have seen tanned skins of animals killed in winter which 
were impossible to identify with any degree of certainty. 

Two geographical races have been described. 

Maktes foina foina Erxleben. 

1777. [Mustela'] foina Erxleben, Syst. Regni Anim., I, p. 458 (Germany). 
1792. Martes domestica Pinel, Aotes Soo. d'Hist. Nat., Paris, I, p. 55 (Prance). 
1801. M[ustela~] f[oina~] alba Beohstein, Gemeinn. Naturgesoh. Deutsch- 

lands, I, 2nd ed., p. 759 (Thiiringen, Germany). 
1857. Mustela foina Blasius, Saugethiere Deutschlands, p. 217. 
1869. Mustela martes var. fagorum Patio, Paune Vert. Suisse, I, p. 318 

(Sweden) : name wrongly attributed to Linnseus, Syst. Nat., i, 

12th ed., p. 67. 
1910. Mustela foina Trouessart, Paune Mamm. d'Europe, p. 72. 

Type locality. — Germany. 

Geographical distribution. — Range of the species except 
southern Spain. 

Diagnosis. — Longer hairs of back tipped with sepia, the 
general hue of upper parts drab. 

Colour. — Underfur very light grey (about grey No. 10 of 
Ridgway) with an evident buffy cast on distal third of hairs ; 
long hairs ranging from wood-brown to mars-brown, becoming 
darker at tips. As the long hairs nowhere conceal the underfur 
the general effect is a light drab resulting from the blending of 
the two colours, very uniform throughout dorsal surface, though 
usually a little paler on neck and darker on posterior portion of 
back. Tail essentially like body, but darkening toward tip, the 
pencil often blackish. Legs and feet washed with dark brown. 
Ear buffy drab externally, creamy white along rim, the inner 
surface greyish white. Muzzle, lips and chin broccoli-brown. 
Throat-patch buffy white, darkening to cream-buff along outer 
edges. 

Measurements. — Adult male from near St. Gallen, Switzer- 
land : head and body, 453 ; tail, 260 ; hind foot, 85 ; ear from 
meatus, 34. For cranial measurements see Table, p. 377. 

Specimens examined. — Twenty-three, from the following localities : — 

France : Manonville, Meurthe-et-Moselle, 2. 

Germany : Ingelheim, Kheinhessen, 2 ; Nuremberg, Bavaria, 1 
(U.S.N.M.) ; southern Germany, 7 (skulls). 

Switzerland : Geneva, 1 (Mottaz) ; Vallee-de-Joux, Vaud, 1 (Mottaz) ; 
Thayngen, Schaffhausen, 3 (B.M. and U.S.N.M.) ; Aarburg, Aarau, 1 
(U.S.N.M.) ; Oberrich, St. Gallen, 1. 

Spain: Silos, Burgos, 1 ; Dofiasantos, Burgos, 1 (U.S.N.M.). 

Italy : Near Rome, 1. 



376 



CARNIVORA 



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380 



CARNIVORA 



Greece : Mt. Vuno, Cephalonia, 1 (colour normal ; size apparently 
rather small). 

9, juv. Manonville, Meurthe-et- Lord Lilford (p). 95. 9. 5. 1-2. 

Moselle, France. 

5. Ingelheim, Rheinhessen, C. Hilgert (o). 8. 11. 2. 17. 

Germany. 

6. Ingelheim, Rheinhessen. G. Barrett-Hamilton 11. 1. 2. 102. 

7 skulls. South Germany. Dr. A. Gunther (o). 1299, a-g. 

2 6 juv. Schaffhausen, Switzerland. 0. Thomas (p). 2. 8. 4. 22-23. 

(E. H. Zollikofer.) 

S. Oberrich, St. Gallen. 0. Thomas (p). 2. 8. 4. 21. 

(E. H. Zollikofer.) 

S. Silos, Burgos, Spain. Rev. Saturio Gonza- 8. 7. 7. 21. 

lez (c). 

1. Rome, Italy. E. Cavendish Taylor 5. 5. 6. 7. 

(r)- 
1. Mt. Vuno, Cephalonia, J. I. S.Whitaker (p). 8.10.1.9. 

Greece. (0. Mottas.) 
Martes foina mediterranea Barrett-Hamilton. 

1898. Mustela mediterranea Barrett-Hamilton, Ann. and Mag. Nat. Hist., 

7th ser., I, p. 442, June, 1898. 
1910. Mustela foina mediterranea Trouessart, Faune Mamm. d'Europe, 
p. 73. 

Type locality. — Sierra de Jerez, Cadiz, Spain. 

Geographical distribution. — Southern Spain. 

Characters. — Longer hairs of back tipped with a light 
yellowish brown, the general hue of upper parts lighter, yellower 
and less drab than in true foina. 

Measurements. — For cranial and dental measurements see 
Tables, pp. 377, 379. 

Specimens examined. — Two, the type, from the province of Cadiz, and 
a second specimen (mummy) from the Sierra Nevada. 

Remarks. — If the two individuals of Martes foina mediterranea 
are not abnormal in colour the race is rather well characterized. 
I have seen only one skin of true foina that approaches them, a 
specimen from Rome. 

1 juv. Sierra de Jerez, Cadiz, Spain. A Chapman (o & p). 98. 3. 18. 1. 

(Type of subspecies.) 

MARTES BUNITES Bate. 

1899. Mustela foina leucolachnsea Barrett-Hamilton, Ann. and Mag. Nat. 

Hist., 7th ser., iv, p. 383 (Not of Blanford). 
1906. Mustela foina bunites Bate, Proc. Zool. Soc. London, 1905, u, p. 318, 

April 5, 1906. 
1910. Mustela foina bunites Trouessart, Faune Mamm. d'Europe, p. 73. 

Type locality. — Kontopalo, Kania, Crete. 

Geographical distribution. — Crete. 

Diagnosis. — In general like Martes foina, but size not so 



MUSTBLA 381 

large (condylobasal length of skull in adult male not reaching 
80 mm.), rostral portion of skull not so broad, and pale throat- 
patch so much encroached on by brown of surrounding parts, 
that it is occasionally absent. 

Colour. — The colour is lighter and more yellowish (less drab) 
than in Martes foina foina, as the long hairs continue a clear 
wood-brown to tips, producing a general effect more nearly a 
light isabella-colour than drab. The difference is, however, not 
very great. Tail and feet essentially as in M. f. foina, though 
not so dark. Throat-patch greatly reduced, absent in one of the 
seven skins,* and in its most extreme development represented 
by a mere horseshoe-shaped mark, one extremity of which lies in 
front of each fore leg, while the anterior portion broadens out 
to form an irregular patch covering entire throat immediately 
behind interramial region. 

Skull and teeth. — The skull and teeth resemble those of 
Martes foina except for their uniformly smaller size and the 
relatively somewhat less breadth of the rostrum. 

Measurements. — External measurements of type (young-adult 
male) : head and body, 403 ; tail, 255 ; hind foot, 79 ; ear from 
meatus, 39. For cranial and dental measurements see Tables, 
pp. 377, 379. 

Specimens examined. — Seven, all from Crete. 

<5, 2. Kanea, Crete. Miss D. Bate (c). 5. 12. 2. 18-20. 

1. Katharo. Miss D. Bate (c). 5. 12. 2. 21. 
6. Kontopalo. Miss D. Bate (c). 5. 12. 2. 22. 

(Type of species.) 
skull. Crete. Miss D. Bate (c). 5. 12. 2. 37. 

2. Crete. H. 0. Jones, B.N. (c). 99. 6. 13. 2-3. 



Genus MUSTELA Linneeus. 

(Synonymy under sub-genera.) 

Type species. — Mustela erminea Linnaeus. 

Geographical distribution. — Northern hemisphere from the 
Arctic coast south in the Old World to northern Africa and the 
Malay Archipelago, and in America to the Andes ; in Europe 
east to Ireland. 

Characters. — Skull in general resembling that of Martes, but 
rostrum so shortened that distance from orbit to gnathion is less 
than width of rostrum between anteorbital foramina ; auditory 
bulla without meatal tube, its outline variable but never flask- 
shaped ; paroccipital process small and flattened, closely applied to 
posterior margin of bulla ; dental formula : i ! ||, c 1=?, pm ~ 61 

* A small tuft of white hair persists behind each angle of mouth, and 
another in front of each fore leg, the largest of ,these about 15 mm. in 
diameter. 



382 



CARNIVOEA 



m ? * = 34 ; upper carnassial as in Martes, its posterior cusp low 
but well developed, the height of its main cusp about half outer 
border of crown ; upper molar between pyriform and pandurate in 
outline, the constriction evident though not deep, the main axis of 
the crown nearly perpendicular to sagittal line ; lower carnassial 
without metaconid, the posterior heel crossed by a longitudinal 
trenchant ridge ; other teeth essentially as in Martes ; external 
form slender, the muzzle obtuse, the ears low and rounded, the 
legs short, the tail variable in length and in quality of hair, but 
never so bushy as in Martes. 

Remarks. — The genus Mustela, the most widely distributed of 
the strictly terrestrial Mustelinse, is also the richest in species. 
These fall into three main groups, extreme members of which 
might be regarded as generically distinct. The existence of 
species with intermediate characters makes it impossible to 
define these groups by anything more than their average 
differences.* They are therefore here treated as sub-genera. All 
three are represented in Europe. 

KEY TO THE EUROPEAN PORMS OP MUSTELA. 

Brain-case broad, the mastoid width decidedly 
greater than distance between basion and 
palation ; auditory bullae triangular in outline 

(Sub-genus Putorius, Polecats) M. putorius, p. 419. 

Underfur usually bufly grey (Central Europe) M. p. putorius t p. 423. 

Underfur usually yellowish (Southern Spain) M.p. aureolus, p. 425. 

Brain-case narrow, the mastoid width less than 
or about equal to distance between basion and 
palation ; auditory bulte not triangular in 
outline. 
Rostrum flattened above ; inner margins of 
auditory bullae strongly divergent pos- 
teriorly ; tail bushy ; habits semi-aquatic 

(Sub-genus Lutreola, Minks) M. lutreola, p. 415. 

Rostrum convex above ; inner margins of auditory 
bullse nearly parallel ; tail not bushy ; habits 
terrestrial (Sub-genus Mustela, Stoats and 
Weasels). 
Tail with conspicuous black terminal area in- 
cluding more than pencil ; skull with ros- 
trum usually not so wide as interorbital 
region ; condylobasal length of skull never 
less than 43 ■ 4 mm. in adult male, or than 
35 ■ 4 mm. in adult female ; hind foot 
usually more than 43 mm. in males and 
34 mm. in females (Stoats). 
Brown of sides noticeably encroaching on 
yellowish or whitish of underparts, the 
line of demarcation irregular ; upper lip 
always brown M. hibemica, p. 398. 



* At least so far as Lutreola and true Mustela are concerned. Putorius 
appears to be sharply circumscribed, so that it might readily be regarded 
as a distinct genus (see remarks on p. 419). 



MUSTELA 



383 



Brown of sides not encroaching on yellowish 
or whitish of underparts, the line of 
demarcation straight ; upper lip wholly 

or partly white M. erminea, p. 385. 

Eegion extending from front of rostrum to 
and including postorbital processes 
relatively broad and short ; size below 
the maximum for the European races ; 
condylobasal length of skull in adult 
males, 43 • 4 to 48 '6 mm., mandible in 
adult males, 23 to 26 • 4 mm. (Scandi- 
navin Peninsula, except extreme south 

of Sweden) M. t. erminea, p. 387. 

Begion extending from front of rostrum 
to and including postorbital processes 
relatively narrow and long ; size maxi- 
mum for the European races ; condy- 
lobasal length of skull in adult males, 
47 to 52 - 5 mm., mandible in adult 
males, 25 to 30 mm. 
Zygomatic arches broadly spreading 
(ratio of zygomatic breadth to con- 
dylobasal length about 59) ; posterior 
extension of palate unusually short 
and broad (Islands of Islay and Jura, 

Scotland) M. e. ricfiue, p. 397. 

Zygomatic arches moderately spreading 
(ratio of zygomatic breadth to condy- 
lobasal length about 54) ; posterior 
extension of palate not unusually 
short and broad. 
White winter ooat normally assumed ; 
condylobasal length of skull in 
adult male seldom over 50 mm. ; 
in adult female seldom over 
45 mm. ; upper carnassial not 
enlarged and thickened (Conti- 
nental Europe from southern 
Sweden to the Alps and Pyrenees) M. e. sestiva t p. 389. 
White winter coat normally not 
assumed ; condylobasal length of 
skull in adult male often over 
50 mm. ; in adult female often over 
45 mm. ; upper carnassial enlarged 

and thickened (Great Britain) M. *. stabilis, p 390. 

Tail without conspicuous black terminal area, 
though a tuft of black or blackish hairs 
may be present in pencil ; skull with ros- 
trum usually wider than interorbital 
region ; condylobasal length of skull, except 
in specimens from the Mediterranean 
region, often less than 40 mm. in males, and 
34 mm. in females ; hind foot (except in 
M. africana) usually less than 41 mm. in 
males and 25 mm. in females (Weasels). 
Size about as in the smaller stoats, condy- 
lobasal length of skull in adult male, 
48 to 50 mm., mandible in adult male 
about 28 mm., its depth at front of car- 
nassial, 4 • 5 to 5 mm. ; tail nearly one-half 
as long as head and body (africanus 
group). 



384 ■ CAENIVORA 

Brown of sides noticeably encroaching on 
yellowish or whitish of underparts, 
the line of demarcation irregular 

(Malta, Azores) M. africana, 412. 

Brown of sides not encroachingon yellowish 
or whitish of underparts, the line of 

demarcation straight (Crete) M. galinthias, p. 414. 

Size less than in the stoats, condylobasal 
length of skull in adult male, 37 to 
47 mm. ; mandible in adult male not 
more than 24 mm., its depth at front 
of carnassial less than 4 - 5 mm. (true 

Weasels) M. nivalis, p 401. 

Winter pelage often white ; size small, hind 
foot of males, 29 to 34 mm. (Northern 

and central Europe) M. n. nivalis, p. 402. 

Winter pelage very rarely white ; size 

medium or large, hind foot of males 

usually 32 to 41 mm. (Mediterranean 

region). 

Skull of adult male not infrequently 

44 to 46' 6 mm. in condylobasal 

length ; hind foot of adult male 34 to 

41 mm. ; colour rather dark (Central 

Mediterranean region) M. n. botxamda, p. 405. 

Skull of adult male seldom if ever exceed- 
ing 44 mm. in condylobasal length ; 
hind foot of adult male 30 to 37 
mm. ; colour rather light (Iberian 
Peninsula) M. n. iberica, p. 407. 

Sub-genus MUSTELA Linnseus. 

1758. Mustela Linnseus, Syst. Nat., i, 10th ed., p. 45 (Type by tautonymy 

M. erminea Linnseus). 
1829. Arctogale Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierw., i, 

p. 30 (erminea). 
1829. Ictis Kaup, Entw.-Gesch. u. Natiirl. Syst. Europ. Thierw., i, p. 35 

(vulgaris = nivalis). Not of Schinz, 1824-1828. 
1841. Gale Wagner, Schreber's Saugthiere, Suppl., n, p. 234 (vulgaris = 

nivalis). 
1857. Fcetorius Blasius, Saugethiere Deutschlands, p. 219 (part). 
1865. Neogale Gray, Proc. Zooi. Soc, London, p. 114 (brasiliensis, aureo- 

ventris and xanthogenys). 
1871. Mustelina, Bogdanow, Tpjx OOm. Ectcctb. Ka3ancit. ymiBepc. I, Mem. i, 

p. 167. 
1899. Eumustela Acloque, Paune de France, Mammiferes, p. 62 (vulgaris 

and erminea). 
1911. Mustela Thomas, Proc. Zool. Soc. London, p. 138, April, 1911. 

Type species. — Mustela erminea Linnseus. 

Geographical distribution. — Northern hemisphere, south in the 
Old World to northern Africa, and in the New "World to the 
Andes ; in Europe west to Ireland. 

Characters. — Form very slender ; tail not bushy ; fur not 
modified for aquatic life ; skull without noticeably projecting 
mastoid processes ; auditory bullse not triangular, the inner and 
outer borders nearly parallel ; posterior border of pin 3 shearing 



MUSTELA 385 

against anterior border of pm 3 ; other small premolars not 
capable of trenchant action. 

Ben/iarks. — The sub-genus Mustela is the most widely 
distributed group in the genus. It is also the richest in species 
and local races. About seventy of these are now recognized, 
eleven of which occur in Europe. 



MUSTELA ERMINEA LinniBUS. 
(Synonymy under subspecies.) 

Geographical distribution. — Europe from the Arctic coasst 
to the Pyrenees and Alps, and from Great Britain eastward 
into Asia. 

Diagnosis. — Skull with narrow brain-case and slightly pro- 
jecting, scarcely angular mastoid region, the mastoid breadth 
less than distance from basion to palation ; rostral width across 
canines usually less than interorbital width ; auditory bullse much 
longer than broad, nearly parallel sided, rounded posteriorly, 
truncate anteriorly ; form very slender, the tail slender and not 
bushy ; colour brown above, whitish below, the line of demarca- 
tion straight ; a white winter pelage assumed in colder parts of 
range ; tail always with black tip including more than terminal 
pencil ; upper lip always at least partly white ; ear usually with 
a whitish rim. 

External characters. — General form long and slender, the 
legs short, the body cylindrical, the neck long and nearly as 
thick as body, tail considerably longer than outstretched hind 
leg. Ear short but appearing distinctly above fur of head, its 
outline evenly rounded, both outer and inner surface densely 
clothed with short hair ; muzzle rather broad and short, the 
nostril pad sharply denned, entirely naked, separated from upper 
lip by a narrow hairy area. Palm and sole completely furred in 
winter, the tubercles bare in summer ; on both palm and sole 
there is a trilobed, heart-shaped tubercular mass at base of 
median digits and a small round pad at base of inner digit ; on 
palm there is an additional posterior rounded pad near wrist. 
Fur dense and soft, the longest hairs on back about 10 mm. 
in winter, less in summer ; tail rather closely haired, the pencil 
full, usually wider than basal portion of tail, its longest hairs 
about half as long as vertebrae ; fur turning white in winter 
except in warmer parts of range. Mammae : i 4 — 4 = 8. 

Colour. — Upper parts and outer surface of legs yellowish 
brown, usually a little darker along middle of back and on head, 
the underfur and bases of the longer hairs much paler ; under- 
parts, inner surface of legs and upper surface of feet whitish or 
yellowish in strong contrast, the line of demarcation between 

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the two colours straight and definite, extending along middle of 
sides of body and neck and passing a little below ear and eye to 
lower edge of muzzle pad, normally leaving upper lip white ; ear 
brown, the rim whitish ; no dark spot behind angle of mouth ; 
tail with terminal fifth and entire pencil black, elsewhere 
concolor with back except for an ill-defined yellowish or whitish 
median area which occasionally extends along under surface from 
base toward or to black tip. In white winter pelage the entire 
animal is white or whitish (often tinged with yellow), with the 





FlO. 81. 
Mustela ei'minea. Nat. size. 

exception of the black area of the tail, which remains as in the 
dark coat. 

Skull. — In general appearance the skull resembles that of 
Mustela putorius (p. 421). This is especially true of the dorsal 
profile, the form of the palate, rostrum and interorbital region, in 
all of which the differences between the two animals are very slight. 
Breadth of rostrum over canines rarely equal to least interorbital 
width. Position of anteorbital foramen as in M. putorius, but 
orifice relatively larger, the width of plate separating it from 



MUSTELA 387 

orbit distinctly less than that of foramen. Postorbital process 
short, not evidently directed backward. Zygomata more widely 
spreading than in M. putorius, strongly and evenly bowed when 
viewed from the side, the orbital process barely indicated and 
posterior widening absent. Owing to shortness of postorbital 
process and virtual absence of orbital process of zygoma, the 
orbit is less margined with bone than in M. putorius. Brain-case 
longer and narrower than in Mustela putorius, the outline when 
viewed from above elongate ovate not distorted by the presence 
of conspicuously projecting mastoid regions ; sagittal crest slight, 
even in old individuals ; lambdoid crest moderately developed, 
scarcely overhanging, the condyles usually visible from above. 
Floor of brain-case nearly flat, though with the usual median 
ridge and lateral and posterior depressions present. Auditory 
bullae moderately inflated throughout, about three-quarters as 
wide as long, nearly parallel sided, squarely truncate in front, 
rounded behind, the meatus not tubular ; anterior extremity of 
bulla not in contact with hamular, and separated from foramen 
ovale by a broad nearly flat area, the length of which is 
about one-half distance between foramina ; inner extremity of 
glenoid surface marked off by a wide, shallow notch. Postorbital 
region abruptly constricted, short, without indication of the 
neck-like elongation which forms so conspicuous a feature of 
the skull of M. putorius. Mandible with lower border slightly 
convex, its posterior fourth somewhat elevated above general 
outline ; angular process small but better developed than in 
Jlf. putorius. 

Teeth. — Except for their smaller size the teeth are essentially 
like those of Mustela putorius. Enamel of lower canine not 
rugose. Second upper premolar and third lower premolar more 
compressed than in the larger animal, and with base of cusp 
shorter, so that there is a slight but evident concavity or 
flattening at both anterior and posterior base. Upper carnassial 
with inner lobe projecting more forward than in M. putorius, so 
that its anterior border extends nearly to level of that of outer 
margin of tooth. Upper molar and posterior lower molar as in 
Mustela putorius, but m 1 with metacone more reduced, and m 2 
with crown area distinctly smaller than that of heel of carnassial. 



Mustela erminea eeminea Linnseus. 

1758. [Mustela] erminea Linnseus, Syst. Nat., i, 10th ed., p. 46 (Sweden). 
1792. M [ustela] erminea hyberna Kerr, Anim. Kingd., p. 181 (name applied 

to the northern true ermine). 
1816. M[ustela] herminea Oken, Lehrb. d. Naturgesch., in, pt. 2, p. 1026 

(Benaming of erminea). 
1827. [Mustela erminea] $ maculata Billberg, Synopsis Faunse Scandinaviae, 

p. 8 (Scandinavia). 
1857. Fcstorius erminea Blasius, Saugethiere Deutschlands, p. 228 (part). 

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CAENIVOEA 



1877. Putorius erminea Coues, Fur-bearing Animals, p. 109 (part). 
1910. Putorius (Ictis) ermineus Trouessart, Faune Mamm. d'Europe, p. 78 
(part). 

Type locality. — Upsala, Sweden. 

Geographical distribution. — Scandinavian Peninsula, except 
extreme south of Sweden ; eastern limits of range unknown. 

Characters. — Region extending from front of rostrum to 
and including postorbital processes relatively broad and short; 
size below maximum for the European races : condylobasal length 
of skull in adult male, 43 • 4 to 48 ■ 6 mm. ; mandible in adult 
male, 23 to 26 • 4 mm. ; basal portion of tail (in summer pelage) 
usually lighter below than above. 

Measurements. — Hind foot in each of two males from the 
vicinity of Christiania, Norway, 43 mm. ; i