Skip to main content

Full text of "Mammals of Eastern Europe and Northern Asia"

See other formats


S.I. OGNEV te^MN 

MAMMALS OF THE U.S.S.R. 
AND ADJACENT COUNTRIES 

Volume IV 



RODENTS 



Translated from Russian 



Published for the Smithsonian Institution, U.S.A. 

and the National Science Foundation, Washington, D.C. 

by the Israel Program for Scientific Translations 



^' ''/ MAMMALS of the U.S.S.R. 

M^^ and 

ADJACENT COUNTRIES 



(Zveri SSSR i prilezhashchikh stran) 



MAMMALS OF EASTERN EUROPE 

AND 

NORTHERN ASIA 

(Zveri vostochnoi Evropy i severnoi Azii) 



Vol. IV 



Rodents 

(Gryzuny) 



Izdatel'stvo Akademi Nauk SSSR 

Moskva-Leningrad 

1940 



Translated from Russian 



Israel Program for Scientific Translations 
Jerusalem 1966 



TT 65-50016 

Published Pursuant to an Agreement with 

THE SMITHSONIAN INSTITUTION, U.S.A. 

and 

THE NATIONAL SCIENCE FOUNDATION, WASHINGTON, D. C 



Copyright © 1966 

Israel Program for Scientific Translations Ltd 

IPST Cat. No. 1254 



Translated by Jean Salkind 
Edited by Professor O. Theodor 



Printed in Jerusalem by S . Monson 
Binding: Wiener Bindery Ltd. , Jerusalem 

$8.04 

Available from 

U.S. DEPARTMENT OF COMMERCE 

Clearinghouse for Federal Scientific and Technical Information 

Springfield, Va. 22151 



IV/6/3 



CONTENTS „ T^ ,. , 

Russian English 

page page 

Foreword 8 ix 

Order Five 
GLIRES UNNAEUS, RODENTS 

First Suborder, DUPLICIDENTATA 12 2 

Family Lagomyidae, Pikas 13 3 

Genus O c hot on a Link 14 4 

Subgenus Pika Lac^p^de 18 8 

149. Ochotona (Pika) al pina Pallas (1773). 

Altai or Alpine pika 19 9 

309. Ochotona (Pika) alpina alpina 

Pall. (1773). Central Altai pika 31 20 

310. Ochotona (Pika) alpina nitida 

Hollist, (1912). East Altai pika 32 21 

311. Ochotona (Pika) al pi na ch an g aic a 

subs. nov. Hangay pika 33 22 

312. Ochotona (Pika) alpina scorodu- 
movi Scalon (1935). Skorodumov's 

Alpine pika 34 22 

313. Ochotona (Pika) alpina svatoshi 

Turov(1924), Barguzin pika 36 24 

314. Ochotona (Pika) alpina 

cinereofusca Schrenck( 1858). Amur pika 37 25 

150. Ochotona (Pika) hype rbore a Pallas (1811). 

Northern pika 38 26 

315. Ochotona (Pika) hyperborea 

hyperborea Pall. (1811). Northern pika 48 35 

316. Ochotona (Pika) hyperborea 
ferruginea Schrenck (1858), Russet 

Northern pika 49 36 

317. Ochotona (Pika) hyperborea 

cinereoflava Schrenck (1858). 51 38 

Okhotsk Northern pika 

318. Ochotona (Pika) hyperborea 
uralensis Flerov (1927), Northern 

Ural pika 51 38 



52 


39 


55 


42 


56 


43 


62 


48 


63 


49 


65 


52 


76 


61 


n 


62 



R. P. E. P. 

319. Ochotona (Pika) hyperborea 

mantchurica Thos. (1909), Central 

Siberian Northern pika 

Subgenus Ochotona Link 

151. Ochotona (Ochotona) daurica Pall. (1776). 
Daurian pika 

320. Ochotona (Ochotona) daurica 
daurica Pall. (1776). Eastern Daurian pika 

321. Ochotona (Ochotona) daurica 
altaina Thos, (1911). Western Daurian pika 

152. Ochotona (Ochotona) rufescens Gray 
(1842). Afghan pika 

322. Ochotona (Ochotona) rufescens 
regina Thos. (1911). Kopet-Dagh pika 

153. Ochotona (Ochotona) pricei Thomas 
(1911). Price's pika 

323. Ochotona S. (Ochotona) pricei 
Thomas (1911). Mongolian pika or Price's 

pika. 82 67 

324. Ochotona (Ochotona) pricei 
subsp. nov. Price's Semipalatinsk pika. . . . 

Subgenus Conothoa Lyon 

154. Ochotona ( C onotho a) m ac rot is Giinther 
(1875). Large -eared pika or Indian pika 

155. Ochotona (Conothoa) rutila Severtzov 
(1873). Red pika 

Species of the Ochotona pusilla and O. thibetana 
group 

156. Ochotona pusilla Pallas (1768). Small pika 
or chekushka 

325. Ochotona pusilla pusilla Pallas 
(1768). European chekushka 

326. Ochotona pusilla angustifrons 
Argyropulo (1932). Siberian chekushka . . . 

Family Leporidae, Hares 

Genus L e p u s Linnaeus ( 1 758) 

Subgenus Eulagus Gray J. E. (1867). Desert and 
Tolai hare 

157. Lepus (Eulagus) europaeus Pall. (1778) 

327. Lepus (Eulagus) europaeus euro- 
paeus Pall. (1778). European common 

hare 140 121 

328. Lepus (Eulagus) europaeus 
hybridus Pallas (1811), Central Russian 

common hare 142 1 23 



84 


69 


85 


69 


85 


70 


90 


74 


97 


80 


98 


82 


104 


89 


105 


90 


106 


91 


108 


93 


109 


93 


112 


96 



R. P. E. P. 

329. Lepus (Eulagus) europaeus 
tesquorum Ogn. (1923). Steppe common 

hare 147 128 

330. Lepus (Eulagus) europaeus trans- 
sylvanicus Matchie (1901). Roumanian 

or Crimean common hare 150 131 

331. Lepus (Eulagus) europaeus 
caucasicus Ognev (1929). Caucasian 

common hare 153 133 

332. Lepus (Eulagus) europae us c yre nsis 

Satun. (1905). Transcaucasian common hare 155 135 

333. Lepus (Eulagus) europaeus caspicus 

Ehrenberg (1830)-. Caspian common hare. . , 158 138 

158. Lepus (Eulagus) tolai tolai Pall. (1778). 

Tolai hare 161 141 

159. Lepus (Eulagus) tib et anus Waterhouse (1841). 

Desert hare 1 72 149 

334. Lepus (Eulagus) tibetanus lehmanni 

Severtzov (1873). Syr-Darya desert hare. . . , 186 162 

334'. Lepus (Eulagus) tibetanus lehmanni 
natio zaisanicus Satun. (1907). Zaisan 
desert hare 190 165 

335. Lepus (Eulagus) tibetanus que rce rus 

Hollister (1912). Chuya hare 191 166 

336. Lepus (Eulagus) tibetanus subsp. 

Kopet-Dagh desert hare 193 168 

337. Lepus (Eulagus) tibetanus desertor- 

um Ogn. et Heptner (1928). Desert sand hare 195 170 

338. Lepus (Eulagus) t ibe tanus buchari - 

ens is Ogn. (1922). Buchara desert hare 197 172 

339. Lepus (Eulagus) tibetanus pamiren- 

sis Blanford (1875). Pamir desert hare 199 174 

Subgenus Lepus s. str. Linnaeus (1758) 201 175 

160. Lepus timidus Linnaeus (1758). Blue hare 201 176 

340. Lepus timidus ti m i dus Linnaeus (1758). 

Northern blue hare 237 207 

341. Lepus timidus kozhevnikovi Ogn. 

(1929). Central Russian blue hare 240 208 

342. Lepus timidus sibiricorum Johans. 
(1923). West Siberian blue hare 242 

343. Lepus timidus tr ansbaik alicus 

Ognev (1929). Transbaikal blue hare 244 212 

344. Lepus timidus gich iganus J. Allen 

(1903). Gizhiga, orOkhotsk blue hare 247 215 



211 



345. Lepus timidus mordeni Goodwin 
(1933), Maritime territory blue hare 

346. Lepus timidus orii Kuroda (1928). 
Sakhalin blue hare 

347. Lepus timidus kolymensis Ogn. 
(1922). Kolyma blue hare 

348. Lepus timidus begitschevi 
Koljuschev (1936). Taimyr blue hare 

349. Lepus timidus tschuktschorum 
Nordquist (1883). Chukchi blue hare 

Genus Allolagus Ognev 

161. Allolagus mandshuricus Radde (1861). 
Manchurian hare 

Genus Oryctolagus Lilljeborg (1873) 

162. Oryctolagus cuniculus Linn, (1758). 

The rabbit ,, 

Second Suborder. SIMPLICIDENTATA 

Family Pteromyidae, Flying Squirrels 

Genus Pteromys G. Cuvier (1817) 

163. Pteromys volans L. (1758), Common flying 
squirrel 

350. Pteromys volans volans Linn. (1758). 
Common flying squirrel 

351. Pteromys volans ognevi Srogan. 
(1936). Western flying squirrel or Ognev's 
flying squirrel 

352. Pteromys volans gubari Ogn. (1934). 
Western Siberian flying squirrel 

353. Pteromys volans betulinus Serebr. 
(1929), Semipalatinsk flying squirrel 

354. Pteromys volans turovi Ogn, (1929). 
Central Siberian flying squirrel 

355. Pteromys volans athene Thos, (1907). 
Sakhalin flying squirrel 

356. Pteromys volans incanus Miller 
(1918). Yakut flying squirrel 

357. Pteromys volans anadyrensis 
subsp. nov. Anadyr flying squirrel 

358. Pteromys volans arsenjevi Ogn. 
(1934). Ussuri flying squirrel 

Family Sciuridae 

Subfamily Sciurinae, Squirrels 

Genus S c i u r u s Linnaeus ( 1 758) 

Subgenus Sciurus Linnaeus (1758) 



R. P. 


E.P 


249 


217 


251 


219 


254 


221 


255 


222 


258 


225 


259 


226 


261 


226 


271 


236 



272 



239 



281 


246 


284 


248 


286 


249 


288 


252 


306 


268 


308 


270 


310 


271 


311 


272 


314 


274 


316 


276 


319 


278 


321 


281 


323 


282 


325 


284 


327 


285 


327 


286 


329 


287 



R.P. 


E.P. 


329 


287 


384 


333 


386 


336 



389 


338 


391 


340 


393 


342 


395 


344 



164. Sciurus vulgaris Linn. (1758). 

The squirrel 

359. Sciurus vulgaris o g n e v i Migulin 
(1928). Central Russian squirrel 

360. Sciurus vulgaris fedjushini Ogn. 
(1935). Belorussian squirrel 

361. Sciurus vulgaris formosovi Ogn. 
(1935). Northern squirrel, or Formozov's 

squirrel 388 337 

362. Sciurus vulgaris varius Barrett 
Hamilton (1899) ex Kerr (1792). Koka 
squirrel 

363. Sciurus vulgaris ukrainicus 
Migulin. Ukrainian rust-colored squirrel 

364. Sciurus vulgaris kessleri Migulin 
(1928). Kessler's squirrel 

365. Sciurus vulgaris bashkiricus 
Ogn. (1935). Bashkiri an squirrel 

365'. Sciurus vulgaris b ashkiri cus natio 
uralensis Ogn. (1935). Ural Mountain 
squirrel 

366. Sciurus vulgaris martensi Matschie 
(1901). West Siberian squirrel 

367. Sciurus vulgaris exalbidus Pallas 
(1778). Teleut squirrel 

368. Sciurus vulgaris kalbinensis 
Selevin (1934), "Borovushka" or the Kalbin 

squirrel 405 353 

369. Sciurus vulgaris jenissejensis 

Ogn. (1935), Yenisei squirrel 406 354 

370. Sciurus vulgaris altaicus Serebren, 

(1928), Altai squirrel 409 356 

371. Sciurus vulgaris j acute nsis Ogn. 

(1929). Yakut squirrel 410 357 

371'. Sciurus vulgaris jacutensis natio 
anadyrensis Ognev (1929), Anadyr 
squirrel 415 361 

372. Sciurus vulgaris f use oni gric ans 

Dwigubski (1804). Transbaikal squirrel 416 362 

373. Sciurus vulgaris mantchuricus 

Thos. (1909), Manchurian squirrel 417 364 

374. Sciurus vulgaris rupestris Thos. 

(1907). Sakhalin-Amur squirrel 419 366 

Subgenus T e n e s Thos 421 367 

165. Sciurus (Tenes) persicus anomalus 



397 


345 


397 


346 


401 


348 



R. P. E. P. 

Gmel, (1778). Caucasian squirrel 422 368 

Subfamily Spermophilopsinae 432 377 

Genus S perm ophilopsis W. Blasius (1884) 433 378 

166. S pe rmophil opsis leptodactylus Licht. 

(1923). Long- clawed ground squirrel 435 380 

375. Spermophilopsis leptodactylus 
leptodactylus Licht. (1823) 452 394 

376. Spermophilopsis leptodactylus 
schumakovi Satunin (1908). Kushka 

or Shumakov's ground squirrel 454 395 

Subfamily Marmotinae. Chipmunks, Ground Squirrels and 

Marmots 

Genus Eut ami as Trouessart (1880) 456 397 

167. Eutamias sibiricus Laxm. (1769). 

The Chipmunk 458 398 

377. Eutamias sibiricus sibiricus 
Laxmann (1769). Central Siberian 

chipmunk 483 421 

378. Eutamias sibiricus striatus Pallas 
(1778). West Siberian or European 
chipmunk 

3 79. Eutamias sibiricus jacutensis 
Ognev (1936). Yakut chipmunk 

380. Eutamias sibiricus orientalis 
Bonhote (1898). Ussuri chipmunk 

381. Eutamias sibiricus lineatus 
Siebold (1826). Amur chipmunk 

SUPPLEMENT TO VOLUME THREE 

Tables 1-61 
Maps I -IX 



485 


422 


486 


423 


487 


424 


488 


425 


490 


427 



FOREWORD 

The fourth volume of "Mammals of the U. S. S. R. " deals with the Order 
Glires (Rodents). 

Numerous books have recently been published on the rodents of the 
U. S. S. R. J which contain extensive and detailed surveys of the ecology and 
geographical distribution of some of the best -known and most interesting 
representatives of the order, e. g. , squirrels, spermophiles, the blue 
hare and the European common hare. 

This book is as broad in scope as the preceding volumes, and can be 
used as a reference book. I have endeavored to use all the available 
taxonomic, zoogeographical and biological literature. Some of the surveys 
are rather extensive and I hope the reader will not mind this. 

The fourth volume contains several innovations. Species are treated in 
detail; subspecies are described separately. This results in greater 
accuracy of the descriptions. Following the suggestion of critics, 
dichotomous keys have been replaced by Swedish keys (instead of the more 
complicated English keys). The amount of material used has been indicated 
in each case. 

The illustrations were prepared with the greatest care. Original 
drawings of skulls were made with a pantograph by S. Yu. Kostylev, 
Scientific Associate of the Moscow State Zoological Museum. Many 
photographs taken by myself are given, as well as half-tone plates by Prof. 
A.N. Formozov, A.N. Komarov, and K. K. Flerov. The color plates 
[reproduced in black and white in this translation] were made by V. A. 
Vatagin. 

Diagrams of cranial measurements of pikas, hares and sperm ophiles 
are given for better understanding. Such diagrams are more instructive 
than descriptions. 

The successful completion of this extensive work is largely due to the 
generous support of V. M. Molotov, Chairman of the Council of People's 
Commissars. 

I wish to record my gratitude to the Scientific Research Institute of 
Zoology of the Moscow State University for providing most favorable 
conditions for work. 

It is hoped that this fourth volume will be received as well as the 
preceding volumes. 

Moscow, 25 November 1936 



10 




11 GLIRES LINNAEUS. 

RODENTS 



ORDER FIVE 
GLIRES LINNAEUS (RODENTIA VICQ. D'AZYR) 

RODENTS 

Rodents are the largest mammalian order, consisting of more than 2800 
species. They are chiefly small pentadactyle, unguiculate mammals with a 
plantigrade or semiplantigrade gait. Their dentition is very characteristic. 
There is only one pair of incisors in the upper and lower jaw. Only 
Lagomorpha and Ochotonidae have a second pair of small incisors behind 
the large pair of upper incisors. The incisors are chisel-shaped, rootless, 
and grow throughout life. Under normal conditions their growth is 
compensated by their permanent wear. Between the incisors and premolars 
there is always a large space called diastema which is caused by the 
permanent absence of canines. The surface of the molars may be plane (as 
in Microtinae), or bear blunt tubercles. The primitive type of molars is 
brachydont or bunodont. They later become hypselodont, and their surface 
becomes lophodont. 

The teeth are adapted to gnawing and grinding a vegetarian diet, including 
hard matter such as wood and bark. The mandibular condyle is generally 
elongate in a longitudinal direction. The tympanic bones form the hollow or 
spongiose wall of the auditory bullae (bullae osseae). The orbit 
communicates with the temporal fossa. The infraorbital canal is often 
very long, as part of the masseter muscle runs through it. The 
premaxillaries frequently have posterior proceases which reach the frontal 
bones, partly separating them from the maxillaries. The clavicles are 
markedly reduced. The tongue has only 3 circumvallate papillae. Intestine 
adapted to vegetarian diet. Stomach relatively simple, intestine very long, 
and cecum well developed. Brain macrostatic and hemispheres usually 
smooth, not covering cerebellum. 
12 Testes lying either permanently in the abdominal cavity or in an inguinal 
sac, rarely in the scrotum. Uterus double (uterus duplex). Placenta 
discoidal and deciduate. 

The external structure of rodents is adapted to greatly variable conditions 
of life. Some rodents are arboreal. Others are burrowing and are adapted 
to underground life in a varying degree. Some rodents are aquatic, and 
are adapted to prolonged swimming and diving. Some species of the order 
are excellent runners, being equipped with very long and specialized 
posterior limbs. Rodents also include a few "flying" species. Skin folds 
between the fore- and hind limbs form a so-called parachuting membrane 
thus enabling them to glide (e.g., flying squirrels). 

Rodent distribution is worldwide. They live in different habitats which 
are reflected in their biological adaptations: highlands, plateaus, steppes. 



deserts, tundras, and forest zones. Aquatic species occur in rivers and 
lakes. 

Rodents are divided into two suborders, distinguished according to the 
following key: 

2 

1 (2), Incisois — , i. e. , an additional small pair of incisors behind the pair of large incisors in the 

upper jaw. Distance between rows of upper molars larger than that between rows of lower molars. Optic 
foramina fused. Cecum with spiral fold. 

Suborder Duplicidentata. 

2 (1). Incisors — , i. e. , one incisor on each side of upper jaw, an additional pair of small incisors 

absent. Distance between rows of upper molars either smaller or equal to that between rows of lower molars. 
Optic foramina separate. Cecum without spiral fold. 

Suborder Simplicidentata. 



First Suborder. DUPLICIDENTATA 

A second pair of smaller incisors present behind the first pair of large 

upper incisors (i - ). Incisors entirely covered with a layer of enamel 

which is very thin on the posterior surface but thick on the anterior surface, 
forming the characteristic cutting edge of the teeth. 

Number of molars varying between - and - . Number of premolars 

constant: "' •'^ ; number of molars varying between - in Leporidae to — in 

PaPi 3 3 

Ochotona. In some extinct Ochotonidae, e.g., Prolagus Pom. (lower 

2 

European Miocene), the number of molars was reduced to - . 

Molars rootless, hypselodont with deep enamel folds on crown; cement 
13 embedded between the enamel folds. F. Major showed that the specialized 
hypselodont teeth of Duplicidentata evolved from primitive bunodont and 
brachydont teeth. The elevation of the crown of the molars in the upper jaw 
is less marked on the inner than on the outer side. As a result, the crowns 
of the upper molars are beveled towards the outside, and in the opposite 
direction in the lower molars. The distance between the rows of upper 
molars is greater than that between the lower rows. Duplicidentata grind 
their food by lateral movements of the rows of molars . The lower rows of molars 
are laterally surrounded by the upper ones. 

The glenoid fossae for the mandibular condyles of Duplicidentata are 
therefore relatively wide transversely. This facilitates lateral movements 
of the mandibula and of the molars. Both halves of the mandibula are 
rigidly connected by the symphysis. 

The optic foramina are fused, and the infraorbital foramina are narrow 
and small. The bony palate is strongly reduced to a narrow transverse 
bridge behind the large palatine foramina. This bridge of the bony palate 
consists mainly of the paired maxillary processes (processus palatini ossis 
maxillaris) in hares. In pikas the greater part of this bony bridge consists 
of the palatines. The maxillary palatine processes are small, narrow, not 
joined with each other in all pikas. They form the anterior (frontal) 
margin of the bony palate. In primitive species of hares the bony palate 



is broader than in more specialized ones. The clavicles may be 
rudimentary or well developed. 

The soft palate bears many folds. The long cecum is equipped with a 
spiral fold. 

Os penis little developed; tip of penis directed backwards. 

All the characters mentioned above prove that the Duplicidentata are 
less specialized and more primitive than the Simplicidentata. 



KEY TO FAMIUES OF DUPUCIDENTATA 

1 (2). Tail inconspicuous; both pairs of legs of about equal length. Nasals widening anteriorly, 

5 
Postorbital processes absent. Walls of bullae osseae spongiose. Molars — . — 

Lagomyidae (Ochotonidae) . 

2 (1), Tail short but well developed; anterior legs about half as long as the posterior ones. Nasals not 
widened anteriorly, Postorbital processes present. Walls of bullae osseae solid, i. e. , not spongiose. Molars 

6 

— .— Leporidae, 



Family LAGOMYIDAE, PIKAS 

1866, Lagomyidae, Lilljeborg, W. , Systematisk ofversigt af de gnagande daggdjuren, Glires. 

Nova Acta Soc. Scienc. , Upsala, Vol. VI. 
14 1897. Ochotonidae, Thomas C, Types in Natural History and Nomenclature of Rodents, Science, 

VI, pp. 485-487. 
1897. Lagidae Schultze Erw., Helios XIV, S. 82; Catalogus Mammalium europaeorum, Ztschr. f. 

Naturwissensch., Bd. 73, 1900, S. 209. 
1910. Ochotoninae (subg. ). , Trouessart E. L. , Faune de Mammiferes d'Europe, p. 213. 

Ochotonidae are characterized by short, rounded ears and short 
posterior legs which are little longer than the anterior legs. Tail strongly 
reduced. 

Nasals widening anteriorly; postorbital processes absent. Lateral parts 
of maxillae not cribriform. Skull relatively low, usually flattened. The 
skull is convex dorsally only in some species of the family. It seems 
much flatter than that of hares in general. The bullae osseae are large, 
elongate, obliquely directed inwards towards the median line of the skull. 
The bony walls of the bullae osseae are spongiose. 

Clavicles well developed. Pubic symphysis absent. 

Dentition: i-;c-; p-; m- (or -)= (26 or 24). 

The family Lagomyidae contains only the genus Ochotona Gray which 
is divided into the 3 following subgenera: Ochotona s. str. , Pika 
Lac6p^de andConothoa Lyon*. 



Pikas of the pusil 1 a - tibe tan a group apparently form an additional subgenus. Ochotona kozlovi 
Biichner constitutes a sharply differentiated subgenus. Ochotona thomasi Argyropulo also probably 
represents a separate subgenus. This species was described by A. I. Argyropulo (in litt. ) from a specimen 
from the vicinity of Lake Alyk-Nor (Northeast Tibet) which was captured by the expedition of P. K. Kozlov 
on 31 May 1901. This specimen differs sharply from all other pikas by its well proportioned, narrow and 
unusually elongate skull. I know of only one skull of this species in the ZMAN ["Zoologicheskii muzei 
Akademii Nauk" (Zoological Museum of the Academy of Sciences)] collection. 



GEOGRAPHICAL DISTRIBUTION. Pikas are distributed from Eastern 
Europe (from the Trans-Volga region) throughout Eurasia. Their range 
extends from the northern limit of the Arctic tundras, through the Boreal 
region, to the mountainous countries of southern Turkmenia, northern 
Iran, Seistan, the Himalayas, Kansu, and Szechwan. They are also 
sporadically distributed in the Nearctic region, from the Tundra-Boreal 
regions of Alaska to California and Colorado (A. Howell, Revision of the 
American Pikas, U.S. Department of Agriculture, Washington. 1924). 



Genus Go hot ona Link 

1773, Lepus Pallas P., Reise durch verschiedene Provinzen des Russischen Reichs, Bd. II, Anhang, S. 

701-702 (partim,'), Lepus Linnaeus, 1758. 
1795. Ochotona Link H. F, , Beitxage zur Naturgeschichte, Bd. I, Theil 2, S. 74 (type Ochotona minor 

Link. = Lepus ogotona Pall. =Lepus dauuricus Pallas). 

1799. Pika Lacepfede Bern. G. E. , Tableau des divisions des Mammifferes, Paris, p. 9(typePika 
alpinus =Lepus alpinus Pallas). 

1800. Lagomys CuvierG., Lemons d'anatomie comparee, Vol. 1, tab. I (Lagom ys Storr, Prodromus 
Methodi Mamm. , 1780, pp. 34-40; this description was apparently made according to an Arctomys 
specimen). 

1867. O gotom a Gray I. E. , Notes on the Skulls of Hares and Pikas in the British Museum, Annals and 
Magazine of Natur. History, Vol. XX, p. 220 (type Ochotoma Pallasii=La gom ys ogotona 
Waterhouse, Lepus ogotona Pallas). 

1904. Conothoa (subgenus.') Lyon M. W. , Classification of the Hares and Their Allies, Smiths. 
1^ Miscellan. Collections, Vol. XLV (I), parts 3-4, p. 438 (type Ochoton a roy 1 e i Ogilby). 

The species of the genus Ochotona have the general characters of the 
family Lagomyidae. Additional characters are as follows: trunk compact, 
rounded, tailless, limbs short, posterior legs only little longer than the 
anterior ones. Paws very furry. Forefeet pentadactyle, hind feet 
tetradactyle. All digits with short, sharp, curved claws. The underside 
of the tip of each digit has a bare callus in summer fur. Ears of moderate 
length, shorter than those of Leporidae, rounded and furry on both sides. 
Eyes small, compared to those of hares. Fur dense, soft, often silky in 
winter. Color of summer and winter fur usually sharply different. Teats 
4 or 6, distributed as follows: p 1-1 (or 2-2); i 1-1*. 

Rostrum of flattened skull narrow, this being a reliable difference 
between the genus Ochotona and the Leporidae. Bony palate shortened. 
Zygomatic bones with backward-directed, pointed processes. The 
ascending part of the mandibula has a strongly developed condyle. The 
anterior surface of the ascending part has no longitudinal fossa such as 
observed in Leporidae. 

The teeth of Ochotona have some typical characters which distinguish 
them clearly from those of hares. The cutting edge of the upper anterior 
pair of incisors has a deep, acutely angular notch. The edge of each tooth 
seems divided into 2 pointed processes. The inner point is much narrower 
and sharper than the outer. In Leporidae the cutting edge of the upper 
anterior pair of incisors also has an angular notch, but the edge of each 
tooth seems divided into 2 rounded processes and the inner process is only 
little smaller than the outer. The posterior lower molar is simple, with a 



* [p= pectoral, i = inguinal. ] 



single enamel cap. In Leporidae it has a double enamel cap. The structure 
of the second upper molar (m^) differs sharply from that of the third pre- 
molar {p3). In Leporidae these teeth are similar. 

The third upper molar is absent as a rule; in Leporidae it is always 
present. 

Type species: Lepus dauuricus Pallas (1773). 

THE GEOGRAPHICAL DISTRIBUTION of the genus is given in the 
description of the family. 

KEY TO SPECIES OF GENUS OCHOTONA 

1 (6). Underside of vomer exposed; lateral margins of premaxillaries not joined under vomer; palatine 
and incisor foramina fused. 

2 (3). Interorbital region of adult skull relatively broad (5. 2-6 mm) and flattened. Summer fur, various 
shades of grayish brown with an ocher tinge on back. Winter fur fluffy, pale grayish on back with a straw- 
yellow tinge. 

Ochotona m a c r o t i s Giinther, Large - eared pika, 

3 (2). Interorbital region of adult skull relatively narrow and convex in profile. 

4(5). Length of skull not over 47 mm. Vibrissae 40-55 mm long; most vibrissae white, few very dark- 
1 6 brown. Ears with well marked pale margins. Color of summer fur on back pale yellowish straw- gray to 
duller and darker straw- gray. Belly pure white. Neck without a broad pale collar. 

Ochotona daurica Pall. Daurian pika. 

5 (4). Length of skull in adults usually 46 mm to 53 mm. Most vibrissae dark brown; longest 
vibrissae with long white tips, 45 to 55 mm long. Ears without well marked pale margins. Summer fur light 
gray on back with straw- yellow tinge and rust- brown hair tips. Belly dirty-grayish white, often with a 
yellowish tinge. Neck with a broad pale ash-gray collar; an indistinctly marked rust-red band behind the 
collar. 

Ochotona rufescens Gray. Reddish pika. 

6 (1). Position of vomer in relation to edges of premaxillaries, and structure of palatine and incisor 
foramina different, 

7 (12). Vomer ventrally covered by premaxillaries, which join each other and separate palatine and 
incisor foramina completely. 

8 (9). Summer fur ocher-rust-gray on back with rusty tinge. Winter fur straw- gray on back with 
a pinkish rust tinge. 

Soles white, sometimes with straw-grayish shades. Pale margins of ears very indistinct. 

Ochotona pricei Thos. Price's pika. 

9 (8). Color of summer fur on back light yellowish ocher to saturated rust- ocher or rust-brown, even 
black-brown-ocher. Winter fur differs sharply from summer fur in its gray or brownish gray color, 
frequently speckled with blackish hair. 

Soles black or dark brown, sometimes grayish white or gray in small forms. Pale margins of ears 
quite well marked. 

10 (11). Skull large, more than 43mm long in fully grown specimens, and may reach 57mm in old 
ones. Rostrum long; upper diastema usually much longer than upper row of molars, and longer than 
distance between inner margins of alveoli of last upper molars*. 

Zygomatic width more than 22 mm; diastema longer than 9 mm. Hind foot (without claws) more than 
30 mm. 

Soles black or black-brown, gray in winter, 

Ochotona alpina Pall. Altai pika. 



These features are not very well marked in small subspecies of O. alpina, viz. O.a. svatoschi, 
O. a. cinere of use a. However, the diastema was always slightly longer than the upper row of 
molars and the distance between the inner margins of the alveoli of the last upper molars in specimens 
examined. 



PI ATE L 




Red pika (Ochotona rutilaSev.) 



Drawing by V. A. Vatagin 




Altai pika (Ochotona alpinaPall. ) 



Drawing by V. A, Vatagin 



17 



11 (10). Skull much smaller, less than 43 mm long in fully grown specimens. Rostrum relatively short ; 
upper diastema as long as row of upper molars and distance between inner margins of alveoli of last upper 
molars. Zygomatic width less than 22 mm; upper diastema less than 9 mm long. Hind foot (without claws) 
less than 30 mm long. Soles grayish brown or gray; whitish gray in winter. 

Ochotona hyperborea Pall. Northern pika. 

12 (7). Vomer not completely covered ventrally by edges of premaxillaries which are not narrowly 
connected, being separated by a narrow slit; the premaxillaries form a bridge between palatine and incisor 
foramina. 

13 (14). Anterior third of nasals with marked spoon-shaped widening. Rostrum long. Upper diastema 
much longer than row of upper molars. 

Vibrissae 94 mm long. 

Summer fur on back rich rust-red; winter fur on back ash-gray. 

Ochotona rutila Severtz. Red pika. 

14 (13). Nasals very little widened anteriorly. Rostrum very short; upper diastema as long as row of 
upper molars. 

Vibrissae short, not longer than 45 mm. 

Summer fur on back deep grayish brown with more or less distinct straw-yellow mottling. Winter 
fur similar, generaly slightly paler. 

Ochotona pusilla Pall. Small pika, or chekushka. 





FIGURE 1. Skull measurements in pikas 
(Ochotona) (dorsal) 

1 — length of skull; 2 — length of nasals; 
3 — zygomatic width; 4 — interorbital 
distance; 5 — width of skull behind zygo- 
matic arches; 6 — width at level of audi- 
tory meatus; 7 — length of suture between 
front als. 



FIGURE 2. Skull measurements in pikas 
(ventral) 

1 — basal length of skull; 2 — condylo- 
basal length; 3 — length of facial 
region; 4 — length of brain case; 5 — 
length of palatine foramen; 6 — dias- 
tema; 7 — length of row of upper 
molars. 



Subgenera of the genus Ochotona 

O. Thomas (The Duke of Bedford's Zoological Exploration in East Asia. 
On Mammals from the Provinces of Shansi and Shensi, North China, 
Proceedings of the Zoological Society of London, p. 982, 1908) proposed the 
following subgenera of the genus Ochotona: 

1) Ochotona Link, 1795 (=C o n o t h o a Lyon). 

Type species: O. daurica Pall. Palatine and incisor foramina fused 
and exposed. 

2) Pika Lacepede, 1799 (=L a go m y s C. Cuvier). 

Type species: O. alpina Pall, nee Storr. , 1780. Palatine and 
incisor foramina separate. Skull flattened in profile. 

3) OgotomaGray, 1867. Type species O . pallasi Gray( = pricei 
Thos. ). Palatine and incisor foramina separate. Frontal region of skull 
markedly convex. 

This classification is based solely on the structure of the palate. This 
character is excellent for the differentiation of species, but is of little use 
for subgenera. Closely related pikas such as Ochotona macrotis 
Glinth. and O. rutila Severtz. which were considered as one species by 
many authors would have been ascribed to different subgenera according to 
the structure of the palate. More accurately, O. rutila Severtz. would 
not belong to the above-mentioned taxon, as its palatine structure is 
intermediate: incisor and palatine foramina are connected by a very narrow 
slit which is sometimes almost occluded. According to its palatine 
structure, Ochotona erythrotis Biichn. would belong to the subgenus 
Pika and would thus be separated from O. roylei Ogilby, with which 
it is very closely related according to all external characters. Ochotona 
pricei Thos. is very closely related to O . daurica Pall, according 
to the structure of the palate, but belongs to a different subgenus. 
Ochotona pusilla Pall, is generally not included in the subgeneric 
system of O. Thomas. 

The division of genus Ochotona into subgenera proposed by O. 
Thomas is thus unsatisfactory in our opinion. 

In his monograph "Ochotonidae (Lagomyidae) SSSR" (Ochotonidae 
(Lagomidae) of the U. S. S. R. (1936)) A.I. Argyropulo offers a much better 
division in my opinion. He divides pikas into the following groups. 

1) Daurica-curzoniae group including O . daurica, curzoniae 
ladacensis, rufescens, pricei. 

2) Roylei group including O . roylei, macrotis, rutila, 
erythrotis, gloveri, wollastoni, nubrica. 

3) Alpina group including O . alpina, hyperborea, princeps, 
schisticeps, collaris. 

4) Pusilla group including O . pusilla, tibetana. 

5) Kozlovi group, considered by A.I. Argyropulo as a separate new 
subgenus. 

Most of these groups can be designated by the former subgeneric names 
to which a new interpretation will have to be given (vide infra). 



Subgenus Pika Lacepede 

1799. Pika Lac^p^de B. G. E. Tableau des divisions des Mammiferes, Paris, p. 9. 

8 



Skull strongly flattened in the interorbital and frontal region; frontal 
fenestrae as a rule absent. Interorbital space wide. Orbits relatively 
small. Vomer in all species of this subgenus covered ventrally by the 
margins of the premaxillaries. Palatine and incisor foramina separate. 

Winter coloration dark brown, rust-brown or dirty-gray. Pads at 
tips of digits not well developed. Vibrissae relatively short, not longer 
than 70 mm. 

Scree -inhabiting, semisylvan and sylvan species distributed in the tundra 
to the shores of the Arctic Ocean. 
19 Type species: Pika alpinus = Lepus alpinu^ Pall. 

The following species also belong to this subgenus: Ochotona 
hyperborea Pall., O. princeps Richardson, O. schisticeps 
Merriam, O. collaris Nelson. 

GEOGRAPHICAL DISTRIBUTION. Tundra and mountain -forest zone of 
North Asia. All North American (Nearctic) pikas belong to this subgenus. 

149. Ochotona (Pika) alpina Pallas (1773). Altai or Alpine 

pika 

1773, Lepus alpinus Pallas P. , Reise durch verschiedene Provinzen des Russischen Reichs, II. S. 701. 
NAMES: Altai pika, mouse hare, cony (English) ; Der sibirische oder Alpen-Pfeifhase (German), 
LOCAL NAMES: pol'skaya koshechka (Polish cat) (in many localities in Siberia e. g. , in the Altai); 
ogoton (Russian, on the Lena); dikaya koshechka (wdld cat) (Russian, in Siberia); shadak or shaidak 
(mountain-taiga region of the Minusinsk Territory); kamennaya koshka (stone cat); pishchukha (pika); 
piskun (squeaker) or senostavets (haybox) (Russian); dzhekulak (Kazakh, Kirghiz, Tatar); tiin-pishkan 
(Kirghiz); agas, chiss, daas-kila (living in screes) (Yakut); tachergana (Buryat); pika (Evenki, Tungus); 
Ogotona, or kuio (Mongolian); snikan (Birar Evenki-Tungus); chaptyrra (Evenki, Tungus on the Vitim); 
fulman (Ostyak, Khanty). 

PRINCIPAL FIGURES: 1) Pallas P,S, Reise durch verschiedene Provinzen des Russischen Rgichs, II, 
1773, S. 701, Taf. II, B. (half-tone plate of entire animal); 2) Pallas P, S. , Novae species Quadrupedum 
e Glirium ordine, 1778, pp. 45-49, tab. II. (entire animal); tab. IV A, 13 (A), 13 (B) (skull, lateral and 
anterior views); 3) Schreberl, Ch. , DieSaugethiere, Erlangen, IV, 1792, S. 910-914, Taf. CCXXXVIII 
(entire animal, in color); 4) Dvigubskii, I. Opyt estestvennoi istorii vsekh zhivotnykh Rossiiskoi imperii 
(Natural History of the Animals of the Russian Empire), 1829, plate 26 (half-tone plate of entire animal); 5) 
Simashko, Yu. , Russkay a fauna (Russian Fauna), II, 1851, p. 791-793, plate 63, fig. 2 (entire animal, 
color plate); 6) Brandt, I. F. , Untersuch, iiber die craniolog. Entwick. und Classification der Nager, 1855, 
PI. XI, Fig. 11-20 (skull and teeth); 7) Schrenck L, Reisen und Forschungen im Amur Lande, I, 1858, 
S. 150, PI. VIII, Fig. 2 (Lagomys hyperboreus var. cinereofusca = Ochotona alpina cinereofusca (entire 
animal, color plate); 8)Brehm, A., Zhizn' zhivotnykh (Animal Life), II, 1874, p. 262 (half-tone plate); 
9) Fokht, K. , Mlekopitayushchie (Mammals), 1885, p. 402 (half-tone plate of entire animal); 10) 
Brehm, A., Zhizn' zhivotnykh (Animal Life), II, 1893, p. 675 (half-tone plate); 11) Tullberg, T. , 
Ueber das System der Nagethiere, Upsala, 1899, Tab. I, Fig. 22-27 (structure of internal organs); 12) Gaake, 
v., Zhivotnyi mir (The Animal World), Vol, II, 1902, p. 33 (half-tone plate); 13) Vinogradov, B. S. , 
Mlekopitayushchie SSSR (Mammals of the U. S. S, R. ), I, 1933, p, 81, fig, 74 (skull, dorsal), fig, 75 (D) 
(incisor foramina of skull) ; 14) Sbomik: "Zveri Tadzhikistana" (Animals of Tadzhikistan), Trudy Tadzhikskoi 
bazy Akademii Nauk, 1935; Pavlovskii, E. N. , Mlekopitayushchie kak vrediteli (Mammals as Pests), p, 54, 
fig, 23 (section through body wall of pika infested with Oestrid larvae). 
MATERIAL EXAMINED: 127 pelts and skulls. 

DIAGNOSIS. Skull large, length over 45 mm in fully grown and old 
specimens, 57 mm in very large specimens. Interorbital space flattened 
and broad in adult and old specimens, 5 to 6.3mm wide. Anterior third 
of nasals without spoon- shaped widening. Lateral contours of nasals 
narrowing posteriorly, their posterior end rounded. Frontals without 



* Species and subsf>ecies are numbered separately in this volume. 



fenestrae. Vomer ventrally covered by margins of premaxillaries; 
palatine and incisor foramina separate. Most vibrissae black or dark 
20 brown, 60 to 70 mm long. Soles black or very dark brown. Ears with 
well marked pale margins. Coloration of summer fur in different 
subspecies light yellowish gray-ocher (on back) to saturated rust-ocher, 
sometimes with very dark brown-ocher tones. Winter fur sharply 
different from summer fur in its gray or brownish gray color, streaked with 
blackish hair. Completely melanistic individuals occur frequently. 

Body length may reach 250 mm (usually about 200 mm); hind foot may 
be 35 mm long (usually about 30mm). 





FIGURES. Skull of Ochotona alpina alpinaPall., 
dorsal. Specimen No. M. 327018 Dec. 18, 1929. Southern 
Altai, near Argat, Katon-Karagai. S. I, Ognev's collection 
Drawing by Yu. A. Kostylev 



FIGURE 4. Skull of Ochotona alpina 
alpina Pall,, ventral (same specimen as 
in Figure 3) 




FIGURES. Skull of Och oton a alpina alpina Pall., lateral (same specimen as in Figure 3) 



10 



21 ADDITIONAL DESCRIPTION. Skull of Altai pika large, flattened; 
interorbital space broad, its margins without lateral facets. Anterior 
and posterior limits of upper contour of interorbital space have approximately 
the same curvature. Posterior quarter of nasal bones flattened but not 
depressed; nasals rounded posteriorly. Sagittal crest of old individuals 
well marked in occipital region, and in posterior part of parietals. 
Mandibular condyle massive and broad (Figures 3-6). 




FIGURE 6. Mandibula of O c h o t o n a alpina alpinaPall. , 
(No. M.3270, Figure 3) 

Age variation in skull structure well marked. Skull of young pikas 
convex in frontal region. Their skull is therefore not as flat as the skull 
typical of adult and old individuals. The parietals are also more inflated 
in young pikas. The posterior region of the skull in young Ochotona 
alpina is therefore more ovoid, and the interorbital space is broader 
than in fully grown individuals. The bullae osseae of young specimens 
not as long but more rounded and shorter. The short rostrum is also 
characteristic for the young skull. 




FIGURE 7. Ochotona alpina Pall. , running 

Drawing by K. K. Flerov. 



22 The skull of the Altai pika is characterized by the structure of the 

vomer and the premaxillary processes. The vomer is completely covered 
ventrally in most specimens by the margins of the premaxillaries; the 

palatine and incisor foramina are 
therefore separate. This character is, 
however, variable. The margins of 
the premaxillaries sometimes do not 
join in young specimens and a narrow 
slit is left, through which the vomer 
is visible. 

COLOR. (Plate I) The summer 
fur varies significantly in different 
subspecies. The general color of the 
back is dull, yellowish ocher-gray 
(tawny-olive, Ridgway, plate XXIX), 
darkened by black or dark brown 
hairtips in specimens from the Narym 
range. The flanks of these specimens 
have a marked rust -red tinge (pinkish 
cinnamon, Ridgway, pi. XXIX), belly 
pale, yellowish ocher, intermediate 
between pinkish cinnamon and 
cinnamon-buff (Ridgway, pi. XXIX) 
although lighter. 

The color of pikas from the Chuya 
Alps (Chegan-Burgazy -O . alpina 
nit i da Hollister) is more intense, of 
a dark brown shade (intermediate 
between mikado-brown and snuff-brown, 
Ridgway, pi. XXIX). Flanks colored 
a brighter rust-brown, close to sayal- 
brownand cinnamon (Ridgway, pi. XXIX). 
Many pikas from Teletskoe are even 
darker. 

Summer color of the back is intense 
chestnut-brown; the middle of back and 
head quite black; rusty and ocher tones 
well marked on flanks (sayal-brown- 
cinnamon). 
Ochotona alpina svatoshi Turov from the Barguzin Range is 
vividly colored. General tone of summer fur rich rust-ocher on back (sayal- 
brown, Ridgway, 1912) with an orange tinge (orange-cinnamon, Ridgway, 
pi. XXIX). Color of flanks almost the same as that of back. 

In paler specimens of the Altai pika winter fur is dirty-ash-gray with a 
marked black tinge caused by the very dark brown hair tips. Lower part 
of back with slight rusty tinge; flanks with clearly marked pale rust- 
yellowish tinge, intermediate between cinnamon and cinnamon-buff, 
(Ridgway, 1912). These pikas are widely distributed in the southern Altai 
(Narym Range), but there are also specimens with definite dull brown 
shades in their gray winter fur (sayal-brown and snuff-brown, Ridgway, 




FIGURES. Skull of Ochotona alpina 
alpina Pall., No. 149, 1897. Central Altai, 
A. A. Silant'ev, KZMAN*. Very large 
specimen 

Drawing by Yu. A. Kostylev 



23 



* (KZMAN — Collection of the Zoological Museum of the Academy of Sciences of the U, S. S. R. ] 



12 



24 



1912, pi. XXIX). Flanks and belly in winter fur much browner with a clear 
ocher tinge (cinnamon and clay color, Ridgway, 1912, pi. XXIX). 

The color of the summer fur of the young differs sharply from that of 
adult specimens in its light gray tone with only a slight rusty tinge. General 
color of back in these specimens intermediate between drab, hair-brown 
and mouse-gray (Ridgway, 1912, pl.XLVI and LI). The light rusty tinge 
(light pinkish cinnamon, Ridgway, 1912, pi. XXIX) is better marked on the 
flanks than on the back. Belly dirty-lead-gray with a slight rusty tinge. 

SYSTEMATIC NOTES. In his earlier work "Reise durch verschiedene 
Provinzen des Russischen Reichs", II, 1773, S. 569, Pallas describes the 
morphology and biology of the Altai pika from specimens caught not far 
from Tigiretskoe in the Tigeretskii Range. This should be considered as 
the locus classicus for this species. In his book "Novae Species 
Quadrupedum e Glirium ordine", pp. 45-59, 1778, Pallas mentions the 
capture of an Altai pika in Sinyaya Sopka on the Kolyvan river (in a locality 
north of Tigiretskoe). This is also briefly mentioned in his earlier book. 

Upon a careful systematic analysis of the Altai pikas, I arrived at the 
conclusion that several subspecies could be differentiated on the basis of 
color differences and characters of the skull. Pikas of the northwestern 
Altai do not differ from those of central Altai which are identical with the 
southern Altai form from the Narym Mountains. This race should therefore 
be considered as the typical O c h o t o n a alpina alpina Pall. Very 
large specimens sometimes occur in the central Altai (Figure 8). Our 
collection has only a few such large specimens and it is therefore 
impossible to determine whether their size is accidental, or whether a 
very large race indeed inhabits the central Altai. Only collection of more 
material will enable us to solve this problem. 

N. Hollister (1912) (Smithson. Miscell. Collect., Vol.60, No. 14, pp. 4-5) 
described Ochotona nitida from the Chegan-Burgazy (the upper 
reaches of the Chuya) which is only a small subspecies ofO. alpina 
distributed along the rivers Chuya, Chulyshman and in the region of the 
Teletskoe Lake. The same subspecies also lives in the Sayan Mountains. 
A well-characterized form, O. a. svatoshi Turov occurs in the 
Barguzin Range in Transbaikalia. O. a. cinereofusca Schrenck occurs 
in eastern and northeastern Transbaikalia, in the upper reaches of the Amur. 
A different form, O.a. scorodumovi Scalon is found south of the 
Gazimur Mountains. Finally, the peculiar subspecies O. a. changaica 
subsp. nov. lives on the Hangay Mountains. 

We give below an analysis of the differences between O . alpina and 
O. hyperborea, which are undoubtedly very closely related. 

It has been recently suggested that O . alpina and O. hyperborea 
should be considered as one species. A.I. Argyropulo has accepted this 
new classification, as far as I know, in his monograph in the series "Fauna 
SSSR" (Fauna of the U. S. S. R. ) which is now in preparation. I should 
like, therefore, to clarify my complete disagreement with this suggestion. 

O. alpina and O . hyperborea are certainly very closely related. 
A taxonomist could, however, identify O. hyperborea by its cranial 
characters and its short feet even if the animal were not labeled. 

O. hyperborea is an east Siberian, boreal species. It occurs 
mainly in the forest of the Taiga and in the Tundra to the coast of the 
Arctic Ocean. The range of the Northern pika extends as far as 
Manchuria (Great Khingan Mountains), northern Mongolia and on the 



13 



25 



Hangay Mountains. The range of O . a Ip i n a reaches from the Altai 
Mountains to Transbaikalia in the east and the Hangay Mountains in the 
south. The Altai pika lives in different habitats, namely screes and rocks. The 
ranges of these forms meet and overlap over a large area. Various 
subspecies of the Northern and the Altai pikas exist together in the same 
locality, where their specific ranges overlap. O. a. nitida HoUister 
and O . h. m an t c hu r i c a Thos. occur together in the Sayan Mountains. 
O. a. cinereofusca Schrenck and O.h. mantchurica Thos. 
occur together in the Gazimur Mountains; O. a. changaica subsp. 
nov. and O . hyperborea subsp. in the Hangay Mountains. 

It is hardly conceivable that all these forms belongto the same species. 
This is impossible from both the taxonomic and zoogeographical points of 
view. 

It is well known that in two closely related species, a small subspecies 
of the larger form frequently becomes similar to the large subspecies of 
the smaller form and vice versa. However, this is no reason to unite 
species. This would produce only confusion. Fortunately, nobody has as 
yet considered uniting S or e X araneus L. and S . mi nut us L. , in 
spite of the fact that both have large and small subspecies. Nor has anyone 
united Pipist re llus n at hu s i i and P i p i s t r e 1 lu s pipistrellus 
in spite of the existence of intermediate types of tooth structure. Even the 
expert and supporter of the specific cycle (Formenkreis), O. Kleinschmidt 
(Die Formenkreislehre, Halle, 1926, S. 46-49), rejected the suggestion 
that Passer domesticus and Passer hispaniolensis be united 
in spite of the existence of transitional forms. 

All this shows how careful one should be before reducing species; 
"simplification" may lead only to errors and confusion. 

GEOGRAPHICAL DISTRIBUTION. Ochotona alpina Pall, is widely 
distributed in the Altai Mountains. The type specimens were caught by 
Pallas in the Tigiretskii Range, and also on Sinyaya Sopka on the Kolyvan 
river. According to Pallas (1773) it is found in very large numbers in 
this place. A.M. Nikol'skii (1884) found a pika between Kuegan and 
Tourak, i.e., in the northern part of the central Altai, and also near 
Uimon (in the southern part of the central Altai). According to A. M. 
Nikol'skii many black specimens are found in the latter locality. N. F. 
Kashchenko (1899) recorded pikas from the following places: vicinity of the 
village of Shebalino; Tuekta in the Ursul' Valley (central Altai); along the 
Malaya Ul'chumen' river in the upper reaches of the Katun river; near the 
village of Kotandy; in the vicinity of Lake Ten'ga; in the vicinity of the 
village of Chergi; on the Ongudai (partly in central Altai, and partly in 
the northern part of the central Altai). I found pikas from the following 
places in the collection of the Tomsk University: vicinity of Tuekta 
(Tuekhta); upper reaches of the Katun river (coll. Prof. Sapozhnikov); 
Malo-Ul'gumenskie snow-covered peak; Kotandinskaya mountain-steppe. 
O. Thomas (1924) examined specimens from the following places: vicinity 
of Berel (southeastern Altai); the Chulyshman river (from Ust'-Bashkaus). 
Prof. N. F. Kashchenko (1902) found the Altai pika in the following places: 
Chulyshman river (near Tartychai); Bele Plot on the eastern coast of 
Teletskoe Lake. A large series of Altai pikas were caught near the 
Teletskoe Lake by the expedition of S. S. Turov and A. M. Kolosov. I have 
examined specimens from their collection from the mountains adjoining 
Teletskoe Lake (near the sources of Teletskoe Lake in the Biya river), from 



14 



26 



Altyn-Tu, from Dzhulu-Kul' etc. Satisfactory material from the Altai 
Reserve has been sent to the Moscow State Museum. 

B.S. Vinogradov (1924) has specimens from the northwestern Altai, and 
from screes in the vicinity of Ridder [Leninogorsk]. B. A. Kuznetsov (1932) 
obtained many pikas in the following places: the alpine-meadow region; 
the Sarym Mountains; the upper reaches of the Sarymsak river (in the 
Narym Mountains of southern Altai). I have a representative series 
of pikas from the mountains adjoining Katon-Karagai (Altaiskaya village) 
collected by V.I. Dotsenko. Pikas were caught in the extreme southeastern 
corner of Altai, near Chegan-Burgazy in the Chuya Alps. A specimen 
caught in this region was used by N. HoUister (1913) as holotype for his 
species Ochotona nitida. A.M. Kolosov obtained a good series on the 
Chuya Alps near Chegan-Burgazy, 

The range of Ochotona alpina in the Altai includes the northwestern 
and northern Altai, and the entire southern and southeastern Altai. This 
species is thus very widely distributed in this area. According to 
P.S. Vinogradov (1933), this species is distributed in northwestern Mongolia 
(on the Hangay Mountains). I have seen specimens from the Hangay 
Mountains in the ZMAN collection (collected by P.K. Kozlov, 1924). 

Ochotona alpina certainly occurs in the Sayan Mountains, where 
it was first recorded by G. Radde (1862). B.S. Vinogradov (1927) found 
O. alpina in the mountains near the estuary of the Kazyr-Suk river. 
According to B.S. Vinogradov and S.I. Obolenskii (1927) this species is 
found in the Minusinsk and Krasnoyarsk areas. He reports that the local 
pikas are related to the Altai species, although they are smaller. According 
to M.D. Zverev (1930) pikas are found in the mountain steppes of the 
southern parts of the Minusinsk and Khakassian districts. 

In Transbaikalia, O. alpina occurs on the eastern coast of Lake 
Baikal, and on the Barguzin Range (O. alpina svatoshi, S.S. Turov, 
Doklady Akademii Nauk, 1924, p. 110), I found a good series of O . alpina 
svatoshi in the material collected by S.S. Turov in the Barguzin Reserve 
(Sosnovka District) and in that collected by N. V. Nekipelov on the Bol'shoi 
Cheremshan and the Barguzin Range. We have, unfortunately, no 
accurate data on the distribution ofO. alpina svatoshi in Transbaikalia, 
especially in its northern part. According to V.N. Skalon (1936, in litt. ), 
O.a. svatoshi reaches the Trans -Kamensk District in the south and 
Chita in the east. 

O. alpina was first recorded from eastern Transbaikalia by L. V. 
Schrenck (1858). He identified material collected by R. Maak from the 
upper reaches of the Amur as O. hyper bore a var. c i ne r e o f u s c a . 
B. A. Kuznetsov (1929) found this species, which he considered to belong to 
species alp in a (subsp. c ine r e of u s c a ) in the Udinskii Range. V.N. 
Skalon (1936, loc. cit, ) thinks that O . alpina cinereofusca is 
distributed along the upper reaches of the Amur, in the Udinskii Range, 
reaching at least the upper reaches of the Gazimur river in the south. The 
steppe and forest-steppe regions of the eastern Transbaikal Ranges are 
inhabited by a peculiar form of the Altai pika, which was described by V.N. 
Skalon (1935) as separate subspecies, O. a. scorodumovi. It is 
found in the Kailastui District, near the Aleksandrovskii Zavod, to the north 
and west of the Borzya District, and south of the Gazimur river. It is 
not exactly known how far east the range ofO. alpina extends. O. 
Thomas (1909) identified the Ussurian pikas asO. cinereofusca. 



15 



This identification is, however, doubtful. A pika of the group hyperborea 
(O . h. mantchurica Thos. ) occurs on the Great Khingan Mountains. 
B. Howell (1928) described the race O . alpina argentata from northern 
Kansu. If this race indeed belongs toO. alpina, the eastern range of 
this species thus reaches southeast to North China. As already stated, the 
range ofO. alpina in the east of the upper reaches of the Amur and to the 
southeast of Transbaikalia is unknown. 

MODE OF LIFE. HABITATS. Screes are the typical habitat of the Altai 
pika. B.S. Vinogradov (1924) observed pikas in such places in the vicinity 
of Ridder. 

A detailed description of the habitats ofO. alpina is given by Prof. 
N. F. Kashchenko "Rezul'taty Altaiskoi zoologicheskoi ekspeditsii, (Results 
of a Zoological Expedition to the Altai) 1899, pp. 51-56: "Pikas are widely 
distributed on the Altai. They apparently occur wherever suitable 
screes are available. They choose only screes with stones about 
1/2 - 1 m large. Pikas occur also in screes of larger stones, though 
less often. They never occur in screes of small stones, apparently 
because there is insufficient space between the stones. 
27 "Screes inhabited by pikas can be divided into 3 types according to their 
origin. To the first type belong stones breaking from rocks as a result 
of disintegration. Such fragments accumulate in hollows between rocks, are 
completely devoid of vegetation and consist of several layers. Screes of 
this kind rarely occur in easily accessible, flat or gradually sloping 
localities. They are usually found at greater heights and on steep slopes. 
Stones are easily displaced and rolled if one walks on them. We found 
many such screes near Ongudai on 17 and 18 June. Many of them 
consisted of small stones. We found pikas only near maral farms of Messrs. 
VarvinskiiandMakar'ev 7 versts* from the village, in a high and little 
accessible hollow between rocks. 

"The second type of screes is represented by rounded mountain peaks 
which are sometimes completely covered with rock debris. These screes 
are sparsely overgrown with low birch and a few other plants of the alpine 
tundra. These were apparently rocks which were transformed to such a 
large broken area during many thousands of years. We found pikas (25 
June) in screes higher than the snow which was not continuous but formed 
patches on the Malo-Ul'gumenskie snow-covered peak. 

"To the third type of screes belong formations which are very similar 
to screes as far as the life of pikas is concerned, namely, man-made 
structures. In many valleys in the Altai, especially along the river Urusul, 
there are ancient tombs (tumuli) made of unhewn stone in various stages 
of ruin: some of them are almost worn to the ground, others are still high. 
Most of the tombs have the shape of a ring, with a central hollow open to 
the ground or even deeper. These hollows, which may not be much more 
recent than the tombs themselves, were probably made by grave-robbers. 
Pikas settle in these remnants of man's past glory, but they choose only 
tombs constructed of sufficiently large stones. " 

B.A. Kuznetsov found many pikas in high-mountain screes in the southern 
Altai (Narym Range). He describes vividly his first encounter with this 
peculiar rodent: "We were climbing in a small winding caravan towards 
the Sarym mountain path to Lake Marka-Kul. We left the wooded taiga 



♦ [Veret: 1.067 km.] 



16 



28 



far below us. The last cedars had disappeared. In the gray mist of the 
cloud around us we saw the enormous bulks of the wild cliffs, separated 
by hollows filled with deep firn. Here and there alpine meadows stretched 
on the southern slopes, on which creeping juniper and dwarf birch alternated 
with areas covered with a lush carpet of various high-mountain grasses 
and flowers. Silence reigned everywhere, such as only exists on snow 
covered mountains. Suddenly, a peculiar shrill whistle from a very close 
scree cut through the fog. I looked a long time for the source of this 
whistle, but in vain. The screes seemed completely desolate. Suddenly 
the muzzle of a small animal with black gleaming eyes and large rounded 
ears looked out from under a huge overhanging slab of limestone. Then the 
animal jumped out and settled on a nearby block, scrutinizing with 
astonishment men who are so rare here. It was a pikal "=■' 

In the southern region of the Teletskoe Lake A.M. Kolosov (1933) found 
pikas not only in their typical habitat, screes, but also in other habitats 
such as dense mixed forests with rich Spire a overgrowth (at 1500 to 
1800m altitude). On another occasion pikas were found above the 
forest zone (at about 2000 m) in arid, sunny, Artemisia and Festuca 
regions. It is of interest that pikas were found in the high-mountain 
alpine tundra, in the vicinity of Lake Dzhulu-Kul' (near the sources of the 
Chulyshman river). According to A.M. Kolosov, pikas in this region 
inhabit moraines which are surrounded with bogs and thickets of polar 
birch. 

Altai pikas persistently inhabit screes even when their external 
appearance is completely changed. N. A. Avrorin ("Materialy o tipakh lesov 
gornogo Altaya" (Forest Types of the Altai Mountains). —Trudy Instituta 
po izucheniyu lesa, Akademiya Nauk, issue I, 1933) recently described the 
formation of screes as follows: screes result at first from eolian 
disintegration of rocks. Such "young" and energetically- growing screes 
are mobile and overgrown only with lichens. Neither mosses nor flowering 
plants can exist in them. In the second stages the screes become 
"quiescent", the stones losing their mobility. Such screes are covered 
with moss and some flowering plants (Sedum hybridum). Disintegrating 
rock particles reach the lower margin of the scree more rarely and 
vegetation is thus more intensive there. The following plants can be found 
at the periphery of the screes: currant (R ibe s nigrum, R. atro- 
purpureum), gooseberry (R ibe s grossularia), very fragrant and 
sweet raspberry (Rubus melanolasius) etc . 

Later, screes become densely overgrown with Caragana 
arborescens, Lonicera coerulea, L. microphylla. 
Spiraea chamaedryfolia, Berberis sibirica etc. The 
afforestation of the "old" scree takes place under the shelter of these 
shrubs . 

According to A.M. Kolosov, pikas do not leave screes, even when screes 
of the first type become gradually transformed into mountain forest regions. 
V. V. Dmitriev kindly placed his interesting observations on the life of 
pikas in such changing habitats at my disposal. He found large pika 
colonies on the eastern coast of Lake Ity-Kul' in the upper reaches of the 
Chul'cha river (a tributary of the Chulyshnaan river), and also in the upper 



* B. A. Kuznetsov, Gryzuny Semipalatinskogo okiuga (Rodents of the Setnipalatinsk District). — Byulleten' 
Moskovskogo obshchestva ispytatelei prirody, 51 (1—2): 115- 116. 1932. 



17 



29 



reaches of the Bidui river (aright-hand tributary of the Abakan river). The 
screes have here completely lost their primitive character: alluvial soil is 
never found, and stones and rocks are covered with moss. The pikas hide in 
burrows. The exits of the burrows are covered with green moss. Large 
cedar groves interspersed with spruce grow on such changed screes. The 
screes are on the northern slopes of the mountains. The ground is 
covered with a thick layer of green moss and thickets of wild rosemary and 
round-leaved birch. Food stores of pikas here consist only of wild rosemary 
and birch leaves. Only rarely do they include some C a lam a gr o s t i s 
grasses, which grow very scantily here. 

In the Altai mountains pikas inhabit heights of from 400- 500m to 2500m. 

GENERAL DESCRIPTION AND HABITS. In places where pikas do not 
(or only rarely) meet man, and where they are not hunted, they are not 
timid and often permit close approach. When the animal is frightened it 
looks out of its burrow slowly and very cautiously, puts out its head, looks 
around, and hides again among the stones. V.N. Skalon (in litt. 1936), who 
found pikas in eastern Transbaikalia, in the Baiolga ravine, reports that 
Ochotona alpina c i ne r e o fu s c a is rather timid. "On seeing man 
the animal either hides without movement, or whistles (though not as 
shrilly as O . alpina*) and disappears in its burrow. If one sits 
motionlessly the animal will soon reappear. It comes out of its burrow, 
looks around and goes about its business". 

Its voice is a strident, sharp whistle. The movements of the pika are 
rapid but not as quick and graceful as those of hares. Compared with them, 
the gait of these short-legged animals is a clumsy waddle. 

DAILY ACTIVITY. According to B.A. Kuznetsov (1932), the activity 
of pikas starts at daybreak and ends at dusk. Their whistles are sometimes 
heard even at night. 

According to A.M. Kolosov's observations in the Teletskoe Lake region, 
the pikas are heard in the morning from 9 a. m. , when the ground has 
already become heated. 

According to V. N. Skalon (in litt., 1936) O . a. c ine r e o f u s c a does 
not leave its underground shelters at definite hours. Its activity is not 
strictly confined to definite hours. Altai pikas are usually active from 
daybreak until the strong heat begins, and then again from 4-5 p. m. until 
sunset. 

V.N. Skalon frequently saw O . a. cinereofusca at noon. He 
thinks that the irregularity of their daily activity may be partly due to the 
fact that they usually inhabit deep narrow ravines where the sun's rays 
hardly penetrate. 

BURROWS, FOOD AND FOOD STORES. The burrows of Altai pikas are 
usually built in screes between stones. Food is stored in typical "stacks" 
near the burrows, often under stones. Food is also hoarded deep inside the 
burrows. B.A. Kuznetsov (1932) reports the following: "In August, when I 
collected material on the snow-coveredNarym Mountains, I observed that the 
pikas were busy preparing hay for the winter. In the morning they could be 
observed scurrying among the stones of the scree carrying grasses to their 
nests. I found large stores of dried hay between the stones, solidly rammed 
into cracks. These hoards consisted of many different plants, but cereals 
were not represented in them at all". 



♦ V. N. Skalon apparently means the typical Altai form. 



18 



V.N. Skalon (in litt. 1936) describes food collection by pikas as follows: 
"The pika moves slowly, its movements like those of a calmly feeding 
rabbit. It examines the ground, paying particular attention to the yellow 
birch leaves which cover it. A suitable leaf is taken at the edge and borne 
to the burrow. The pika repeats this several times, and discards dozens of 
unsuitable leaves. When it becomes tired and hungry it chooses a leaf, 
calmly sits down and eats rapidly, moving its muzzle like a rabbit. " 

V.N. Skalon adds: "Food -hoar ding pikas do not only drag leaves into 
the burrow. They also store leaves on the ground, apparently using 
leaves driven by the wind under fallen trees. " 

A. M. Kolosov (1933) found the following plants in food stores of pikas 
in the Teletskoe Lake region on the Altai: Rubus idaeus, Rosa 
acicularis, Athyrium filix femina, Pteridiura. 
aquilinum, Saxifraga crassifolia, Lycopodium annotinum, 
Vaccinium vitis idaea, Betula verrucosa. Sphagnum, 
Archangelica descerrens, Cystopterus fragilis, Carex 
sp. 

A.M. Kolosov found many pika bite-marks on shrubs. In some cases 
finger -thick twigs had been gnawed. He assumes that pikas sharpen their 
teeth on such twigs. 

REPRODUCTION. We know very little on the reproduction of alpine 
pikas. No data on this subject have been published. The collection of 
pikas made by A. M. Kolosov in 1936 contains a series of young specimens 
which are two -thirds as long as adults (caught in the first ten days of July). 
The dates and number of litters per year are unknown. 

B.A. Kuznetsov (1929) states that O . a. c ine r e of us c a has at 
least two litters per year. 

Kuznetsov writes as follows: "Working on the Udinskii Range in late 
June, I saw young pikas of the first litter (16-17 cm long) already living 
in separate burrows. At the same time, young of the second litter were 
still suckled, only occasionally coming out of the burrows to get warm in 
the sun. They were the size of a vole. A litter apparently consists of 
4-6 specimens". 

Our knowledge of molts is also very incomplete . Examination of 
„ material from collections gave the following conclusions. Molting into 
summer fur begins between May and June. Males molt earlier than 
females. Molting is delayed in females, apparently in connection with 
gestation. Some males have almost finished their molting by the first half 
of July. The spring-summer molt takes place in a definite sequence. The 
flanks, head and front part of the back molt first, and the posterior part 
of the back later. There is one female specimen in my collection (No. M. 
3979) caught at an altitude of 2300m, near Katon-Karagai on 6 July 1931, 
still bearing her complete warm winter fur. Beginnings of molting were 
observed in some females caught in late July and early August. Molting in 
females normally starts in late July (e.g.. No. 240 ?, 27 Jul. 1936 Kuraiskaya 
Steppe, Altai, A.M. Kolosov) or in early August (No. 2358$, 13 Aug. 1929. 
Katon-Karagai, B.A. Kuznetsov). 

The replacement of the short gray fur of half-grown specimens by the adult 
rust-colored fur takes place in August -September, sometimes beginning 
even earlier, in late July. The fur of the flanks, cheeks and anterior part 
of head is shed first, and then that of the back. The fur of the nape is the 
last to be shed in some specimens. 



19 



Winter fur is substituted for summer fur in late September and October in 
adults. Specimens caught in late November and December had already 
completed their molt and bore full winter fur. 

In yearlings which had recently assumed their adult summer coat 
(August-September), molt and growth of winter fur set in rapidly. This 
process was not found in these yearlings. It is, however, probable that 
the full molt into winter fur closely follows the early fall molt in which 
the young assume their adult fur. The process is therefore almost 
continuous. 

This subject is very interesting and should be further studied on a 
large amount of material. 

These pikas do not hibernate. 

ECTOPARASITES. A.M. Kolosov (1933) states that a small parasitic 
tick which lays many eggs is very common on the ears of O . a 1 p i n a . 
The frequent scratching of the ears with a hind paw observed in them is 
apparently due to the itching caused by these parasites. 

E.N. Pavlovskii (in "Zveri Tadzhikistana" (Animals of Tadzhikistan). 
pp. 54-55, 1935) writes that the skin of O . alpina is parasitized by larvae of 
Oestromyia pallasii. He gives a good drawing of these larvae 
in a section of the body wall of the Altai pika (Figure 23, p. 54) 



Subspecies of Altai or Alpine pika 

309. Ochotona (Pika) alpina alpina Pall. (1773). Central Altai 
pika 

1773. Lepus alpinus Pallas P. S. , Reise durch verschiedene Provinzen des Russischen Reichs, II, S. 569, 

701. 
1778. Lepus alpinus Pallas P. S. , Novae species Quadrupedum e Glirium ordine, p. 45-59. 
1811. Lepus alpinus Pallas P. S. , Zoographia Rosso- Asiatica, p. 150. 
1842. Lagomys ater Eveismann E. , Addenda ad celeberrimi Pallasii Zoograph. Rosso-Asiatica, p. 3 

(type from vicinity of Uimon). 
1899. Lagomys al pi nus Kashchenko, N. F. , Rezul'taty Altaiskoi ekspeditsii (Results of an Expedition 

to the Altai), pp. 54-57. 
1924. Ochotona alpina Vinogradov B. S. , Nablyudeniya nad gryzuaami Zapadnoi Sibiri (Observations 

on Rodents of West Siberia), Izvestiya Sibirskogo entomologicheskogo byuro, No. 3, p. 36. 
1932. Ochotona alpina Kuznetsov B. A. , Gryzuny Semipalatinskogo okruga Kazakhstana (Rodents of 

the Semipalatinsk District of Kazakhstan), Byulleten' Moskovskogo obshchestva ispytatelei prirody, XII 

(1-2):115-116. 

TYPE LOCALITY. Vicinity of Tigiretskoe in the Tigiretskie Mountains of Northwestern Altai. Type 
unknown. 

MATERIAL EXAMINED: 42 pelts (most of them with skulls). 

DIAGNOSIS. Skull large, length in adult specimens 47-57 mm (M. 50.2); 
condylobasal length 44-53 mm (M.46.8); zygomatic width 22-26 mm 
(M. 23.6); width of skull at level of auditory meatus 22-25.7 mm (M. 23.6). 

Color of summer fur on back in adult specimens dull yellow-ocher-gray 
(tawny-olive), darkened by black or very dark brown hair tips. Flanks with 
marked rust-red tinge (pinkish cinnamon); belly pale, yellow-ocher 
(pinkish cinnamon to cinnamon-buff). 

Winter fur dirty-ash-gray with marked black tinge caused by 
very dark brown hair tips. Lower part of back usually with rusty tinge. 
Belly dull dirty-grayish with a marked rusty tinge. 



20 



33 



The general gray tone of the back in the winter fur of some specimens 
is mixed with dull brown (sayal-brown and snuff-brown). Flanks and belly 
of such specimens with marked ocher tinge*. 

SYSTEMATIC NOTES. There are no pelts and skulls from the type 
locality of O c hot on a alp in a, namely the Tigiretskie Mountains, in our 
zoological museums. I examined pelts from the Ust-Kamenogorsk vicinity 
(collected by Kinderman, 1853) in the ZMAN collection and also specimens 
from the vicinity of Ridder [Leninogorsk] i.e., from localities near the 
typical region of O . alpina Pall. 

Their general color is quite bright, and they are distinguishable from 
the southern Altai specimens from the Narym Range. The northwestern 
Altai and its central and southern parts are thus apparently inhabited by 
the typical form. 

GEOGRAPHICAL DISTRIBUTION. From the Tigiretskie Mountains in the 
north to Ust-Kamenogorsk, Ridder [Leninogorsk] in the west, to the 
Katun river in the east, and the Uimon and Narym Mountains in the south. 

310. Ochotona (Pika) alpina nitida Hollist. (1912). East Altai pika 

1902, Lagomys alpinus Kashchenko, N. F. , Mlekopitayushchie Altaiskoi ekspeditsii Ignatova (Mammals 
Collected by Ignatov's Expedition to the Altai), Ezhegodnlk Zoologicheskogo muzeya Akademii Nauk, 
VII, p. 300. 

1912. Ochotona n i t i d a HoUister N. , New Mammals from the Highlands of Siberia, Smithson. 
Miscellan. Collect. , Vol. 60, 14, p. 4. 

1913. Ochotona nitida HoUister N. , Mammals Collected by the Smithson. Harvard Exped. , Proceedings 
U.S. National Museum, pp. 522-523. 

TYPE LOCALITY AND DEPOSITION. Chegan-Burgazy mountain pass in the upper reaches of the Chuya 
Alps, southeastern Altai. Type No. 7539 in the collection of the U. S. National Museum, 
MATERIAL EXAMINED: 60 pelts with skulls, 

DIAGNOSIS. Skull smaller than in the typical race, its length in adult 
specimens 15-51 mm (M.47.6); condylobasal length 41-48 mm (M. 43.7); 
zygomatic width 21. 2-24 mm (M. 22.5); width of skull at level of auditory 
meatus 21.2-23 mm (M. 22.3). 

Color of summer fur on back generally dark brown, mikado-brown to 
snuff-brown; color of summer fur on back even darker, in some specimens 
intensely chestnut-brown with marked blackness; rust and ocher shades very 
marked on flanks (sayal-brown and cinnamon). Belly gray-ocher (cinnamon- 
buff with a strong gray tinge). 

SYSTEMATIC NOTES. N. HoUister (1912) described this form from a 
specimen*'!" from the Chegan-Burgazy mountain pass. He stresses the dark 
color of the summer fur ofO. nitida on the back ("general color of 
upper parts dark mummy-brown, a mixture of russet and black; nose, 
cheeks and area behind ear lighter"), and also the smaller size of O 
nitida as compared with north Altai forms. 

O. alpina nitida is certainly well differentiated as a subspecies. 

GEOGRAPHICAL DISTRIBUTION . Ochotona nitida lives in the 
eastern Altai, in the Teletskoe Lake region, along the Chulyshman, 



* Completely black (melanistic) specimens often appear among central Altai pikas, and also among those 
from the northern and southern Altai. 
** HoUister had a series from the Chegan-Burgazy mountain pass and from Tapuchi {31 pelts) in addition to 

this type. 



21 



Chul'ga and Bashkaus rivers, in the upper reaches of the Chuya river 
(Chegan-Burgazy), reaching the Sayan Mountains in the east. 

311. Ochotona (Pika) alpina ahangaica subsp. nov. Hangay pika 

TYPE LOCALITY AND DEPOSITION. Mongolia, Hangay, vicinity of Subur-Hayrhan, Type No. 26, 

1925 (24) 14Aug. 1924, P. K. Kozlov, in the collection of the Zoological Museum of the Academy of Sciences. 
MATERIAL EXAMINED. The following specimens were examined in addition to the type: 2) No. 125, 

1926 (7) cf northern Hangay, basin of the Orhon, 1925, P. K. Kozlov; 3) 20 220 5, 8 Jul. 1929, southwestern 
Hangay, screes in Ohton-Tengri; 4) No. 59 5, 19Jul. 1929Hangay, rocks along river Bombotu (an 
eastern tributary of the Buyanta Gol river; 5) No. 26, 1925(23), 15Aug. 1924, Subur-Hayrhan Hangay, 
P.K, Kozlov. 

DIAGNOSIS. This subspecies is closely related to the typical 
alpina, closely resembling a series from the Narym Mountains. Series 
ofO. a. changaica in summer fur paler than typical form; upper 
surface of light yellowish collar duller than inO. a. alpina. Color 
of winter fur more grayish white than in nominal race. 

Dimensions smaller than in O . a. alpina. Skull length 46-48.9 mm 
(M.48); condylobasal length 42.7-45.2 mm (44.3); zygomatic width 22.5- 
23mm (M. 22.7); width of skull at level of auditory meatus 22. 7-23. 2mm 
(M. 23). 
34 SYSTEMATIC NOTES. This subspecies is closely related to O . a . 

nit id a in its dimensions; it resembles the southern Altai O . a. alpina 
Pall, in its pale color, though both its summer and winter fur is paler 
(which is obvious in the series). 

GEOGRAPHICAL DISTRIBUTION. Hangay in western Mongolia. 

312. Ochotona (Pika) alpina scorodumovi Scalon (1935). 
Skorodumov's Alpine pika 

1935. Ochotona alpina scorodumovi Skalon V. N, , Nekotorye zoologicheskie nakhodki v yugo- 
vostochnom Zabaikal'e (Some Zoological Discoveries in Southeastern Transbaikalia), Sbomik Rabot 
Protivochumnoi organizatsii Vostochnosibirskogo kraya, Vol. I, pp. 85-87. 

1936. Ochotona alpina scorodumovi Skalon V. N. Dal'neishie issledovaniya po sistematike i 
biologii mlekopitayushchikh (Further Studies on the Taxonomy and Biology of Mammals), Izvestiya 
Gosudarstvennogo protivochumnogo instituta, Vol. IV, pp. 180-192. 

TYPE LOCALITY AND DEPOSITION. Northern Kailastui, Borzya District, Transbaikalia, type No. 25,009 
(289) male 14Sept. 1934, in the collection of the Zoological Museum of the Academy of Sciences. 

MATERIAL EXAMINED. 1 have examined only one specimen of O. A. scorodumovi Scalon. The 
author of this new subspecies described it from a series of 47 specimens from Kailastui, and gave an additional 
more detailed description from a new series of 244 specimens. 

DIAGNOSIS. V.N. Skalon gives the following diagnosis for O . a . 
scorodumovi; "Differs from O. a. alpina Pall, by its smaller size 
(length averaging less than 200mm). Length of type 189.5mm, average 
length of the whole series 182.8 mm; skull large, length of skull of type 
46.6mm, average skull length 46.66mm; condylobasal width of type 43.8mm, 
average 43.5; interorbital width 5.5mm, average 5.4 mm; zygomatic 
width of type 21.8mm, average 22.7mm; occipital height of type 12.1mm, 
average 12.0. Color gray-brown and rust -brown to blackish brown, 
always with marked streaks of black hair; color never with the rich rust 
tones of Oc hot on a alpina alpina Pall. Color of flanks usually 
different from that of belly and resembling that of back (Figure 9). 



1254 

22 



ADDITIONAL DESCRIPTION. According to V.N. Skalon, the top of the 
head and spine of specimens ofO. a. scorodumovi were either 
uniformly colored, or the middle of the spine was darker, especially 
posteriorly. In some cases the spine is almost completely black with a 
metallic luster. Color of sides of head lighter, tending toward grayish. 
Flanks always with a rusty tinge. Belly uniformly colored, as a rule paler 
and more grayish than the flanks. Ears with a well-marked white margin. 
Inside of ears grayish, outside almost black. 

Young specimens almost mouse-gray. 

Melanistic individuals never occur. 





FIGURE 9, Skorodumov's alpine pika, 
Ochotona alpina scorodumovi 
Scalon, type No. 25,009, ZMAN 
collection 



FIGURE 10. Bargiuinpika, Ochotona alpina 
svatoshi Turov, No. M. 3989 § 19 Aug. 1923, 
Shumilildia river, Sosnovka, Baikal, S. S. Turov; 
coUectionof S. I. Ognev 

Drawing by Yu. A. Kostylev 



35 



The following is a description of the color of a type of O. scorodumovi 
from a specimen from the Zoological Museum of the Academy of Sciences. 
Color dark. General shade of back dark brown with marked blackness 
caused by the black tips of the guard hairs. Blackness particularly strong 
on the lower and middle parts of back, less so on top of head. Flanks with 
marked brownish tones, close to cinnamon; a black tinge is also observed 
here, but less marked. Belly grayish rust, relatively pale. Muzzle 
behind eyes and on ears streaked with gray hair which is well marked on 
basal parts of ears. Chin and upper part of throat grayish. 



23 



36 



SYSTEMATIC NOTES. Ochotona a. scorodumovi is most 
closely related to O . a . cinereofusca; it differs from it by its 
deeper color with black tones on the back. According to V.N. Skalon, 
O. a. scorodumovi is also somewhat larger than O. a. cinereofusca. 
These facts should be carefully verified. On the Gazimur Range, where 
V. N. Skalon collected his series of O. a. cinereofusca, O. 
hyperborea m ant c hu r i c a Thos. also occurs. It may be concluded 
from measurements of O . a. cinereofusca (published by V. N. 
Skalon) that he dealt with heterogeneous material. Adult specimens with a 
skull length of 39.2-42 mm, which appear in the table (No. 10-11), 
undoubtedly belong to hyperborea. 

The only certain difference between O. a. scorodumovi and O . a . 
cinereofusca is thus the color . It may be more correct to regard 
O. a. scorodumovi as a natio of cinereofusca. 

GEOGRAPHICAL DISTRIBUTION. Borzya District of Transbaikalia. 

313. Ochotona (Pika) alpina svatoshi Txirov. (1924). Barguzin 
pika 

1923. Lagomys al pinus Turov, S. , K voprosu o Barguzinskom sobolinom zapovadnike (The Barguzin 
Sable Reserve), Izvestiya Vostochnosibirskogo otdeleniya Russkogo geograficheskogo obshchestva, XLI 
(2): 13 (bare peaks along the Burtui river). 

1924. Ochotona svatoshi Turov S.S, O faune pozvonochnykhzhivotnykhsevero-vostochnogo 
poberezh'ya oz. Baikala (The Vertebrate Fauna of the Northeastern Coast of Lake Baikal), Doklady 
Rossiiskoi Akademii Nauk, p. 110 (Bargiuin Range). 

1929, Ochotona svatoshi Kuznetsov B. A. , Gryzuny vostochnogo Zabaikal'ya (Rodents of Eastern 

Transbaikalia), Izvestiya Assotsiatsii nauchno-issledovatel'skikh institutov, 1 (1): 88-89. Moscow. 
1933. Ochotona hyperborea svatosc hi Vinogradov B. S. , Mlekopitayushchie SSSR (Mammals 

of the USSR), Gryzuny, p. 83. 

TYPE LOCALITY AND DEPOSITION. M 3985, summer 1922, Shumilikha ravine, Sosnovka, 
Bargiuin Reserve. Type in S. I. Ognev's collection. 

MATERL^L EXAMINED: 15 pelts and skulls. 

DIAGNOSIS. Skull small, length in adult specimens 44-47.2 mm 
(M. 15.6) [sic]; condylobasal length 40.7-45 mm (M. 42.8); zygomatic width 
21.1-22.2 mm (M. 21.6); width of skull at level of auditory meatus 21.2 - 
23.1mm (M. 22.1). 

Summer fur on back of fully grown specimens rich reddish ocher 
(sayal-brown) with an orange tinge (orange-cinnamon). General color of 
flanks very slightly different from that of the back. Winter fur reddish ocher 
with a marked black tinge caused by black tips of the long guard hairs. 

Fur of young markedly grayer and duller, generally dull gray-rust-ocher. 
Sides of body and cheeks with well marked ocher tinge. 

SYSTEMATIC NOTES. S.S. Turov (1924) described the Barguzin pika 
as a separate species. B. A. Kuznetsov ("Gryzuny vostochnogo Zabaikal'ya" 
(Rodents of Eastern Transbaikalia), pp. 88-89, 1929) confirmed the 
specific status of O . svatoshi. B.S. Vinogradov (1933) considered O . 
svatoshi as a subspecies of hy p e r b o r e a . This form occupies in fact 
an intermediate position between O . alpina and O . hyperborea. 
A true, small form of hyperborea occurs in the Transbaikalia mountain 
taiga. Furthermore, the skull structure and size of svatoshi are more 
closely related to alpina. It is therefore, more correct to consider O. 
svatoshi as belonging to alpina. 



24 



GEOGRAPHICAL DISTRIBUTION. O.a. s vat o s h i is found on the 
Barguzin Range in the Sable Reserve area. Due to lack of material, it is at 
present impossible to define the northern limit of this form's range, and 
whether it occurs on the Ikatskii and Yuzhno-Muiskii ranges. V.N. 
Skalon (in litt. 1936) assumes that it reaches Chita in the east; this is 
on quite possible. It is also unknown how far south this form occurs. An 
alcohol specimen of O c ho t on a alpina caught 327 km from 
Troitskosavsk [Kyakhta] may belong to this form, but I doubt it. This 
specimen was mentioned by N. F. Kashchenko (1910). A badly preserved 
specimen (Zoological Museum of the Tomsk University) caught by P. 
Mikhno in the Aga steppe (Alkhanai bald peak) is closely related to O . a . 
svatoshi. It is possibly intermediate between O . a . svatoshi and 
O.a. cinereofusca, to judge by its color. It is however difficult to 
define the systematic position of this form from a single pelt. 

314. Ochotona (Pika) alpina cinereofusca Schrenck (1858). 
Amur pika 

1858. Lagomys hyperboreus var. cinereofusca Schrenck, L. , Reisen und Forschungen im 

Amur-Lande, I,S. 150, Taf. VIII, Fig. 2. 
1923. Ochotona hyperborea cinereofusca Soweby, A. de C. , The Naturalist in Manchuria, 

II -III, p. 174. 
1929. Ochotona (Pika) cinereofusca Kuznetsov, B. A. , Gryzuny vostochnogo Zabaikal'ya 

(Rodents of Eastern Transbaikalia), Izvestiya Assotsiatsii nauchno-issledovatel'skikh institutov., 

2 (l):87-89, 
1933, Ochotona hyperborea c inereof use a Vinogradov B. S. Mlekopitayushchie SSSR. Gryzuny 

(Mammals of the USSR. Rodents) p. 83. 

TYPE LOCALITY. Described from a specimen caught by R. Maak in the upper reaches of the Amur 
river. 

MATERIAL EXAMINED: 8 skulls and pelts, 

DIAGNOSIS. Skull small, length in adult specimens 44-49 mm (M.45.8); 
condylobasal length 40-44.9 mm (M.42.6); zygomatic width 21-23 mm 
(M. 21.7); width of skull at level of auditory meatus 21-23 mm (M. 22.2). See 
Figure 11, p. 27. 

General color of summer fur on back dull grayish brown with a slight 
ocher tinge. This ground color is markedly darkened, especially along 
middle of back, by black or blackish brown tips of hair; overall color of 
back intermediate between tawny-olive, sayal-brown and snuff-brown 
(Ridgway, 1912, pi. XXX). Color of front part of head with distinct gray 
tones, especially marked in the anterior part of the forehead, on the upper 
parts of the cheeks, and near the ears. Many blackish brown hairs on 
the occiput. Flanks dull rust (intermediate between pinkish cinnamon and 
cinnamon -buff, Ridgway, 1912, pi. XXIX), but more dirty-grayish. Belly 
lighter and grayer; sides of belly more yellowish. 

Winter fur* dirty and dull gray with rich straw-rusty tinge. 

Color of young more grayish and dull; it generally resembles the color 
of young O. a. svatoshi, but is paler and grayer. 

SYSTEMATIC NOTES. B.A. Kuznetsov (1929) observed that O.a. 
cinereofusca and O. a. svatoshi differ from each other as a rule 
only in color. However, Kuznetsov's opinion that the color of O . a . 



* My knowledge of its color comes only from patches of winter hair retained in the lower part of the back 
by incompletely molted summer specimens. 



25 



cinereofusca is "richer" than that of s v at o sh i is incorrect. The 
color ofO.a. cinereofusca is duller, grayer and yellower than that 
ofsvatoshi, and lacks the intense ocher, reddish, and orange tones 
characteristic of the summer fur of fully grown svatoshi. These 
38 differences can be clearly seen in the series. 

V.N. Skalon ("Nekotorye zoologicheskie naJchodki v yugovostochnom 
Zabaikal'e" (Some Zoological Discoveries in Southeastern Transbaikalia), 
1935, p. 85) described a peculiar subspecies, O . a . s c o r o dumo vi from 
near the village of Kailastui (Borzya District). The skull of this form is 
larger than that ofOch. a. cinereofusca according to him (vide supra) 
the mean skull length ofO. a. scorodumovi was 46,45mm; the skull 
of specimens from the Baiolga Valley (north of the Gazimur Range) 
identified by V. N. Skalon as O . a . c i ne r e o fu s c a , is much smaller, its 
mean length being 42.75 mm. The skull of specimens of O. a . c i ne r e o f u s c a 
collected by B. A. Kuznetsov on the Udinskii Range is not smaller than that 
ofO. a. scorodumovi; it is even slightly larger (see tables). The 
Udinskii Range lies north of the Gazimur Range and the Baiolga valley. 
The cranial differences between O. a. scorodumovi and O. a. cinereo- 
fusca should therefore be further studied. 

GEOGRAPHICAL DISTRIBUTION. The exact boundaries of the range of 
this subspecies are not clear. L. Schrenk described cinereofusca from 
specimens obtained in the upper reaches of the Amur river. B. A. 
Kuznetsov obtained a series of these pikas from the Udinskii Range south of 
Nerchinsk. V. N. Skalon recorded c i n e r e o fu s c a from the Baiolga 
Valley, north of the Gazimur Range. 

This subspecies is apparently widely distributed in eastern and northern 
Transbaikalia, reaching the upper reaches of the Amur river in the east. 
G. Radde (1862) found a specimen in the upper reaches of the Digan river 
(in the Bureya Mountains). He notes its relatively large size and considers 
it a cinereofusca. 

O. Thomas (1909) found a specimen of this subspecies in the Ussuri 
Valley. His observations should, however, be verified. 

150, Ochotona (Pika) hyperborea Pallas (181 1). Northern pika 

1811. Lepus hyperboreus Pallas P. S. , Zoographia Rosso- Asiatica, p. 152. 

NAMES: Northern Pika, North-Siberian Pika (English); der Nord-Sibirische Pfeifhase, der nordliche 
Pfeifhase (German), 

LOCAL NAMES. Piskun [squeaker] (Russian, on the Lena river); Kamennaya pishchukha [stone pika] 
(Russian, on the Amur river); chis (onomatopoetic); khaya-kutuyakh (mountain mouse); tas-kutuyakh 
(stone mouse) (Yakut); chiptyrra, or chantyrra (Tungus- Evenki); ogotnon (Buryat); pika (Lake Baunt Evenki, 
TungTis); chipa (Orochi on the Amur); teperkan (Orochi on the Gilyui); piski [squeaks] (Orochi in the 
Far East). 

PRINCIPAL nCURES: 1) Schrenck L, V. , Reisen und Forschungen im Amur Lande, I. 1858, S. 150, 
Taf. VII, Fig. 1 (Lagom ys hyperboreus var. normalis), Taf.VII, Fig, 2(L, h. var. ferruginea), 
Tai, VIII, Fig, 1 (L. h. var, cinereoflava); 2) Flerov K. K. Pishchukha sevemogo Urala (The North 
Ural pika), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, Vol. XXVIII, 1927, pp. 138-144, figs. 1-3, 
pp. 141-142 (entire animal); pi, IV, 1-4, (photograph of skulls), 

MATERIAL EXAMINED: 206 pelts (most of them with skulls). 



39 



DIAGNOSIS, Skull small, length in old animals usually less than 45 mm, 
42,5mm in very large and old individuals, Interorbital space of adult and 
old specimens flattened and 4,3-5.7 mm wide. Anterior third of nasals 
without spoon-shaped widening. Lateral contours of posterior part of nasals 
very slightly narrowing posteriorly (less than in Ochotona alpina), 

26 



with rounded ends. Frontals without fenestrae. Vomer ventrally covered 
by the margins of the premaxillaries; palatine and incisor foramina 
separate. Vibrissae mostly blackish brown, with whitish tips, 42-55mm 
long. Soles covered with grayish white brownish or even blackish hair. 
Ears with very narrow whitish margins. 




FIGURE 11, Amur pika, Ochotona 
alpina cinereof use a Schrenk, 
No. 588 23/ VI 1928, Nerchinsk 
District, B. Kuznetsov 

Drawing by Yu. A. Kostylev 



FIGURE 12-13, Skull of Northern pika, Ochotona hyperborea 
cinereoflava Schrenck. (dorsal and ventral). No. M. 3989 7/III 
1928. Ol'skiy Island, Rassvet Bay, G, D. Dul'keit, S. I, Cgnev's 
collection 

Drawing by Yu. A, Kostylev 



The color of the summer fur varies greatly in different subspecies. The 
general tone of the back is quite pale in the nominate form, grayish brown with a 
light straw rusty tinge. Reddish tones (sayal-brown, Ridgway, 1912 pi. XXIX) 
well marked on lower part of back, upper part of back more straw colored. 
Flanks darker than back; belly whitish with a distinct straw-yellow tinge. 

O. a. cinereoflava has an even paler fur. Its summer fur is distinguished 
by a rich gray-yellow color with a weak rusty tinge. Flanks rich rust-yellow. 
O. h. f e r r ugine a has a saturated rust-red-ocher fur, which becomes intense 
red-rust-brown in the middle of the back. The Amur-TransbaikalianO. h. 
mantchurica has a relatively dull chestnut -brown fur. 

The winter fur differs sharply from the summer fur in its grayer or dirty- 
grayish color with a brownish tinge. 

Color of young which have reached half the size of adults dull gray. 
Body length may be 210 mm, usually about 150-160 mm; hind foot 30 mm 
long, usually 26-27 mm or 22-30 mm. 

SYSTEMATIC NOTES. O. hyperborea is sytematically very closely 
related to O. alpina. In B.S. Vinogradov's (1933) opinion the only 
difference between them is the small size of the former. This is a good 
criterion, but it does not exclude other specific differences. 



27 



The skull of O. hy p e r bo r e a has many characteristic features (see 
Figure 12-13, p. 27): 

1) Most important is the short rostrum ofhyperborea. The distance 
between the inner margins of the alveoli of the last upper molar is about 
equal to the length of the upper diastema, measured to the alveoli of the 
small posterior incisors; in alp in a the diastema is much longer than the 
distance between the alveoli of the molars. 

2) The shape of the nasals ofhyperborea differs from that of 

alp in a. This can be seen in a series. The width of the nasals is almost 
equal anteriorly and posteriorly in hyperborea while the nasals of 
alp in a are markedly widened anteriorly. 

3 ) The brain case of O. hyperborea is shorter and more rounded 
than in a 1 p i n a . 

4) The orbits ofO. hyperborea are more rounded, not as elongate 
as in alp in a . 

All these differences can be clearly seen in a series. The differentiation 
between the two species is difficult, because the subspecies of alpina 
contain some rather small forms, e.g.,0.a.svatoshi, O.a. 
cinereofusca etc. However, the above-mentioned characters, 
particularly the rostral structure, always determine the species of a specimen. 

The following table of measurements may prove useful for general 
orientation. 

O. alpina O. hyperborea 

Length of skull , more than 43 mm less than 43 mm 

Condylobasal length " " 41 " 

Zygomatic width ii " 22 " 

Width of skull at level of auditory meatus .... « « 21 " 

Diastema " " 9 " 

Row of upper molars . " " 8 " 

Body length usually more than 190 mm usually less than 190 

Length of hind foot without claws " " " 30 " " n n 30 

Length of vibiissae " " " 60 " " n m go 

There are five subspecies of Ochotona hyperborea in the U. S. S. R., 
which are described below. 
41 GEOGRAPHICAL DISTRIBUTION. The Northern pika is widely 

distributed, mainly in the mountain taiga, in the boreal zone of Asia. Its 
northern boundary penetrates the tundra, and in the south it occurs in 
screes, as on the Hangay and in northwestern Mongolia. The range 
extends roughly between the borders of the Chukot Peninsula in the 
northeast to the Urals in the west; southern Altai, northwestern Mongolia, 
Hangay and Manchuria in the south. 

Let us now examine the distribution of this species over its wide range 
in detail. 

The Northern pika was first described by Pallas (1811) from the Chukot 
Peninsula. The "Vega" expedition caught pikas which (O. Nordquist (1883)) 
identified as the typical form. Specimens from Anadyr are found in the 
collection of the Zoological Museum of the Academy of Sciences and in my 
collection (collected by S.V. Kertselli). An excellent series of 27 
specimens of the Chukot pika from the vicinity of the village Markovo (on 
the Anadyr river) and from Cape Dezhnev was obtained by L.A. Portenko 
during his expeditions to Chukot Peninsula, where this species is widely 
distributed. L.O. Belopol'skii (in litt. 1936) reports that he found a colony 



28 



near Observation Cape (Anadyr Estuary) on a small hill in the talus zone 
of the bald peaks in the fall of 1930. Similar colonies were recorded near 
Cape Nerpichiy (Anadyr Estuary) and on hills between the Bay of the Cross 
and Anadyr Estuary in 1931. Similar colonies were recorded from the 
valley of the Anadyr river, near the estuary of the Belaya river, near the 
village of Markovo and in the estuary of the Yeropal river. Further 
west it occurs on the Kolyma river from where it was described by I. A. 
Allen (1903). O.kolymensis = O . hy pe r b o r e a . S.A. Buturlin 
found burrows and traces of pikas along the beach northeast of the village 
of Sukharnyi (near the northeastern part of the estuary of the Kolyma river), 
and frequently heard their voices onCape Epra-Ukun. He also found pikas 
in the tundra near the village of Pokhodskii (85 km from Nizhne Kolymsk 
alongthe Kolyma river). According to A. Bunge (1887) this species reaches the 
Yana and Adycha rivers north of 70°N. The collections of the Zoological 
Museum of the Academy of Sciences contain specimens from the following 
localities: Verkhoyansk Range; delta of the Lena, valley of the 
Olenek river; Adycha river. I.D. Cherskii also caught the Northern pika 
on the Indigirka. O. hyperborea is undoubtedly widely distributed 
throughout Yakutia. A.I. Ivanov (1934, in " Yakut skaya ASSR" (Yakutian 
Autonomous Soviet Socialist Republic)) records this species from the 
following districts: Lena-Khatanga; Lena-Yana; Indigirka-Kolyma. 
According to R. Maak (1886) it occurs along the middle and lower reaches 
of the Vilyui river to Suntar. It lives among rock outcrops along the 
banks of this river 50-60 kilometers above the village of Suntar (K.E. 
Vorob'eva). According to R. Maak, pikas live among rocks in the upper 
reaches of the Vilyui river. They were first found by R. Maak on his way 
to the Olenek river on the Tas plot on the bank of the Khaazngya river. 
In the collection of the Zoological Museum of the Academy of Sciences are 
specinmens from Yakutiya, from the right bank of the Amga river (opposite 
the summer camping site of Khokhur-Terde, estuary of the Khokhur river, 
and from the summer camping site of Ytyk); from the road from the Lena 
river to Burtul, and from the Eastern Kangalas district. I saw specimens 
in the ZMAN collection from the following localities in the Verkhoyansk 
district: Ken-Yuryakh; the bank of the Nel'chekhe river (atributary of the 
42 Adycha river); Tumara river (a tributary of the Aldan river); Nam District 
(Arbynskii community). 

Further south, in the southeastern part of the Lena river basin, it occurs 
in very large numbers along the Irbo, Choiongro and Yanude rivers, and 
also alongthe Yuzhno-Muiskii Range (I. Polyakov, 1873). To the east it 
is found alongthe Yablonovoi Range, where B.A. Kuznetsov (1929) caught 
an excellent series near Kyker. The expedition of B.S. Vinogradov and 
S.I. Obolenskii (1927) caught a series on the Alkhanai Mountain* south of 
the Bol'zino railroad station (former Chita Region). Further south it was 
found in the following localities: Khamar-Daban Range (specimen at the 
ZMAN collection); near the village of Gazimurski Zavod (S.I. Ognev's 
collection); the mountains in the vicinity of Okonon (Yablonovoi Range). 
In the east it occurs in the mountains of the Amur river basin. According 
to G.N. Gassovskii (1927) it is very numerous on the slopes of the 
Turkuringra and Gilyui ranges. S.I. Obolenskii sent several specimens 
from the Turkuringra Range to the Zoological Museum of the Academy of 



* A specimen from the Alkhanai mountain labeled by N. F. Kashchenko as O chotona sp. is also a 

Northern pika. (Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, vol. 17, 1912 (1913), p. 397-398.) 

29 



Sciences. R. Maak (1859) mentioned that it is frequently found on the Shipka 
river and in the mountains near the estuary of the Argun river. A. A. 
Emel'yanov (1927) found this species in the region of the Orochis along the 
banks of the Kop river. According to Middendorf (1867) it is widely 
distributed on slopes of the Stanovoi Range. It is also found in Kamchatka, 
and in the collection of the Zoological Museum of the Academy of Sciences 
are specimens from the Khalzan Mountains. My collection contains 
specimens from Ol'skii Island (Sea of Okhotsk), and from the coast of 
Penzhina Bay and the mountains near the village Penzhino. The Northern 
pika occurs farther into the mountains near the village Udskii Ostrog, and 
also in the mountains along the lower and middle reaches of the Amur 
river. In my collection are specimens from the vicinity of De-Kastri 
Bay. I saw specimens from the Ussuri Valley in the ZMAN collection 
(collected by V. K. Arsen'ev). N.T. Zolotarev (1936) mentions specimens 
from the Kolumba river. I have specimens from the Ussuri Territory in 
my collection. It is apparently common in some localities on the Sikhote- 
Alin Range. The ZMAN collection has specimens from Sakhalin Island. 
Dr. Suprumenko (1899) writes that the Northern pika lives on coastal cliffs 
and also on mountains on the island. One specimen was caught near Cape 
Aleksandrovskii (Dui-Zhonkier). 

The Northern pika occurs in Manchuria, and was found near Tsitsihar 
(vide infra), and also on the Great Khingan Mountains. Specimens caught 
by the American expedition 45 miles north of Ulan-Bator were identified 
by Gl. Allen (1927) as O. h. mantchurica. However, the local 
specimens are smaller than the real m an t c hu r i c a and form, apparently, 
a separate geographical race together with those from the Hentey and Hangay 
Mts. (Ohton-Tengri) (vide infra). 

Let us now describe the distribution of the Northern pika throughout east 
Siberia, from Transbaikalia and the watershed between the Lena and 
Yenisei rivers up to the Chukot Peninsula. The distribution in northern 
Siberia is as follows. 

O. hyperborea was found by N. P. Naumov (1934) in the Yenisei 
Basin in the mountains of the basins of the Lower Tunguska and Stony 
Tunguska rivers (Ugami trading station, vicinity of Baikit). In my 
43 collection are specimens from the Kureika river (caught by F. F. Shillinger). 
According to N. P. Naumov (1934) it lives along the Kureika and Khatanga 
(Kotui) rivers. It does not occur north of 71°N along the Khatanga. It is 
found in the upper reaches of the Volochanka river (near 69°30' N) and in 
the Norilsk mountains (at the same latitude) (S.I. Orlov, 1930). Along the 
Yenisei it does not occur north of the Khantaiskii settlement (68°30' N). 
Further south, was found in the Yenisei Basin by A. Ya. Tugarinov near 
Krasnoyarsk (the specimen is in the ZMAN collection), and also on the left 
bank of the Angara river. It was recorded from the river Ana (a tributary 
of the Abakan river in the western Sayan Mountains in the Minusinsk 
territory) (vide M.D. Zverev, 1930). The ranges of hyperborea and 
alp in a overlap both in the vicinity of Krasnoyarsk and in the Minusinsk 
Territory. It has not yet been found on the Altai. 

Nobody has found the Northern pika in the Tobolsk Region of western 
Siberia so far. The discovery of a litter in the upper reaches of the Voikar 
river (a tributary of tne Ob) by V. Yu. Fridolin (1925) was therefore of 
special interest. K. K. Flerov, who took part in the expedition of the 
Academy of Sciences to the northern Ural, found pikas on the eastern slope. 



30 



in the upper reaches of the Synya river (a tributary of the Ob). Specimens 
were also found in the valley of the Bol'shaya Kharuta river. This record 
is very interesting and considerably extends the range of this species to the 
west. 



44 




FIGURE 14. Northern pika, O chotona hyperborea uralensis 
Flerov 

Drawing by K. K. Flerov 

MODE OF LIFE. HABITATS. B.A. Kuznetsov (1929) found the Northern 
pika in Transbaikalia in the vicinity of the Kyker river in the mountain 
taiga of the main range of the Yablonovoi Range. He states: "This species 
inhabits screes exclusively. As soon as heaps of debris and detritus are 
deposited by the torrents of the spring waters, they become inhabited by 
pikas." N.P. Naumov (1934) reports from the Turukhansk Region: "Pikas 
occur mainly in screes either at peaks of ranges or in valleys, sometimes 
close to the water. They are more numerous on screes covered with shrubs, 
ferns, reindeer moss and other plants. In such places stones are hardly 
visible, as they are covered by a thin layer of soil. Pikas do not inhabit 
screes consisting of large, bare stones. However, stony areas are not the 
only habitat of pikas. I frequently found them in large piles of driftwood 
along river banks, in which conditions of life are very similar to those in 
screes. I found a large colony of pikas 7-8 km from the rock outcrops on 
the bank of the Yenisei river (along its lower reaches, 2 km from 
Turukhansk). They lived here among old driftwood. " K.K. Flerov gives 
the following interesting data on habitats of pikas on the Urals (1927): 
"Pikas inhabit all the gabbro screes in the northern Urals, They do not 
prefer forests or tundra. The deep taiga, shrubs on mountain slopes and 
bare spaces on peaks are inhabited by pikas to an equal extent. They live 
only on gabbro, never penetrating the dunite-peridotite massifs, probably 
because the gabbro screes are rich in cavities which provide many suitable 
hiding places. The Malyi [Lesser] Ural Range is therefore mainly inhabited 



31 



by them, as it consists entirely of gabbro. Pikas are the most widely 
distributed animals in the area; no other animal is as common. " 

The habitats of this species in Manchuria and on the Great Khingan 
Mountains are quite unusual. According to A.S. Lukashkin (in litt. ) pikas 
live there mainly in river valleys, along the banks and on small, arid 
mounds among tussocks. They occur rarely on bare slopes of mud 
volcanoes. A.S. Lukashkin found them in the floodland of the Nonni [Nun 
Kiang] river near the Tsitsihar [Angangki] station of Heilungkiang Province, 
which is an unusual habitat. 

In the northern tundra, e.g., in the lower reaches of the Kolyma river, 
pikas were encountered by S. A. Buturlin in screes (such as near Cape Enra- 
Ukun, and also among rocks along the right, raised, bank of the Kolyma 
river). Pikas are also found in the floodlands of the delta of the Kolyma 
river. S. A. Buturlin often saw pikas in the unflooded parts of the tundra 
during the spring floods. The animals ran about willow roots across 
puddles. In the Chukot Peninsula, L.O. Belopol'skii (1936) found colonies 
in the valley of the Anadyr river in cedar thickets along river banks (such as 
along the Ust -Belaya) or on slopes and islands covered with weeds and 
grass (near Markovo and in the estuary of the Yeropol river). 

GENERAL DESCRIPTION AND HABITS. The pika is a beautiful, lively 
animal. Its movements are elegant and it runs and jumps well, and has 
typical harelike behavior. K.K. Flerov (1927) states that the Ural 
pikas are extremely agile and alert; it is difficult for eagles and hawks to 
catch them unawares. At the slightest danger they instantly take to their 
burrows. He says: "They spend most of the time in their burrows. They 
emerge at regular intervals, leaping up on a stone and squeaking sharply. 
One animal's squeak is echoed by others, which sometimes stay inside 
their burrows and squeak for some time. This chorus then gradually 
ceases. Pikas normally squeak 15-30 times upon each emergence from the 
burrow. A frightened animal emits one or two short, shrill, sometimes 
nearly inaudible squeaks. The sound is sharp, shrill and ringing, 
resembling the call of the woodpecker. In order to "sing", the pika chooses 
a stone, usually the highest, which affords a full view of the surroundings. 
Sight is apparently as well developed as hearing. A person approaching them 
in the open against the wind is noticed from a great distance, and the pikas 
flee. The weather does not affect them much. They leave their burrows in 
rain, cold, and wind but they prefer quiet sunny evenings after the rain. " 

I can add that the Northern pika does not fear water (like many hares). 
S.A. Buturlin saw a pika swim across a small stream (about 10 m wide) in 
the Kolyma delta. 

DAILY ACTIVITY. According to K.K. Flerov, pikas of the northern 
Urals are always awake, their squeaks audible day and night. B.A. 
Kuznetsov (1929) observed in Transbaikalia that the Northern pika is mainly 
diurnal, though he heard its squeak repeatedly during the night. 

BURROWS AND NESTS. K.K. Flerov (1927) gives a detailed description 
of nests and burrows of pikas in the northern Urals: "Pikas live in large 
colonies, but each individual keeps to its own territory, usually one of the small 
46 separate screes which cover all slopes. Burrows are dug, and passages 
run in all directions among the stones with no definite system. Sometimes 
these passages are open ti-acks, sometimes they are concealed under stones 
and covered with earth, like tunnels. The nest is usually constructed among 
stones, in a crevice or a hollow. Sometimes one animal has several burrows . 



32 



45 



True burrows are dug in the forested taiga, among shrubs in river valleys 
where screes are covered with moss or earth. The passages of these 
burrows are flat, 9 cm wide and 6 cm high. Burrows are usually small. 




FIGURE 15. Postures of the Ural Northern pika, O c h o t o n a hyperborea uralensis Flerov 

Drawing by K. K. Flerov 

1-1.5 m long, without many branches but with several entrances, connecced 
by well trodden paths. The grass on the path is gnawed and removed. The 
paths usually run in depressions so that the animals are protected. They 
are 9-10 cna wide. Sometimes there are small open squares at the junction 
of paths or before entrances. On these squares pikas carry out all sorts of 
household chores, e.g., they stack the collected food, gnaw off long stalks 
before carrying them into the burrow, etc. Heaps of excrement may be 
found at a distance from the entrances, under stones or roots. They are 
found only rarely on the paths and in the burrows. " 

B. A. Kuznetsov (1929) observed the following in Transbaikalia: "In 
screes consisting of large rocks, pikas usually do not dig burrows, but 
build their nests of dry grass under the rocks. In screes consisting of 



33 



47 



small stones where the space between the stones is filled with alluvial 
soil, they dig deep burrows with several exits. " The ecology of the 
Manchurian pika is apparently quite specialized. A.S. Lukashkin (in litt. ) 
observed that it builds spherical nests of grass under shrubs or on tussocks 
in summer. More exact information on these habits would be very desirable. 

FOOD AND FOOD STORES. The pikas are purely vegetarian. Only 
once did K. K. Flerov (1927) find remains of spiders in the stomach of a 
pika. The preferred diet of pikas in the northern Urals is Ve rat rum 
lobelianum. They usually cut off the upper part of the stalk. The 
stalk is cut into pieces 5- 10 cm long, and carried to the square in front of 
the burrows. According to N. P. Naumov (1934) pikas in the Turukhansk 
territory feed on forest grasses, preferably chickweed (Stellaria). In 
Transbaikalia (on the Yablonovoi Range) B.A. Kuznetsov (1929) found 
dry twigs of the following plants in stored food: Filipendula, raspberry, 
river birch and Vi c i a cracca. According to I. Safronov (1926) food 
hoards of pikas in the region of the Ivanovsko-Arakhleiskie farms 
(Transbaikalia) consisted of twigs of the following plants: birch, larch, 
aspen, wild rosemary, black crowberry, raspberry and red bilberry. 
Like all other species ofOchotona, the Northern pika collects food 
hoards. Such hoards were observed in Transbaikalia by I. Safronov (1926) 
and B.A. Kuznetsov (1929). N.P. Naumov (1934) observed that in the 
Turukhansk territory pikas store food by placing small bundles of dry grass 
under roots of fallen trees, under stones etc. Food is stored at a distance 
from the passages between burrows. K.K. Flerov remarks that he never 
saw true "stacks" in the northern Urals. Crevices between stones 
occasionally contained heaps of dry grass, but there were no large stores. 
Flerov assumes that large stores are made only in late fall. 

REPRODUCTION. According to N. P. Naumov (1934) Northern pikas 
in Turukhansk territory have their estrus in late May or early June, 
apparently before the molt, but sometimes at the same time. A male 
caught on 8 June 1930 on the Tura (Lower Tunguska) still wore its winter 
fur, but molt was at its height (the fur fell out in tufts). The testes of this 
specimen were much enlarged. According to observations in Transbaikalia, 
B. A. Kuznetsov (1929), assumes that the Northern pika of the Yablonovoi 
Range has two summer broods. Three embryos 45-48 mm long were found 
in the uterus of a female killed on 23 July. At the same time young 
specimens 14-15 cm long were found. My collection contains some very 
young specimens, caught by G. D. Dul'keit on Aug. 17, 1928 on Ol'skii island 
(Sea of Okhotsk). 

MOLT. N. P. Naumov describes the molt of the Northern pika as follows: 
"The spring molt in O . hype rbor e a occurs very late. A specimen in 
molt was caught on Jun. 8, 1930. There is a specimen in the ZMAN collection 
which had just begun its fall molt (caught on Jun. 27, 1928), and a similar 
specimen caught on Jul. 15, 1927 (Verkhoyansk District). I found completely 
molted specimens only in July (Jul. 7, Lower Tunguska). The molt takes 
place in the following sequence. The fur of the neck is shed first. The 
molted part surrounds the entire neck, later proceeding to the legs. At 
the same time the eyes become ringed (No. 18658, Jun. 27, 1928, Verkhoyansk, 
and No. 16733, Jul. 15, 1927, Verkhoyansk district, ZMAN collection). The 
molt then spreads over the muzzle to the ears, descending from neck to 
back, sides, and flanks to the buttocks. Occiput and the lower part of the 
back still bear winter fur (specimen No. 141, 1928, collection No. 255, 



34 



Aug. 7, 1927, Verkhoyansk District, ZMAN collection). The occiput and 
lower part of the back molt later. Winter fur remains on the ears and 
buttocks, which are last to molt. Judging from the material in the ZMAN 
collection, the fall molt goes in the opposite sequence, beginning in early 
September. " 

Examination of specimens from the Anadyr Territory revealed that the 
fall molt takes various and different sequences. The anterior part of the 
body, head and shoulders sometimes molt first. Sometimes the lower part 
of the hind quarters molts at the same time. The back always molts first 
and the flanks later. Specimens caught in mid- and late August were in the 
middle of the molt. Pikas caught in mid-September bore an almost complete 
winter fur; the series from Anadyr, however, contains specimens still 
bearing summer pelage in October. 

The fall molt of females apparently takes place later than that of males. 
48 The same was observed in the Alpine pika. 

WINTER LIFE. According to N. P. Naumov (1934) burrows of pikas 
in the Turukhansk territory are covered with snow, and no life is observed 
on the surface. All data in the literature indicate that the Northern pika 
does not hibernate. R. Maak (1859) observed in the Am^ur territory that 
pikas do not hibernate, coming to the surface of the snow during severe 
frost. He found a pika at a temperature of -15°. Pikas in the Kolyma- 
Anadyr Territory also do not hibernate during the winter. 



315. Ochotona (Pika) hyperborea hyperborea Pall, (1811). 
Northern pika 

1811. Lepus hyperboreus Pallas P. , Zoographia Rosso- Asiatica, p. 152, 

1858. Lagomys hyperboreus var. normalis Schrenck L. , Reisen imd Foischungen im Amur Lande, 
I, S, 148, VII, Fig, 1. 

1882. Lagomys litoralis Peters W. , Sitzungsb, der Gesellschaft Naturforsch. Freunde zu Berlin, S. 95 
(near Aran, on Cape Chukotski). 

1883. Lagomys hyperboreus Nordquist O. , Anteckninger och studier till Sibirska ishafskustens 
D^ggdjursfaima, S. 83 (Berings Sund). 

1903. Ochotona kolymensis Allen J. , Report on the Mamm. Collect, in North- East. Siberia, Bull. 

Amer. Mus. Nat. History, XIX, pp. 154- 155. (Upper Kolyma). 
1926. Ochotona hyperbore a Ognev, S. I. , Mlekopitayushchie Severovostochnoi Sibiri (Mammals of 

Northeastern Siberia), pp. 70-71. 
1926. Ochotona kolym ensis Ognev, S, 1. , Loc, cit. , pp. 68-70. 
1926. Ochotona litoralis Ognev, S.I,, Loc. cit,, pp. 71-72. 

TYPE LOCALITY AND DEPOSITION, The "Zoographia Rosso-Asiatica' gives the following indication: 
"Plura ejus sjjecimina et pelles e terris Tschuktschicis, ubi hyeme collecta fuerunt". We should therefore 
consider the Chukot Peninsula as the type locality for the nominal race. It is not kn own whether the type 
has been preserved. 

MATERIAL EXAMINED: 32 pelts (most of them with skulls), 

DIAGNOSIS. General tone of summer fur on back rather light, grayish 
brown with a slight straw- rusty tinge. Rust tones (sayal-brown) extending 
to lower part of back. Flanks much lighter and more yellow than back 
(with a gray tinge). Belly white with straw-yellow tinge. 

Winter fur on back uniformly colored, pale, grayish brown with a yellow 
tinge. Flanks with a purer straw-yellow tone. Belly lighter than flanks. 



35 



49 



This subspecies is relatively small. Length of skull 35-40.7 mm (M. 
37.3); condylobasaf length 32.8-38.3 mm (M. 34.4); zygomatic width 1 7.3- 
20.7mm (M. 18.5): width of skull at level of auditory meatus 17.2-21.3 mm 
(M. 18.5); length of row of upper molars 6.1-8 mm (M.6.7). Body length 
119-160mm (M. 140); hind feet 20-23. 5mm (M. 21.8). 

ADDITIONAL DESCRIPTION. General tone of back of adult 
specimens rather light grayish brown with a slight straw -rusty tinge. 
Rusty tones (sayal -brown, Ridgway, 1912, pi. XXIX) well marked on 
lower part of back; upper part of back more straw -yellow (buffy- 
brown, Ridgway, pl.X). Top of head, cheeks and area around eyes 
with an ash-gray tinge. Flanks much lighter and more yellow than 
back (cinnamon-buff with a gray tinge). Belly whitish with a distinct straw- 
yellow tinge (pale pinkish buff and pinkish buff). Feet white. 

Anterior inner margin of ears covered with blackish gray hair. Inner 
surface of ears straw-white; posterior parts of ears dull gray. 

Color of summer fur of young specimens paler, and sometimes more 
reddish than in adults; hair of young specimens shorter and thinner. 
Young specimens are easily recognized by their short, very soft fur. 

WINTER FUR. Top of head and back of adults uniformly colored, 
pale, grayish brown-yellow. Flanks with distinct, straw-yellow tones 
with a slight grayish tinge. Belly slightly paler than flanks. 

SYSTEMATIC NOTES. O. h yp e r b o r e a from the Chukot Peninsula 
is undoubtedly identical with O . litoralis which Peters (1882) described 
from the Avana river on the Chukot Peninsula. Peters apparently described 
young specimens bearing fur transitory between the summer and winter 
fur. 

O. kolymensis J. Allen (from Verkhne Kolymsk) is identical with 
O. hyperborea. The former was described by J. Allen (1903) from 
two alcohol specimens. The Kolyma pikas do not differ externally from 
those found on Chukot Peninsula. I had a large series of Chukot pikas and 
only a few from Kolyma. A comparison of skull measurements of these 
two forms showed that the maxima of the Kolyma and Chukot specimens are 
equal, but the skull of the Kolyma form is usually slightly larger. This 
should be verified on the basis of more material. 

GEOGRAPHICAL DISTRIBUTION. The range of pikas on the Chukot 
Peninsula extends from its northeastern parts to the Kolyma Basin, 
apparently to the Indigirka Basin. The Verkhoyansk Territory is inhabited 
by another subspecies. 

316. Ochotona (Pika) hyperborea ferruginea Schrenck (1858). 
Russet Northern Pika 

1858. Lagomys hyperboreus var. lerruginea Schrenck L. Reisen und Forschungen im Amur Lande, 

1,S. 148-149 (South Kamchatka, Khalzan Mountains). 
1887. Lagomys alpinus Bunge A. , Berichte iiber die Expedition nach Neusibirisch. Ins., S. 40 (nee 

Lepus alpinus Pall.'). 
1922, Lagomys kamtschatica Dybowski B. , Arch. Tow. Nauk. Lwowie, III, p. 10 (nomen nudum! ) 
1934. Ochotona (Pika) hyperborea turuch ane nsis , Naumov, N. P. , MIekopitayushchie 

Tungusskogo okruga (Mammals of the Tungus District), Trudy Polyamoi Komissii Akademii Nauk, Issue 

17, pp. 38-42 (Lower Tunguska, Uchami trading station). 

TYPE LOCAUTY AND DEPOSITION, Kamchatka, Khalzan Mountains. Type at the ZMAN collection. 

MATERIAL EXAMINED: 57 pelts (most of them with skulls). 



36 



50 



DIAGNOSIS. General color of summer fur on back rich, rust-red-ocher, 
more intensely red-rust-brown along middle line of back (mikado -brown); 
black hair tips markedly developed on midline of back, sometimes forming 
an indistinct blackish stripe on backbone. Top of head with deep chestnut 
tones. Flanks more yellow-ocher (cinnamon). Belly very pale, reddish 
ocher-yellow. 

Winter fur pale, dirty-yellowish gray (intermediate between drab and 
buff, with a tinge of deep chestnut); middle line of back slightly darker as 
a result of development of dark chestnut hair tips. Top of head and nape 
with rich ash-gray tinge, contrasting to color of back. Flanks rust-red 
with an olive tinge, intermediate between tawny- olive -clay (Ridgway, 1912, 
pi. XXIX). Belly dull whitish yellow. 

This subspecies is larger than the typical form. Skull length 38-41 mm 
(M. 40). According to N. P. Naumov (1934), the skull length of O . h . 
t ur uc hane ns is reaches 43.2 mm; condylobasal length 37-38.6 mm (M. 37) 
(according to Naumov to 40.2mm); zygomatic width 19.03-21 mm (M, 19.9); 
width of skull at level of auditory meatus 19-21.2 mm (M. 19.8); length of 
row of upper molars 7.2-7.3 mm (M. 7.3). 

SYSTEMATIC NOTES. The North Siberian pika (with the exception of 
those in the Kolyma-Anadyr Territory) has a rich, saturated color. L. 
Schrenck (1858) described the Kamchatka population as a separate richly 
colored "variation" offerruginea (according to specimens collected 
by Voznesenskii). I could not find any difference between the Kamchatka 
specimens and a series from, the Verkhoyansk Range (Olenek river), from 
Baikit (Lower Tunguska), and also from other adjacent localities in 
Siberia. They are very similar in coloring. 

N. P. Naumov (1934) described a subspecies, Ochotona hyperborea 
turuchanensis, from specimens from Uchami (Lower Tunguska) and 
from Baikit (Stony Tunguska). Its color is very similar to that of 
ferruginea. N. P. Naumov (1934) compares t ur uc h ane n s i s with 
O. svatoshi and O.h. uralensis, but he omits the Kamchatka 
ferruginea. This form differs from the typical and other forms in the 
same characters as turuchanensis . 

We regard t uruchanensis as synonymous with ferruginea; however, 
there is very little material on Kamchatka pikas. It is therefore possible that 
differences between the Kamchatka and continental Ochotona will be 
found when a large series is examined. 

GEOGRAPHICAL DISTRIBUTION. In addition to the Kamchatka pikas, 
specimens from the following localities belong to this subspecies: coast of 
Penzhina Bay; Verkhoyansk Range; mountains along the right tributaries 
of the Aldan river (e.g. , onthe Tumara river); the Khandyk river (a right 
tributary of the Alden river); eastern slopes of the Verkhoyansk Range*; 
the delta of the Lena (the village of Bulun, sent by Romanov to the ZMAN 
collection); the Olenek river (collected by R. Maak); the Chunya river 
(100 km from its estuary); Lower and Stony Tunguska. Thus, the richly 
colored O.h. ferruginea is distributed throughout the northern region 
of eastern Siberia except in the Kolyma-Anadyr Territory. 



* Specimens from the right bank of the Anga river (a tributary of the Aldan river) belong to this form. The 
ranges of O.h. ferruginea and O. h. cinereoflava (vide infra) lie very close together here, but 
the boundaries have not been exactly determined. 



37 



51 317. Ochotona (Pika) hyperijorea cinereoflava Schrenck 

(1858). Okhotsk Northern pika 

1858. Lagomys hyperboreus var. cinere of 1 a va Schrenck L. , Reisen und Foischimgen im 

Amur Lande, I. S. 150, Pl.VlU, fig. 1. 

TYPE LOCAUTY AND DEPOSITION. No, 689, 18 Feb. 1845, Udskii Ostrog, A. Middendorf. Type in 
ZMAN collection. 

MATERIAL EXAMINED: 26 pelts and skulls. 

DIAGNOSIS. General color of summer fur on back rich yellowish gray 
with a slight rusty tinge. Top of head with dark brown tones, especially 
marked on the hair tips. Indistinct pale yellow on occiput. Flanks with rich 
yellowish rusty tones. Belly yellowish with a slight ocher tinge. 

Winter fur of back generally yellowish gray with an ocher tinge. Black- 
brown hair tips little marked, so that the color is rather light. Color of 
head slightly grayer than back; the ground color is yellowish gray with 
brown hair tips. Flanks more yellow than back; belly whitish with a light 
yellowish tinge. 

Skull length 40-42 mm (M. 40.8); condylobasal length 37-39.21 mm 
(M. 38.1); zygomatic width 20-20.7 (M. 20.3); width of skull at level of 
auditory meatus 19.2-19.6 mm (M. 19.1) [sic]; length of upper tooth row 7.3- 
7.6mm (M.7.4). 

SYSTEMATIC NOTES. This subspecies is easily distinguished in series 
from other subspecies ofO. hyperborea by its light yellowish gray 
color with a slight rusty tinge, and also by the light, pure color of the 
flanks . 

GEOGRAPHICAL DISTRIBUTION. This subspecies was described from 
a specimen from the vicinity of the Udskii Zavod. In L. Schrenck's 
opinion (1858) the same form lives along the Maya and Olenek rivers. 
Richly colored forms of the ferruginea type live in the basin of the 
Olenek river. I examined a series of 19 specimens of this subspecies in 
the ZMAN collection from the former Yakutsk District caught in the 
vicinity of the Yelanskaya station (on the Yakutsk-Irkutsk highway). 

The light color of pikas from Ol'skii Island (collected by G. D. 
Dul'keit) shows that they belong to cinereoflava (these specimens are 
in my collection). This light-colored form apparently inhabits the central 
and northern zones of the coast of the Sea of Ochotsk, but the same light 
colored form occurs in Central Yakutia. The lower Amur basin is inhabited 
by another form which does not differ from the Transbaikal form. 

318. Ochotona (Pika) hyperborea uralensis Flerov (1927). 
Northern Ural pika 

1927. Ochotona hyperborea uralensis Flerov K. K. , Pishchukha Sevemogo Urala (The Northern 

Ural pika). - Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, Vol. XXVIII, pp. 138-144, fig. 1-3; 

pp. 141-142, pi. IV, fig. 1-4. 

TYPE LOCAUTY AND DEPOSITION. No. 16107 9 10 August 1926, eastern slope of the Ural Range, 
valley of the Bolshaya Kharuta (Syn'ya river), K. K. Flerov. Type at ZMAN collection. 

MATERIAL EXAMINED: 4 pelts with skulls. 



38 



52 DIAGNOSIS. General color of summer fur on back yellowish-rust, 
darkened by blackish brown hair tips*. Top of head with a distinct dark 
brown tinge; fur on occiput more yellowish. Color of flanks and cheeks 
rather light, ocher-yellow. Belly pale, ocher-yellowish gray. 

Color of young specimens paler and grayer than in adults, without 
ocher shades. 

Skull length 37-39 mm (M. 38); condylobasal length 35.3-36.6 mm (M. 
38.1); zygomatic width 18-20mm (M. 19.3); width of skull at level of auditory 
meatus 18. 8-19. 7mm (M. 19.2); length of upper tooth row 7.1-7.5 mm (M. 
7.3). 

Body length 167- 183 mm (M. 175.5); hind feet 22-27 mm (M. 25). 

SYSTEMATIC NOTES. The Northern Ural pika differs clearly from 
the central and south Siberian pika (vide infra) in the lighter color of 
its summer fur which lacks the saturated rust-red-ocher tones of 
ferruginea. According to its general color the Ural pika is closely 
related to O . h . cinereoflava. But more material is necessary to give 
an exact description of the Ural pika and its characters as given in K. K. 
Flerov's work. The small number of syntypes in K. K. Flerov's collection 
did not make it possible to give a more accurate description of the Ural 
pika. 

GEOGRAPHICAL DISTRIBUTION. K. K. Flerov (1927) found this 
subspecies on the eastern slope of the northern Ural along the upper reaches 
of river Synya (a tributary of the Ob) and along the river Lyapin (a 
tributary of the Sosva). V. Yu. Fridolin (1925) collected a litter along the 
upper reaches of the Voikara river (a tributary of the Ob). 

319. Ochotona (Pika) hyperborea mantchurica Thos. (1909). 
Central Siberian Northern pika 

1859. Lagomys hyperboreus Maak, R. , Puteshestvie na Amur ( A Journey along the Amur). — St. 

Petersburg, p. 108. 
1862. Lagomys hyperboreus Radde G, , Reisen im Siiden von Ost-Sibirien, S. 232-233 (partim.'). 
1873. Lagomys alpinus Polyakov, I., Geograficheskoe rasprostranenie zhivotnykh v yugo-vostochnoi 

chasti Lenskogo basseina (The Geographical Distribution of Animals in the Southeastern Part of the 

Lena Basin). — Zapiski Geograficheskogo obshchestva, III, pp. 52-53, 
^909. Ochotona hyperborea m antchurica Thomas O. , A Collection of Mammals from Northern 

and Central Manchuria, Ann. and Magaz. Natur. History, ser. 8, pp. 504-505. 
1927. Ochotona cinereofusca Vinogradov, B.S. and Obolenskii, S.I,, Materialy po faime gryzunov 

yuzhnoi chasti Eniseiskoi i Irkutskoi Gubemii i Zabaikal'ya (Material on the Fauna of Rodents of the 

Southern Part of the Yeniseisk and Irkutsk Provinces and Transbaikalia), Izvestiya Sibiiskoi stantsii zashchity 

rastenii. No. 2, p. 15 (taiga regions of Transbaikalia, nee. cinereofusca Schrenck.'). 
1927. Ochotona hyperborea m antchuric a Allen G, , Lagomorphs Collect, by the Asiatic Exped. , 

Amer, Mus. Novitates, No. 284, pp. 1-2 (a series of 30 specimens caught 45 miles north of Urga). 

1929, Ochotona (Pika) hyperborea Kuznetsov, B, A. , Gryzuny vostochnogo Zabaikal'ya (Rodents 
of Eastern Transbaikalia), Izvestiya Assotsiatsii Nauchno-issledovatel'skikh institutov, 2 (l):86-87. 

1930. Ochotona hyperborea Zverev M. D. , Obzor gryzunov yuzhnoi chasti Minusinskogo i Khakasskogo 
okrugov (A Survey of Rodents in the Southern Part of the Minusinsk and Khakassian Districts), Sbomik 
Srednesibirskogo geograficheskogo obshchestva, p, 85 (River Ana, a tributary of the Abakan, western 
Sayans). 

TYPE LOCALITY AND DEPOSITION: No. 23 d", 18 May 1908, Great Khingan, British Museum. 
MATERIAL EXAMINED: 87 pelts, mostly with skulls. 



* K. K, Flerov (1927) describes the fur of the Ural pika as "reddish". This seems not as accurate as the 
description given here. 



39 



53 DIAGNOSIS. General color of summer fur on back relatively dark, 

brown-chestnut, often with a strong development of black hair tips. Flanks 
rust-brown (cinnamon, Ridgway, pi. XXIX) with a gray tinge. Belly dirty- 
rust-gray. 

Winter fur on back dirty-grayish brown, straw-brownish gray on flanks 
and dirty-gray with a straw-white tinge on belly. 





FIGURES 16,17. Skull of Central Siberian Northern pika, Ochotona hyperborea mantchurica 
Thos. 593 $ 16 July 1928. VicinityofKyker, SretenskDistrict, Transbaikalia. B. A. Kumetsov 

Drawing by Yu. A. Kostylev 



54 



Skull length in fully grown specimens 39.3-43.3 mm (M.41); condylo- 
basal length 36.7-40.2 mm (M. 38); width of skull at level of auditory 
meatus 19.2-21.1 mm (M. 20.3); length of upper tooth row 7-8 mm (M, 7.4); 
body length 150-195mm (M. 172.8); hind feet 20-30mm (M. 24.9). 

ADDITIONAL DESCRIPTION. It should be added that the color of the 
summer fur varies in different specimens. In light colored specimens the 
back generally has a rather intense chestnut-rust-brown color (closely 
similar to snuff-brown, Ridgway, 1912, pi. XXIX), but has a slight rusty 
tinge. Top of head of the same color. Color of flanks lighter and purer, 
with a well marked rusty tinge (intermediate between cinnamon and sayal- 
brown). Ground color of belly lead-gray with a marked rusty tinge, which 
is especially marked in the middle of the belly. 

Specimens in which the summer fur has strongly developed black hair 
tips, especially along the midline of the back and on the head, occur 
together with light colored specimens. These specimens generally have a 
dark, blackish chestnut-brown fur. Flanks more rusty than back, but also 
darkened by well marked blackish brown hair tips. Belly lead-gray with a 
rusty tinge. 



40 



Color of winter fur on back more intense than in other subspecies of the 
Northern pika; its gray ground color is mixed with chestnut-brown. 




FIGURE 18. Skull of Central Siberian Northern pika, Ocho ton a hyperborea 
mantchurica Thos. , lateral (same specimen as in Figures 16, 17) 



SYSTEMATIC NOTES. O.Thomas (1909) described O. h. mantchurica 
from 10 summer specimens from the Great Khingan. The color of these 
specimens was rather pure, without any marked blackish shades. According 
to Thomas the ground color of this subspecies (summer fur) is intermediate 
between cinnamon and russet. He also states that the measurements of 
O.h. mantchurica are generally maximal for the species 
hyperborea: skull length 43.4 mm; condylobasal length 41mm, 
maximum width 22 mm; nasals 14.2 mm long; interorbital distance 4.3 mm; 
bony palate 15.5mm long; palatine foramen 6.4mm; row of upper molars 
(measured along alveoli) 8.1mm long. 




FIGURE 19. Mandibula of Central Siberian Northern pika, Ochotona 
hyperborea m antchuric a Thos. , (same specimen as in Figures 16-18) 

Body length 178mm; hind feet 28.5mm; ear 18mm. 

When I first examined the material I had the impression that O.h. 
mantchurica differs from Transbaikalian and Amur Northern pikas, 
and that it is closely related toO. alpina in some characters. After 
comparing the material I found quite similar specimens from the Yablonovoi 
Range, Great Khingan, and also from near Krasnoyarsk! 

In general, the following tendency was noted. The color becomes deeper, 
less rich and the animals larger from the north of Siberia (Verkhoyansk 



41 



55 Territory, rivers Lena, Yana, Olenek, etc.) southwards. The differences 
between typical O.h. hyperborea from the eastern parts of the Chukot 
Peninsula and O.h. mantchurica Thos . from the type locality (Great 
Khingan) are especially marked. These two forms are antipodal, if their 
large range is taken into account. 

However, these forms have gradual series of intermediate forms like 
real subspecies. These intermediate forms obliterate the sharp differences 
between subspecies from far removed localities. 

GEOGRAPHICAL DISTRIBUTION . This subspecies occurs in the Yeniseisk 
Territory, in the taiga near Krasnoyarsk, western Sayans (river Ana, a 
tributary of the Abakan), near Lake Hobsogol in the Irkutsk District, the 
Vitim Olekma upland, Transbaikalia (Yablonovoi Range, Gazimur Mountains) 
the Amur and Ussuri territories. A.S. Lukashkin (in litt. ) reports that 
this subspecies occurs on the eastern slopes of the Great Khingan (e.g., 
near the Yalu station of the Chinese Eastern Railway. It was also found 
in the floodland of the river Nonni [Nun Kiang] near the Tsitsihar [Angangki] 
Station in the Heilungkiang Province. 

G. Allen (1929) received a series of pikas caught 45 miles north of Urga 
(Ulan- Bator). He identified them as O.h. mantchurica but pointed out 
that they are smaller. I saw a series of 10 pikas from southwestern 
Hangay, from Ohton-Tengri in the ZMAN collection. The same collection 
also has specimens from Hereleng Gol river (eastern Hentey). All these 
specimens differ from O . h . m ant c hur i c a in their light, pure, rust- 
yellow color similar to that of Central Yakutian and North Ural specimens. 
It is probable that this is a separate geographical race, which should be 
carefully studied on a large material. 



Subgenus Ochotona Link 

1795. Ochotona Lin, H. F. , Beitrage zur Naturgeschichte, I, Abt. II, S. 52, 74. 

Skull of most species elongate, slender, arched in the region of the 
interorbital space and frontals. Fenestrae frontales absent. Interorbital 
space narrow. Orbits relatively large. Structure of palatine and incisor 
foramina varying: palatine and incisor foramina in O. daurica, 
curzoniae, rufescens fused and vomer ventrally free; in O . 
ladacensis and O . pr i c e i these foramina are separate, and the vomer 
is ventrally covered by the premaxillaries. Fenestrae triangulares rostri 
small and rounded. 

Color light, straw-sandy, reddish sandy, brownish in summer fur and 
gray in winter fur. Pads of digits not markedly developed. 

Vibrissae short, not more than 60mm long. 

Species of this genus live in open landscapes, steppes, deserts or high 
plateaus. 

Type of subgenus: Ochotona minor = O. daurica Pall. 
56 GEOGRAPHICAL DISTRIBUTION. The subgenus occupies a wide range, 
from central Iran in the west, Iranian Baluchistan, mountainous Turkmenia 
(Kopet-Dagh), Afghanistan, and further north from the Chuya river to 
northern and central Mongolia, to the Daurian steppes. The range of the 
subgenus also includes Ladakh, Nepal, Sikkim, Seistan and Tibet. 



42 



Ochotona curzoniae Hodgson, O.ladacensis Giinther, O. 
rufescens Gray, O. price i Thos. also belong to this subgenus. 

151. Ochotona (Ochotona) daurica Pall. (l776).Daiirian pika 

1776, Lepus dauuricus Pallas P. , Reise durch verschiedene Provinzen des Russischen Relchs, III, S, 692, 

NAMES: Daurian pika (English); Otogono or Ogotona (German). 

LOCAL NAMES: pishchiikha-senokoska [hay-cutting pika]; kuyun; "kamennaya koshka" [stone cat]; 
"kamennyi krot" [stone mole]; ogotona (Transbaikalian Cossacks); ogotona (Mongolian); Umuki (Evenki, 
Tungus). 

PRINCIPAL FIGURES: 1) Pallas P. S. Novae species Quadrupedum e Glirium ordine, 1778, pp. 59-70, 
tab. Ill (whole animal), tab. IV A, fig. 16 A and 16 B (skull front and lateral); 2) Schreber I. Ch. , Die 
Saugethiere, Erlangen, IV, 1792, S. 915-918, Tab. CCXXXIX (in color); 3) Simashko Yu. , Russkaya fauna 
(Russian Fauna), Vol.11, Mlekopitayushchle, 1851, pp. 793-794, pi. 63, fig. 3 (in color) ; 4) Bikhner, E. A. 
Nauchnye resul'taty puteshestvii N. M. Przheval'skogo (Scientific Results of N. M. Przheval'skii's Journeys), 
otdel Zoologicheskii, Vol.1, Mlekopitayushchle, Issue 5, 1894, pi. XXV, fig. 1-5 (photograph of skull), 
pi. XXII (whole animal in color); 5) Kobel't V., Geograficheskoe rasprostranenie zhivotnykh (Geographical 
Distribution of Animals), 1903, p. 209 (whole animal, half tone plate); 6) Brehm's Tierleben, Saugetiere, 
II, 1914, Taf. S. 15 (color plate); 7) Formozov, A. N. , Mlekopitayushchie severnoi Mongolii (Mammals of 
Northern Mongolia), 1929, p. 85, fig. 13 (head), p. 86, fig. 14 (skull ventrally); 8) Vinogradov B. S. , 
Mlekopitayushchie SSSR (Mammals of the U. S. S. R. ), I, 1933, p. 81, fig. 74 (B) (skull dorsal), pLVIII, 
fig. XV (photograph of entire animal). 

MATERIAL EXAMINED: 47 pelts, mostly with skulls. 

DIAGNOSIS. Skull rather large, its length exceeds 40 mm in fully grown 
specimens. Interorbital space narrow, 3.2-4.2 mm; surface of interorbital 
space convex, with two weakly marked converging crests in the middle of 
interorbital space in old specimens. Nasals slightly widened anteriorly, 
without constriction at the base of the wider part. Lateral contours of 
nasals narrowing posteriorly, their posterior ends rounded. Frontals 
without fenestrae frontales (vide Figure 20). 

Vomer completely free ventrally, not covered by premaxillaries; 
palatine and incisor foramina fused (Figure 22). 

Most vibrissae white, a few with a black-brown tinge, 40-55 mm long. 
Soles either pure white, or gray or whitish brown. Ears with distinct 
white borders. 

Color of summer fur on back varying in different subspecies from pale, 
yellowish straw-gray to duller and deeper straw-gray. Flanks light and 
yellow; belly pure white, hair bases gray. 
57 Winter fur differs markedly from summer fur in its long and lighter 
hair, and gray color with a straw-yellow tinge. 

Body length to 195 mm; hind feet 31mm. 

ADDITIONAL DESCRIPTION, In addition to the above-mentioned 
characters, the orbits are rounded (not oval as in O . rufescens, the skull 
of which resembles that of daurica); base of triangular fenestrae 
relatively low, elongate longitudinally. 

The base of the triangular fenestrae ofO. rufescens lies higher, 
is broader and shortened in the longitudinal direction. 

Bullae osseae rounded and slightly flattened laterally. The outer contours 
of the sides of the bullae are more distinctly flattened inO. rufescens. 
Rostrum of moderate length, less elongate than inO. rufescens. 

The skull structure varies markedly according to age. Rostrum of young 
skull markedly abbreviated. The distance between the inner margin of the 
alveoli of the posterior molars is much greater in very young specimens 
than the length of the upper diastema, as it is in fully grown and old 



43 



specimens ofO. pusilla, in which the lower margins of the 
premaxillaries are much closer together than in adults of the Daurian pika. 

In spite of the converging margins of the 
premaxillaries, the vomer is clearly 
visible from beneath, completely separate 
from the margins of the premaxillaries 
which lie close to it. 

The structure of the bony palate is 
similar in general to that ofO. pusilla. 
The bridge of the bony palate which 
consists of the pterygoids and palatines 
is broader than in adults. Bullae osseae 
more rounded, swollen and short. The 
brain case is very rounded and short. 
When skulls of young and adults in profile 
are compared the parietal bones of the 
former appear swollen while those of O. 
pusilla are more flattened and slant 
posteriorly. 

SYSTEMATIC NOTES. The Daurian 
pika is clearly distinguished from related 
forms by a number of characters. It 
includes the following subspecies: the 
typical subspecies O. d. daurica (from 
Dauria); O. d. altaina Thos. (vicinity 
of Lake Achita Nur, to the southeast of 
the Saylyugem Range in northwestern 
Mongolia); O.d.bedfordiae Thos. 
from Shansi (Ningwufu) . O.huangensis 
from Kansu is apparently closely related 
to O. daurica, and is possibly only a 
subspecies of this species. The close 
relation between huangensis ani 
daurica was pointed out by O. Thomas 
(1908). The description of O . 
huangensis by P. Matschie (Wissenschaftliche Ergebnisse der Expedition 
Filchner, Bd. 10, 1908, S. 214-217) is very vague although it seems 
reliable. The material used by the author was also in bad condition. 

GEOGRAPHICAL DISTRIBUTION. N. F. Kashchenko in his "Key to 
Mammals of the Tomsk Territory" (Opredelitel' mlekopitayushchiksh 
Tomskogo kraya) (1900, pi. 30) states that O. daurica does not occur 
in the Tomsk Territory and in West Siberia. In his opinion, Brandt's 
record of O . daurica in West Siberia is erroneous. The northwestern 
range of the Daurian pika includes the southeastern corner of West Siberia. 
This species occurs in the Altai Chuya Alps, where a gray pika was caught 
in summer 1936 by A. M. Kolosov in the Chegan-Burgazy region. In my 
collection are several specimens from the Gorno-Altai Autonomous Region 
caught in the vicinity of Toshakty (1934). All these specimens belong to 
O. d. altaina Thos. which was described by O. Thomas from specimens 
from the vicinity of Lake Achita Nur and from Suok (on the Altai). The 
same author later recorded O. d. altaina from the vicinity of Lake Ureg 
Nur (north of Achita Nur). There are specimens from Hobdo [Jirgalanta] 




58 FIGURE 20. Skull of Daurian pika, 

Ochotona daurica daurica Pall. 
No. M.3997 9June 1925. Near Borzya 
Station of the Chita Railway. E. S. 
Vinogradov and S. I, Obolenskii, S. I. 
Ognev's collection 

Drawing by Yu, A. Kostylev 



44 



59 



in the ZMAN collection. A.N. Formozov (1929) recorded the Daurian pika 
from the vicinity of Lake OrogNur (45° N) on the eastern slopes of the Great 
Altai. This species also occurs near Uliassutai [Jibhalanta] on Lake HobsOgOl, 
inhabiting the valleys of rivers Onshin Gol, Tuin Gol, Baydarag Gol, 
Dzabhan Gol and the basin of Lake Dzhargalante. G. Allen (1927) mentions 
that O. daurica occurs south of Ulan -Bator (Urga, near Turin). 
Apparently, O. daurica is widely distributed throughout northern 
Mongolia to Lake Dalai-Nor, from where an excellent specimen was sent 
tomebyA.S. Lukashkin. G. Radde (1862) caught a large series near Tarey Nor. 
R. Maaik (1859) recorded it from the steppes of the valley of river Onon 
and from the vicinity of the Mokhotui frontier post. N. F. Kashchenko 
(1913) collected a series in the Aga Steppe, and also recorded O. daurica 
from the former Troitskosavsk [Kyakhta] and Verkhneudinsk [Ulan Ude] 
counties (1910). B. S. Vinogradov and S.I. Obolenskii (1927) found this 
species in the vicinity of Troitskosavsk [Kyakhta] and Durena, near the 
village Mukhor-Shabar. A.N. Formozov (1929) writes that this rodent 
is so numerous on the Selenga river near Ust -Kyakhta and Troitskosavsk 
[Kyakhta] that in some places, its fecal pellets are scattered in all 
depressions of the ground. 

Detailed information on the distribution of pikas in Dauria is given in an 
article by A.S. Fetisov (1936):"The Daurian pika inhabits all the steppes of 
the so-called Selenga Dauria. This region lies along the valley of the river 
Selenga. It is found in almost all the steppes and forest steppes of the 
Selenga and Dzhida regions. The Daurian pika reaches, at present, the river 
Ulikchin in the west along the Dzhida river. It is not found in the Trans - 
Kamensk District. This species is frequent near the villages Enkhory and 
Tsagan-Usun in steppes and forest steppes of the Dzhida Range, and along 
the Mongolian border. It is very rare in other parts of the Dzhida Range, 
never reaching the Mongolian border. We caught it only in the estuary of the 
river Zheltura. It was caught near the middle reaches of the river Khuldat, 
and it is very common along the left bank of river Dzhida from the estuary 
to its left tributary Ulikchin. The most northern point in which we found it 
was near the village Udung (on the left bank of the river Temnik) and in the 
valley of the middle reaches of the river Armak. In the northeastern region 
it is found in the vicinity of Gusinoe Lake, the town Novoselenginsk and 
village Verkhnii Ubukun". 

If O . bedfordiae Thomas is considered as identical with O . daurica, 
the eastern range ofO. daurica extends to the Shansi Province in 
China, but this is still not certain. 

N.M. Przheval'skii (E.A. Buchner, Nauchnye rezul'taty. Issue 4, 1890, 
p. 173) found this species in the southeastern Gobi and Ordos, i.e. , the most 
fertile parts of the Gobi desert. This southern range is apparently not 
connected directly with the northern range mentioned above. 

MODE OF LIFE. Habitats. In the northern part of its range, the 
Daurian pika lives mainly on high steppes (rarely on firm sands) where it 
forms large colonies. According to A.N. Formozov (1929), it preferably 
inhabits ravines and depressions in southern Transbaikalia steppes, rarely 
mountain meadows (e.g., near Uliassutai [Jibhalanta] on the Hobsogol). 
Colonies ofO. daurica occur sporadically along river valleys in the semi- 
steppe and desert. Their favorite habitats are thickets of Iris biglumis 
and Stipa splendens near water. Small plots with fresh grass are 
usually found between the scattered hillocks in which the pika builds its 



45 



60 



btirrows. A.N. Formozov adds that: "This species does not avoid the 
proximity of water, and often settles in damp or very low places. Its 
burrows become, therefore, completely flooded when the water rises. " 

B.S. Vinogradov and S.I. Obolenskii (1927) found colonies of this species 
in the open steppe or in thickets of peashrub (Caragana arborescens) 
in the vicinity of Troitskosavsk [Kyakhta] and village Durena. Only once 
did they come across a pika in a ravine overgrown with shrubs. 

According to M.N. Przheval'skii (E. A. Buchner, 1890) the Daurian 
pika occurs in the most fertile parts of the Gobi desert (northeastern 
Mongolia and Ordos). 

M.N. Przheval'skii states: "North of the Ural, in the part of Mongolia 
where many forests and shrub grow, Ogotona is found in large numbers in 
thickets of shrub along rivers. " 

GENERAL DESCRIPTION AND HABITS. M.N. Przheval'skii (E. A. 
Buchner, 1890) gives the following description: "The behavior of the pika 
is strongly influenced by curiosity. If it has discovered a passing man or 
dog, it lets him approach to about 10 steps and then suddenly takes flight 
into its burrow. However its fear is overcome by curiosity. After a few 
minutes, the head of the animal appears out of the burrow. If the object of 
alarm withdraws, the Ogotona immediately comes out and sits on its hind 
legs at the entrance of the burrow. " 

It is very easy to kill a pika. However, even when fatally wounded, it 
still has time to escape into its burrow. Therefore, only specimens killed 
on the spot become the hunter's booty. 

G. Radde (1862) states that the voice of the Daurian pika is much more 
sonorous than that of the Alpine pika. According to A.N. Formozov, 
O. daurica emits a long, rolling trill gradually subsiding and becoming 
weaker at the end, sounding like: "pit'-pit'-pit'-pif- pit', pi-pi-pi-pi". 
This resembles the warble of some birds, e.g., that of Ant bus 
trivialis, and it can be heard at regular intervals. 

A.N. Formozov writes that: "The pika is regularly active during the 
day, but it also often keeps awake during the night. We caught one at night 
in the hilly sands near Orog Nur while looking for hedgehogs with a 
searchlight. We often heard its whistle even in rainy and cloudy weather 
in complete darkness." 

G.I. Radde (1862) also often heard the voice of these pikas at night. 

BURROWS, NESTS, AND FOODSTORES. Based on observations in 
Mongolia and Gobi, N.'M. Przheval'skii states "The burrows of this pika 
are usually built in colonies. Therefore, it one finds one burrow, one may 
find in the same area, tens, hundreds and sometimes even thousands of 
burrows. " 

A.N. Formozov (1929) found so many burrows in escarpments near 
Troitskosavsk [Kyakhta] that it was almost impossible to see the limits 
between different nests or colonies. The burrows of the Daurian pika on 
even ground had 2 to 5 exits with small mounds crossed by footpaths. 

N. V. Nekipelov (1935) gives a different description of the burrows of the 
Daurian pikas: The burrows consist of many shallow corridors with 
numeroi s exits. The area of the burrow of one rodent is large, it may 
reach 25-35 sq.m. The number of exits is also very large, 15-20. The 
entrances are about 5 cm wide. The burrows of pikas apparently were 
inhabited previously by voles because a) old stores of voles could be found in 



46 



excavated burrows; b) the galleries were narrow (3 cm wide). The pikas 
widened these galleries later and adapted them for their use. 

G.I. Radde (1862) states that burrows of pikas are often built near the 
large burrows of tarbagans (Marmota bobak sibirica). In winter 
long winding burrows dug through the snow with air holes are often seen. 

The Daurian pika prepares small "haystacks" for winter in front of the 
entrance of the burrow. The hay serves both as bedding and as food in 
winter. The efforts of the rodents are often wasted as this hay is eaten by 
cattle. In this case the rodent has to live throughout the winter on dry 
grass which it finds under the snow. 

A.N. Formozov (1929) found stems of the following plants in the "hay 
stacks": Medicago ruthenica, Artemisia dracunculus, 
Potentilla t anace t if o li a, Artemisia siversiana, 
Potentilla s ibir ic a and Me di c ago falcata (the order of the plants 
in this list is according to their decreasing frequency). 
61 A.N. Formozov says; "When the tips of the iris leaves have hardly begun 
yellowing in autumn, the pikas begin cutting them and piling them in large 
heaps near the burrows. The leaves are arranged in a star pattern and are 
moved from time to time. However, this does not prevent decay of some 
leaves deep in the heap. In addition to iris, the haystacks contained some 
grasses and artemisia. The stacks were about 50 cm wide and 35-40 cm 
high. Along the river Dzabhan Gol, pikas, the burrows of which were on low 
land under willow roots, arranged their haystacks near stones, trees and 
on stumps. " 

N.M. Przheval'skii found in the haystacks: grasses, artemisia, 
Conapositae, Leguminosae, and twigs of peashrub. The food piles were 
carefully built like hayricks, always near the entrance of the burrow. 
G.I. Radde (1862) says that pikas cover their haystacks with branches 
of peashrub or Chenopodium to prevent scattering by the wind. 
Ogotona can live without water. N.M. Przheval'skii writes as follows: 
"In winter the animal obtains its water from occasional snow, and in 
summer from rain. If there is no rain, it uses dew, although this is 
rather rare here. 

"It is a questionwhat Ogotona drinks in spring and autumn when drought 
and extreme dryness of the air prevail often for months on the Mongolian 
upland. " 

The animals apparently use the moisture reserve of desert plants. 

REPRODUCTION. A.N. Formozov writes as follows: "This rodent has 
several litters each summ.er. We continuously found young specimens during 
our expedition. A female caught on 2 Aug. (river Tuin Gol) had five young 
ready to be born. A female caught on 2 Sept. had 5 embryos 15 mm long, and 
it was still accompanied by 3 young of the previous litter which were about 
half as large as the mother. " 

J. Allen (1927) reports that he caught very young pikas, only 105 mm 
long, 20 miles south of Ulan-Bator (Urga) on 18-19 May 1922. 

A.M. Kolosov caught a young pika only 75 mm long on 8 Jul. 1936 in the 
Chuya Alps along the river Chegan-Burgazy (2000 m). 

There is a young specimen, 130 mm long, in my collection, which was 
caught on 2 5 Jul. 1934 in the floodland of river Toshtugovu, a tributary of 
the Shibita, 10 km from Toshakty. B. A. Kuznetsov caught a young specimen, 
134 mm long, on 10 Aug. 1928 in the Aginskoe District near Lake B. Tsaga- 
Nor. These data confirm the assumptions of A.N. Formozov mentioned 
above . 

47 



The knowledge about molts in this species is fragmentary. A.M. 
Kolosov caught adults on the Chuya Alps on 5 Jul. 1936 and 9 Aug. 1936. The 
head and the greater part of the back of these two specimens was covered 
with summer fur. Winter fur was present only on the lower part of the 
back, and less on the flanks. A male caught on 10 Jul. 1936 in the same 
place was entirely covered with summer fur. The fur of a pika caught in 
Dalai Nor (17 May 1935) was intermediate though it had already much summer 
hair as a result of its early molt (on the head and part of the back). Pika 
No. 1762 in my collection caught near Troitskosavsk [Kyakhta] on 21 Oct. 
1924 has a complete winter fur. 

POPULATION DYNAMICS. The population of pikas, like that of many 
Lagomorpha, is subject to considerable yearly fluctuations. B.A. Kuznetsov 
(1929) writes that: "The population of Daurian pikas in the steppes of 
„ southern Transbaikalia apparently undergoes considerable yearly changes. 
The Cossacks of the Chindant and Kulusutai villages were well acquainted 
with this animal by the name senokoska ("haystacker") or kuyun. They 
told me that the animal was very numerous in the adjacent steppes in 1927, 
but I only caught 3 specimens in 1928. " 

ENEMIES. The Daurian pika has many enemies. In winter it is hunted 
by snow owls, corsac foxes and wolves. According to N.M. Przheval'skii 
pikas suffer a lot from eagles and buzzards. This author writes: "The 
agility of these birds of prey is amazing. I witnessed buzzards repeatedly 
swooping so rapidly on an ogotona that the animal did not have time to dash 
into its burrow. We once saw the same done by an eagle who swooped on a 
pika sitting near the entrance of its burrow from a height of at least 30- 
40 sajenes*. Buzzards (A r c hi b ut e o hemilasius, i.e., Buteo 
ferox hemilasius Temm. et Schleg. ) feed on pikas to such an 
extent that their hibernation sites in the Gobi are mainly chosen in 
connection with the population of pikas. " 



Subspecies of the Daurian pika 

320. Ochotona (Ochotona) daurica daurica Fall. (1776), 
Eastern Daurian pika 

1776. Lepus dauuricus Pallas P. , Reise durchverschiedene Provinzen des Russischen Reiches, III, S. 692. 
1778. Lepus ogotona Pallas P. S. , Novae species Quadrupedum e Glirium Ordine, pp. 59-70 (cliffs along 

rivers Selenga, Chikoyu and Dzhida). 
1811. Lepus Ogotona Pallas P.S., Zoographia Rosso- Asiatica, p. 151. 

1862. Lagomys ogotona Radde G. , Reisen im SUden von Ost-Sibirien, S. 226-232 (Tarei-Nor). 
1913. Ochotona daurica Kashchenko, N. F. , Novye issledovaniya po mammologii Zabaikal'ya (New 

Studies of Mammals of Transbaikalia), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, XVII, 

p. 397 (Aga Steppe), 
1927. Ochotona daurica Allen Gl. , Lagomorphs collected by the Asiatic Expedition, Amer. Mus. 

Novit. , No. 284, pp. 2-3 (20 miles southwest of Urga). 
1927. Ochotona daurica Vinogradov, B. S. and S. I. Obolenskii, Materialy po faune gryzunov yuzhnoi 

chasti Eniseiskoi i Irkutskoi gubemii i Zabaikal'ya (Material on the Rodent Fauna of the Southern Part 

of the Yeniseisk and Irkutsk Provinces and Transbaikalia), Izvestiya Sibirskoi stantsii zashchity rastenii, 

No. 2, p. 15 (vicinity of Troitskosavsk [Kyakhta] and Durena). 

TYPE LOCAUTY AND DEPOSITION: In the famous work of Pallas "Reise" etc, , BD. Ill, 1776, p. 692, we 
lead: "Vivlt in campis, montiumqae declivibus arenosis apricis, per totam Dauriam. . . ". 



* [Sajene: 2.134 m.] 



48 



63 



In another work of Pallas, "Novae species Glirium", 1778, p. 60, the first locality mentioned in the 
description of the geographical distribution is mountains along river Selenga (Transbaikalia), This may be 
considered as "locus classicus". 

It is unknown whether the holotype has been preserved, 

MATERIAL EKAIvDNED: 22 pelts, most of them with skulls. 

DIAGNOSIS. Color of summer fur on back rather pale, straw-gray 
with a slight ocher tinge which is especially marked on top of head. Ocher 
tones on flanks and cheeks distinct. Belly white, hairs with dark lead-gray 
bases and white tips; chest between forelegs with an indistinct pale straw 
spot; a shaded stripe extends from this spot to the middle of the belly. 
Winter fur on back and on top of head gray with a straw tinge; middle of 
back with blackish brown speckles caused by the black tips of the guard 
hair. Straw tones more distinct on flanks. 

Length of skull 38,5-44 mm (M.42.4); condylobasal length 36.3-41 mm 
(M.39.6); zygomatic width 20.4-21 mm {M.20.8); width of skull at level of 
auditory meatus 19. 2-20. 2mm (M. 30); length of upper tooth row 7.7-8.6 mm 
(M.7.9). 

Body length up to 180mm (M. 175.5); hind feet to 28mm (M. 27). 

ADDITIONAL DESCRIPTION. The fur on top of the head straw -gray in 
summer with a more or less marked ocher tinge varying between pinkish 
buff and cinnamon-buff (Ridgway, 1912, pi. XXIX). Neck paler, with a 
more yellowish tinge (pinkish buff, Ridgway, pi. XXIX). Back pale, straw- 
gray with a slight rusty tinge, and with distinct dark brown speckling 
caused by the dark hair tips. Color of flanks more pure, ocher -straw 
(pinkish buff); same color present on cheeks. Spot on chest pinkish buff. 

Winter fur much more grayish. Back generally straw-gray, with 
distinct blackish brown speckling caused by the dark tips of the long hair. 
Color of flanks similar to that of back but with less marked dark speckling 
and more distinct straw tones. 

GEOGRAPHICAL DISTRIBUTION. Range including northern and central 
Mongolia, southeastern Mongolia and Ordos, southern Transbaikalia 
(Troitskosavsk [Kyakhta] and Verkhneudinsk [Ulan Ude] districts, valley 
of the river Onon, Aga and Daurian steppes, vicinity of Tarei Nor and 
Dalai Nor). 

321. Ochotona (Ochotona) daurica altaina Thos. (1911). 
Western Daurian pika 

1911, Ochotona daurica altaina Thomas O. , New Mammals from Central Asia mostly collected 

by D. Carruthers, Annals and Magaz, of Natur. History, Ser. 8, pp. 761-762; Thomas O. , Annals and 

Magaz. of Natur. History, XIII, 1924, p. 164. 

TYPE LOCALITY AND DEPOSITION: vicinity of Lake Achita Nur (northwestern Mongolia). Type 
No. 205 (collector's number) 27 Aug. 1910 (British Museum). 

MATERIAL EXAMINED: 25 pelts with skulls. 

DIAGNOSIS. Summer fur on back markedly grayer and duller than in 
typical form; ocher-straw tones weaker. Color of top of head like that of 
back. Rust-straw shades on flanks and cheeks very slight, covered by 
grayish tone. Straw spot on chest markedly paler and duller than in the 
typical form. Color of winter fur not much different from that of typical 
form, but perhaps paler. 



49 



Length of skull 44-47 mm (M. 45.9); condylobasal length 40.7-45.2 mm 
(M.43.3); zygomatic width 21.7-23 mm (M.22.2); width of skull at level of 
auditory meatus 21-21.5 mm (M. 21.2); length of upper tooth row 8.3-8.8 mm 
(M.8.6). See Figures 21-25. 





FIGURES 21, 22. Skull of Daurian (western) pika, O chot on a daurica altainaThos. 9 July 1936. 
Collected by A, M. Kolosov from the Chuya Alps, Chegan-Burgazy 

Drawing by Yu. A. Kostylev 



65 Body length to 195 mm (M. 191.7); hind foot to 31 mm (M. 29.7). 

This subspecies differs from the typical form in its grayer and 
duller coloration and its larger size. 

GEOGRAPHICAL DISTRIBUTION. Chuya Alps along the Chegan-Burgazy 
(2000m) by A.M. Kolosov (1936) near Tashakty (in the Gorno-Altai 
Autonomous Region), and near Lake Achita Nur (type locality), Suok (on 
the Altai) near Lake tjreg Nur (to the north of Lake Achita Nur), in Hobdo 
(specimens in the ZMAN collection). 



50 




FIGURE 23. Skull of Daurian (western) pika, Ocho ton a daurica altaina 
Thos. , (lateral) and mandible of same specimen as in Figures 21, 22 





FIGURES 24, 25. Skull of young Daurian (westem)jpika, Ocho tona daurica altaina 
Thos. , Altai, Chuya Alps, A. M. Kolosov Drawing by Yu. A. Kostylev 



51 



152. Ochotona (Ochotona) rufescens Gray (1842), Af^an pika 

1842. Lagomys rufescens GrayJ., Annals Magazine of Natural History, p. 266. 

NAMES: The Afghan pika (English). 

LOCAL NAMES: Gurukses-sichan (Turkmenian). 

PRINCIPAL HGURES: 1) Blanford W. F. , Eastern Persia, H, Zoology, 1876, pp. 83-84, pi. VI, fig. 2 
(entire animal in color); 2) Blanford W. F. , The Fauna of British India, 1888, p. 458, fig. 148; p. 455 (skull) ; 
3) Major Forsyth, On Fossil and Recent Lagomorpha, Transactions of the Linnean Society of London, Vol. VII, 
Zoology, 1896-1900, pi. 38, fig. 14, 29; pi, 39, fig. 34 (parts of the skeleton); 4) Argyropulo A. I. , Materialy 
po faune gryzunov Srednei Azii (Material on the Rodent Fauna of Soviet Central Asia), II. Pishchukhi 
Srednei Azii i Kazakhstana (Pikas of Soviet Central Asia and Kazakhstan), Trudy Zoologicheskogo instituta 
Akademii Nauk, 1, 1932, p. 46, fig. 5 (dorsal, ventral and lateral views of skull); 5) Flerov K. K. and I. M. 
Gromov, Mlekopitayushchie dolin Sumbara i Chandyra (Mammals of the Valleys of the Sumbar and the 
Chandyr), Trudy Karakalinskoi i Kzyl-Atrekskoi parazitologicheskikh ekspeditsii, 1934, p. 331. fig. 9 
(structure of legs). 

MATERIAL EXAMINED: 22 pelts with skulls. 

66 DIAGNOSIS. Skull large in fully grown and old specimens. Skull length 
over46mnri, reaching 53 mm. Interorbital space narrow, 3-3. 8mm in 
adults, convex. Well developed lateral crests present on both sides of 
interorbital space of adult and old specimens; the older the animal, the 
more do the lateral crests converge; the crests run along the lateral parts 
of frontals, forming the inner contours of the orbit. Nasals widened 
anteriorly, narrowing posteriorly after a slight constriction, their 
posterior part rounded with a narrow incision in the middle. Frontals 
without frontal fenestrae. Vomer visible, not covered ventrally by 
premaxillaries. Nasal and incisor foramina fused (Figures 26-29). 

Most vibrissae dark brown; longest vibrissae almost completely white; 
vibrissae relatively short, 45-55 mm long. Soles whitish. Ears without 
distinct light margins. 

Color of summer fur on back mainly pale gray (light-drab and even drab- 
gray, Ridgway, 1912, pl.XLVI), often mixed with a straw tinge (wood- 
brown, Ridgway, 1912, pi. XVI); tips of hair on back rust-brown; general 
color rust-brown (sayal-brown, Ridgway, 1912, pi. XXIX). 

Color of top of head like that of back;, a pale stripe on the occiput. 
Flanks much grayer than back; belly dirty-whitish gray, often with a 
yellowish tinge. 

Winter fur duller, grayer and more brown than summer fur. 

Body length up to 207 mm (M. 205); hind feet to 35.2 mm (M. 34.5); ear 
to 27 mm (M. 22). 

ADDITIONAL DESCRIPTION. The bullae osseae of the Afghan pika are 
compressed laterally and narrowing anteriorly. Rostrum rather long; 
sagittal crest of adult, and especially of old pikas, well developed and 
clearly convex. 

Skull of young pikas more convex in profile; interorbital area more 
strongly convex. The parieto-occipital region and the whole rostrum is 
more flattened, elongate and slanting in adults and old specimens; rostrum 
also more slanting. The width of the brain case in younger specimens 
measured behind the zygomatic arches either greater than that of old 
specimens or nearly as great. Ratio of this measurement to maximal length 
of skull 3.69 in old specimens and 2.99 in young ones. 

Ground color of summer fur on back rather light gray (light-drab or 
drab-gray); this ground color is often mixed with a straw tinge (wood- 
brown); rust-brown shade distinct in middle of back (sayal-brown). Rusty 
tinge well marked on top of head (cinnamon-buff or cinnamon). 

52 



67 





FIGURES 26, 27. Skull of Afghan pika, Oc ho ton a rufescens reginaThos. , dorsal and ventral, 
No. M.4020 d' 23 June 1925. Germab, Kopet-Dagh, S.I. Ognev's collection 

Drawing by Yu. A. Kostylev 




FIGURES 28,29. Skull and mandible of 
Ochotona rufescens regina 
Thos. cC specimen No, M. 4020. (lateral) 




53 



68 Cheeks and stripe behind the ears around the neck light ash-gray. A rust- 
red stripe posterior to this stripe (between cinnamon-buff and cinnamon). 
This stripe widens markedly under the ear, and forms an elongated spot. 
In A.I. Argyropulo's opinion (1932) this spot is characteristic for this 
species. Ears covered with whitish gray hair; posterior parts and inner 
side of ears with rust tones. Flanks duller and grayer than back, sometimes 
with a slight yellow tinge. Belly dirty whitish, sometimes with a distinct 
straw-yellow shade. Feet whitish both dorsally and on the soles; claws 
black. 

The richest colors are seen in old pikas, especially in males. The 
general color of the back is accompanied by a strongly varying reddish 
color of forehead and parietal area. The reddish tinge is relatively 
indistinct in some young females, but its rich tone can be seen from a 
distance in adult specimens. There is a yellow shade on the belly of old 
males which is usually absent in females of the same age. Females 
generally smaller and paler than males. 

Color of young pikas nearly without any reddish tinge. Hair of back with 
well marked brown tips giving a bone -brown color to the whole back 
except the pale collar. Hair much softer than in adult specimens. 

Winter fur duller and grayer than summer fur, intermediate between 
drab and hair-brown (Ridgway, 1922, pi. XLVI) with a slight reddish tinge 

GEOGRAPHICAL DISTRIBUTION. I wrote with V. G. Heptner: "This 
pika is widely distributed along the Kopet-Dagh and the Bolshie Balkhany 
("Mammals of Central Kopet-Dagh" "(Mlekopitayushchie Srednego Kopet- 
Daga), 1925, p. 77) According to our observations the distribution of this 
species is very irregular and sporadic We noticed on our expedition 
that pikas were rare in the mountains near the village Verkhneskobelevskii. 
We did not find any on the road along the border from Verkhneskobelevskii to 
Mikhailovskii (Germab) Near Mikhailovskii (Germab) it is already common, 
occasionally occurring in large numbers. We found the following colonies in this 
region: l) in the estuary of river Sakis-Yab (Germab); 2) a large colony in a 
ravine to the west of the village, near the so-called "Cossack camps". We also 
found it alongthe precipitous walls of the broad valley from the Germab frontier 
post to the Kheirabad frontier post (which is now ruined). 

"Near the village Dmitrievskii (Kurkulab) Ochotona is numerous along 
a ravine of the river Mergen-Uli, about 8 versts above the village and most 
common near the village itself. We did not find this species near the 
village Sulyukli. " 

A.I. Argyropulo (1932) recorded the following localities for O. 
rufescens from material in the ZMAN collection: vicinity of Kizil- 
Arvat; the Bol'shie Balkhany Range and Kazandzhik Mountains (near the 
junction of the Malye Balkhany Mountains with the Kyuren-Dagh Range). 
P. Varentsov (1894) states that it occurs on the Kopet-Dagh near Gaudan 
and in the Firyuza Ravine. 

69 K.K. Flerov and I.M. Gromov (1934) recorded the following localities for 
this species. The largest colony was found on the Chandyr mountain pass in 
a narrow ravine at the bottom of which runs the Karakala-Chakankala Road. 
A second, smaller colony, was found 15 km from Kara-Kala, where the 
Sumbar river flows between rocky ranges on both sides The third, 
smallest colony was found on the slopes of the Skalistoe Ravine between the 
Naarli outpost and Kizil-Imam (in the Chokhrok plot). 



54 




FIGURE 30, Ochotona rufescens reginaThos., calling 

Drawing by K. K. Flerov 

This species was found by A.M. Nikol'skii in northeastern Iran (near 
Aber, Nardin and Bojnurd). O. rufescens is undoubtedly widely 
distributed throughout Iran. It was recorded by W.T, Blanford (1876) from 
Korud north of Isfahan, According to Blanford (1888) it is found in 
Afghanistan near Quetta, and in northeast India; according to Thomas 
(1920) it is found near Kalat in Central Baluchistan. 

O. rufescens is widely distributed, but occurs in Russia only at 
the extreme northeastern tip of its range. 

MODE OF LIFE. HABITATS. The irregular distribution of the species 
is connected with the specific habitats of these animals. They inhabit almost 
exclusively slopes of ravines, and avoid flat country and valleys. The 
70 favorite habitats of these animals are gradual slopes, but they sometimes 
occur on steep escarpments. They occasionally occur near cultivated land 
if they find stones and rocks. I found numerous pikas near the village 
Kurkulab in adobe fences of settlements and along roads. These adobe walls 
("duvals") were completely perforated by burrows, like sponges. 

P. Varentsov ("Fauna of the Transcaspian Region" (Fauna Zakaspiiskoi 
oblasti) Ashkhabad, 1894, p. 14) found a colony of pikas in a plain. He 
writes as follows: "I found a large colony of pikas in the Firyuza Ravine 
in the Kopet-Dagh 5 versts from the road from the Ak-Tepe military camp 
to the village Kozelnoe. The burrows of the pikas were under stones, not 
on mountains as in Guadan, but in the foothills. Burrows were occasionally 
dug in soft ground of a large valley covered with lush vegetation. The 



55 



Firyuzinskii Brook (Izgand-Su) runs along this valley. The valley is about 
6000 ft above sea level, and it is surrounded by high stony mountains without 
any vegetation. " 

GENERAL DESCRIPTION. HABITS. I wrote with V. G. Heptner (1925): 
"This animal is usually very trusting and graceful. Its appearance, behavior, 
and facial expression remind one of a small rabbit, quite different from the 
drawings of this species given by Miitzel et al. Its movements are rather 
phlegmatic. Quietly coming out of its burrow, it sits for a long time in 
front of its entrance with its four legs folded. 




71 



FIGURE31. Afghan pika Ochotona rufescens reginaThos., on an adobe fence in the 
Kopet-Dagh 

Photograph by S. I. Ognev 

"It moves quickly when disturbed and settles again observing. Sometimes 
it behaves quite differently, demonstrating how rapidly it can move. It 
runs in curves. Its movements are between the running of a mouse -like 
rodent and the leaps of a very short-legged rabbit. Both feet of the animal 
touch the ground when it runs, as in hares. After having run some distance, 
it invariably stops near its burrow, folds all four legs and observes 
calmly. This is a very characteristic posture in which it is very similar to 
a hare. A hasty movement of the observer suffices to make it jump to its 
burrow in one bound, so rapidly that it cannot be followed by the human eye. 
However, the animal is curious and will come out of its burrow again after 
a short time, " 

Strangely enough, we heard its voice only once. It was a very weak, 
melodious squeak. P. Varentsov (1894) describes the voice of this species 
as monotonous. If alarmed it gives a peculiar prolonged cry. 



56 




FIGURE 32. Afghan pikaOchotona rufescens reginaThos. , in the Kopet-Dagh Mountains 

Photograph by S. I. Ognev 



72 



73 



K.K. Flerov and I.M. Gromov (1934) write as follows on the voice of 
Ochotona rufescens: "The voice .. . is very weak, and can be 
heard only for a very small distance. It has nothing in common with the 
true bird songs of the Daurian pika or with the loud cry ofO. hyperborea 
It is a short, almost soundless warble starting with a very short, hardly 
audible whistle followed by a series of dry, jerky sounds resembling the 
suppressed rattling of an alarm clock. Its voice sounds about like "fyu- 
trrrrrrrr". One specimen ended the warble very rapidly, drumming the 
ground with its hind feet. " 

DAILY LIFE. According to our observations in the Kopet-Dagh (June 
1925) the pikas are diurnal. There was a prolonged drought and hot 
weather in 1925. Pikas appeared early in the morning, soon after sunrise. 
They fed and were active to 8-8:30 a.m. when the heat became intense. 
They could also be seen after 5-6 p.m. when the heat subsided. However, 
they were more sluggish than in the morning. In spring and fall, when the 
weather is cooler, the animals are generally much more active than in 
summer. 

BURROWS AND NESTS, I wrote with V. G. Heptner (1925): "On stony 
slopes, the entrance of the burrow is always under a large stone, or it 
leads into a deep crevice. Burrows are generally built near a group of 
several stones and the animal often sits on the stone overhanging the nest. 
There is always a small, flat, beaten and completely clean landing in front 
of the burrow (about 1 sq. ft. ) which is usually roofed by an overhanging 
stone. This is the balcony of the animal where it sometimes sits for long 
periods in the shade of the overhanging block. It apparently does not 
move far away from its burrow. We never saw a pika more than 50-100 
steps from its burrow. There is always a small heap of excrement near 



57 




FIGURE 33. Burrows of Afghan pika in an adobe fence in the Kopet-Dagh Mountains 

Photograph by S. I. Ognev 




FIGURE 34. Near the entrance of the burrow of the Afghan pika. The burrow is in a ravine among stones. 
Excrements are visible near the entrance. Germab Ravine, Kopet-Dagh 

Photograph by S. I. Ognev 



58 



74 



the burrow. Pellets accumulate in rather large quantities, sometimes 
large handfuls. They look like pellets of hares, but are much smaller 
(3-4 mm). Only rarely did we find fresh feces near inhabited burrows. 
The excrements were usually completely dry. However, they dry very 
rapidly here. " 

". . .The pika burrows with preference in adobe walls near roads. The 
burrows in this habitat are of a peculiar type which is worth describing. 
The fence is about 1 meter high. The exits of the burrow open at the 
base of the fence at both sides. Their diameter, as that of the galleries, 
is about 7 cm. The exits are sometimes widened because the soil crumbles. 
There is rarely only a single exit. There are usually 3-4, which apparently 
belong to the same network. Burrows built among the rocks have several 
exits, but single burrows also often occur there. This is rare in fences. 
The softness of the ground apparently enables the animal to build numerous 
galleries with exits. There are often simple transverse tunnels under the 
wall. The main gallery usually leads from the base of the fence into the 
ground. In broken walls galleries often run along the wall for a long 
distance. Exits are sometimes l/2 meter above the ground. The animal 
apparently loses its direction and makes an accidental opening. This 
crumbles and eventually becomes very large, but it is apparently not used 
as an entrance. 

"The pika is very agile and jumps very well. It can jump onto a wall 
about 1 meter high without any apparent effort. 




FIGURE 35. Dry hay reserves near the entrance of the burrows of the Afghan pika, Germab Ravine in Kopet-Dagh 

Photograph by S. I. Ognev 



59 



We found a burrow near the Dmitrievskii village in a crack in a monolithic 
rock. The animal had, each time, to jump onto a small platform (about one- 
tenth sq. m) before the entrance into the crack 1/2 m above the ground. 

"A considerable quantity of soil accumulates near burrows in walls along 
roads. Small balconies are formed by this soil in front of the entrances, 
especially in places where a gutter runs at the base of the wall. Along the 
base of these walls there are cornices (about 7 cm wide) formed by the 
rejected soil. These cornices transform the sloping wall of the gutter into 
a horizontal surface along which the animals run along the wall. Such 
tracks occasionally run for a long distance, covered here and there by 
weeds or shrubs. 

"An unusual type of burrow was found once near the Dmitrievskii village 
(Kurkulaba). Several exits were located under a C appar i s shrub on a 
small mound of earth. The form of the exits seemed to be that of a desert 
hare but small pellets were found near the burrow. " 

FOOD AND FOOD STORES. O. rufescens is modest in its food 
requirements. In food stores near burrows we found dry Artemisia 
and Ephedra. It cuts the long thorny stalks of ephedra and dries them 
by pushing bunches into cracks between stones near the burrows. The 
store is sometimes under the stones overhanging the entrance. A large 
part of the stores is apparently taken deeper into the burrows. Food 
stores are undoubtedly collected twice a year: a) in spring, when the rich 
vegetation comes to an end because of the onset of heat; b) in autumn, 
when the vegetation is revived by rain. The autumn reserves are prepared 
for the winter. Pikas living in "duvals" and near plantations eat alfalfa 
avidly. We also found apricots whose flesh had been eaten but the kernels 
had been left intact. 

We have no data about the reproduction of the Afghan pika. We did not 
find any young specimen of the same year during our expedition to Kopet- 
Dagh (1925). According to local hunters, the females suckle their young in 
June in the burrows and never, or only very rarely, come outside. They 
feed then on stores collected in the spring. The males live in separate 
burrows at this time and have no responsibility towards their family. They 
frequently come outside. This is probably the reason why there are so many 
more males than females in our collection, 

MOLT. In spite of the hot climate of Turkmenia, the spring molt takes 
place rather late. We caught a female near Germab in June 1925 (M.4011) 
with winter fur persisting on the lower part of the back and on the flanks. 
Most of the back of a male (M, 4017) caught on 21 Jun. 1925 (Kurkulab Ravine) 
was covered with strongly worn winter fur. The molt proceeds from the 
head to the hindquarters, as it is typical in the spring molt of Ochotona. 
Finally, the winter fur in a male (M, 4016) caught on 30 June 1925 (Germa.b 
Ravine) persisted in a few traces on the lower part of back and on the flanks. 
Based on the examiination of material from the Sumbar and Chandyr Valleys, 
K. K. Flerov and I.M. Gromov (1934) came to the following conclusions 
about molt of the Afghan pika. The summer fur grows during June and in 
early July. In the second half of July and first half of August, the animals 
bear the summer fur. The winter fur begins growing from the second half 
of August, and even earlier. This ends in the second half of September. 
The young are born in winter fur. The summer fur begins growing in the 
second half of July. Immediately after the summer fur has grown, the 
molt into winter fur starts again. The higher the pikas live, the longer they 



60 



76 



probably wear their winter fur, and the later begins the spring molt. 

LIFE IN WINTER, The Afghan pika does not hibernate like its relatives. 
With the onset of winter, the animals block their burrows. They come out 
only to take hay from their "haystacks" or to catch a breath of fresh air 
on a nice sunny day. On the 20th of December 1892, P. Varentsov saw 
many pikas on the Gaudan jumping over the rocks. Eleven were shot and 
were found to wear a complete winter fur. 

LIFE IN CAPTIVITY. According to P. Varentsov (1894), pikas are very 
delicate and have to be treated carefully in captivity. It is difficult to keep 
them in a room because they are very unclean. Their copious urine has a 
strong, very disagreeable smell. 

322. Ochotona (Ochotona) rufescens regina Thos. (1911). 

Kopet-Dagh pika 

1885. Lagomys rufescens Nikol'skii, A.M., Materialy k poznaniyu fauny pozvonochnykh severo- 

vostochnoi Persii i Zakaspiiskoi oblasti (Material on the Vertebrate Fauna of Northeastern Iran and the 

Transcaspian Region), Trudy Sankt-Peterburgskogo obshchestva estestvoispytatelei, p. 7, 
1890. Lagomys rufescens Radde G, , Walter A, , Blasius W. , DieSaugethiereTranscasp. , Nauchnye 

rezul'taty, Tiflis, p. 7. 
1894. Lagomys rufescens. Varentsov, P., Fauna Zakaspiiskoi oblasti (The Fauna of the Transcaspian 

Region), pp. 11-14, 
1911, Ochotona rufescens re gina Thomas O, , On New Mammals from Central and Western Asia. 

Annals and Magaz. of Natur. History, p. 762. 
1929, Ochotona rufescens re gina Ognev S, I. and Geptner, V, G, , Mlekopitayushchie srednego 

Kopet-Daga; prilezhashchei ravniny (Mammals of the Central Kopet-Dagh and the Adjoining Plain), 

Trudy Nauchno-issledovatel'skogo instituta zoologii, 3(1): 75-81, 
1932, Ochotona rufescens Argyropulo A, I. , Materialy po faime gryzunov Srednei Azii, Pishchukhi 

Srednei Azii i Kazakhstana (Material on the Rodent Faima of Soviet Central Asia. Pikas of Soviet Central 

Asia and Kazakhstan), Trudy Zoologicheskogo instituta Akademii Nauk, 1:45-48. 

TYPE LOCALITY AND DEPOSITION: In the Kopet-Dagh Mountains, to the west of Ashkhabad. Type in 
the British Museum, 

MATERLA.L EXAMINED: 22 pelts with skulls, 

DIAGNOSIS, Resembling the species in general (vide supra), O. 
rufescens Gray differs from the nominal Afghan race by its larger 
size*. Length of skull 44-53.2 mm (M. 50,3); condylobasal length 44,5- 
48,7mm (M,46,4); zygomatic width 23-25mm (M,23.8); interorbital 
space 3-3,8 mm (M,3,4); length of upper row of molars 9, 4-10. 2mm 
(M.9,5); Length of body and head 150-219 mm (M. 205); hind feet 31-36 mm 
(M,34,5); ear 19-25, 2mm (M. 22). 

SYSTEMATIC NOTES. I consider the Turkmenian race of reddish pikas 
as belonging to Ochotona rufescens regina. This subspecies was 
described by O, Thomas from collections by Littledale from a locality west 
of Ashkhabad (Kopet-Dagh Mountains), The typical O , rufescens 
rufescens Gray was described from the east Afghanistan Mountains 
(Kabul Region). According to Thomas, O, rufescens regina differs 
from the typical form in its larger size and large skull with typical bullae 
osseae, O. Thomas states that the bullae of the Turkmenian pika are 
larger than those of the typical race, and those of the central Iran O. 
rufescens vizier Thos, The holotype of O , rufescens vizier 
Thos, is the specimen described and drawn by Blanford, ("Eastern Persia 



* According to data of O. Thomas, I have not seen typical rufescens specimens and cannot give an 
opinion. 



61 



77 



78 



II, 1876, pi. VI, Figure 2). This specimen was caught in Korud, north of 
Isfahan*. According to Thomas, O. r. vizier differs from the typical 
form in its small size and richer and more saturated color. Its ears are 
shorter and its bullae osseae are smaller than those of the typical race. 
O. Thomas described O. r. vulturna Thos . , from the vicinity of Kalat 
(Baluchistan) which is smaller than the typical form but larger than 
vizier . The color of its winter fur is more monotonous yellowish red 
than that of the closely related subspecies. 

I can discuss these systematic questions, unfortunately, only from the 
literature. I have only material of the Turkmenian pika and no pelts from 
Iran, Afghanistan, and Baluchistan. 

A.I. Argyropulo (1932) considers the Tibetan O . curzoniae Hodgson 
as definitely identical with rufescens. 

This seems doubtful. W.T. Blanford (Fauna of British India, 1888, 
p. 457) definitely states that O . curzoniae Hodgson and O. roylei 
Ogilby are closely related. According to O. Thomas (On Some New Forms 
ofOchotona, Annals and Magaz. ofNatur. History, ser.9. Vol. IX, pp. 189- 
190, 1922), curzoniae differs sharply from r u fe s c e n s in a number of 
characters. These questions can be solved only by examination of more 
material. 

I have discussed the geographical distribution ofO. r. regina Thos. 
in detail above. 

153. Ochotona (Ochotona) pricei Thomas (1911). Price's pika 

1911. Ochotona (Ogototna) pricei Thomas O, , Mammals from Central and Western Asia, Annals 

and Magazine of Natural History, ser. 8, No. 48, Vol. VIII, p. 760. 

PRINCIPAL FIGURES: 1) Formozov A. N. , V Mongolii (In Mongolia), Moskva, 1928, pp. 75 and 98 
(drawings of entire animal); 2) Formozov A, N. , Mlekopitayushchie sevemoi Mongolii po sboram ekspeditsii 
1925 goda (The Mammals of North Mongolia in Collections of the 1925 Expedition), 1929, p. 91, fig. 15 
(skull doisal and ventral); p. 93, fig. 16; p. 94, figs. 17- 18 (characteristic postures of pikas, sketches from 
nature); 3) Argyropulo A. I. , Materialy po faune gryzunov Srednei Azii (Material on the Rodent Fauna of 
Soviet Central Asia), II. Pishchukhi Srednei Azii (2. pikas of Soviet Central Asia), Trudy Zoologicheskogo 
instituta Akademii Nauk, Vol, 1:49, 1932, fig. 6 (skull dorsal, ventral and lateral). 

MATERIAL EXAMINED: 36 pelts (most of them with skulls), 

DIAGNOSIS. Skull in general similar to that ofO.daurica, of average 
size. Length 44-49 mm. Interorbital space relatively narrow (3.5-4.7 mm) 
and arched in the middle. There are median crests branching anteriorly 
and posteriorly in old specimens which may pass to the parietals. Nasals 
gradually widening anteriorly. Lateral contours of nasals gradually 
narrowing posteriorly, their posterior part rounded. Frontals without 
fenestrae. Vomer covered ventrally by the margins of the premaxillaries. 
Palatine and incisor foraniina separate (Figures 36-38). Vibrissae dark 
brown; longest vibrissae with long, white tips, 60 mm long. Soles whitish, 
sometimes with straw-grayish tinge. Light margins of ears very indistinct. 

Color of summer fur on back varying in different subspecies between 
ocher-rust-gray (cinnamon-buff, Ridgway, 1912, pi. XXIX) with some 
grayer and rust tones (cinnamon and clay color, pi. XXIX) and duller, 
grayish brown tones with a slight yellow tinge; the combination of these 
colors creates a general impression of drab. 



* K. K, Flerov and I. M. Gromov (1934) consider the Asterabad pikas as belonging to this form. 



62 





79 



FIGURE 36. Skull of Price's pika, Ochotona price! Thos. , No. M.3974 lOSeptember 1926. 
Central Mongolia, Tsagan-Adzir Mountain, 100km south of Uliassutai [Jibhalanta], A.N. 
Formozov, S.I. Ognev's collection 

Drawing by Yu, A. Kostylev 

Winter fur on back beautiful straw- gray with a pinkish rust tinge 
(avellaneous). 

Body length to 200 mm (usually about 180 mm); hind feet to 33 mm 
(usually about 30mm). 

GEOGRAPHICAL DISTRIBUTION. This species was described by O. 
Thomas (l91l) from 3 subadult specimens collected by D. Carruthers' 
expedition near Lake Achita-Nur and near Suok, in the upper reaches of the 
river Hobdo Gol basin (northwestern Mongolia), at altitudes of 6700 
and 8000 ft. 

According to data from A.N. Formozov's expedition (1929) it was found 
in north and central Mongolia between Urga and Lamangegen. Numerous 
remains of this species were found in the disgorged pellets of the eagle 
owl on slopes near the river Shargain Gol, southwest of the intersection of 
46°N and 102°E. Farther to the west, this species was found near 
the Lamangegen Monastery. Near Uliassutai [Jibhalanta] it reaches the 
Tsagan-Adzirga Mountain in the north. In the south it occurs on the Gobi 
Altai, where specimens were caught in Yihe-Bogdo. 

Glover Allen (1927) recorded the species from the following localities: 
Gun Burpe; Ussuk; Artsa-Bogdo; and 40 km from Tsetsen Han [Ondor Han].* 
A.I. Argyropulo mentions that Price's pika is found near the village Choirin 
on the Kalgan (Changkiakow) highway. A.M. Belyaev (1933) states in an 



' In Allen's work, Tze-Tzen Wang. 



63 



article on the rodents of Kazakhstan that this species occurs in the southern 
part of the Kazakhstan Plateau, near the valley of the river Syry-Su and the 
Ulu-Tau Mountains to the river Tokrau. Its range reaches approximately 
48° latitude in the north. 





FIGURES 37, 38. Skull and mandible of Ochoton a priceiThos. (lateral) 
No. M.3974 



!0 



There are two specimens of this species in the ZMAN collection caught 
in the former Kedei subdistrict of the Karkaralinsk county of the 
Semipalatinsk Region. V.A. Selevin (in litt. , 1935)kindly put the following 
data on the distribution of this species at my disposal. Price's pika is 
replaced in the desert byO. pusilla south of Karaganda, northwest of 
Balkhash in the direction of Berty-Su. V. A, Selevin recorded O . p r i c e i 
near the Karaurun Hills. It is also numerous in the valley of the river 
Emek. A colony was found in eastern Bet-Pak-Dala near Kara-Ungur 
along the Chakchagailydzhel (in the northeastern part of the desert), and a 
skull was found in disgorged pellets of the eagle owl. It was also found 
on the Altai in the Chuya Steppe (ZMAN collection, Romanov, 1841) and near 
the Talduchinskii Glacier in southeastern Altai, west of the Chuya Steppe). 
O. Thomas (A New Pika from the Altai, Ann. and Magaz. of Natur. History, 
XIII, 1924, pp. 163-164), inhis description of Oc hot ona sushkini 
(=pricei) claims that it occurs in the northeastern Altai. This is incorrect 
because the specimen of P.P. Sushkin was from the southeastern Altai. 

MODE OF LIFE. The only data on the biology of this pika are given in 
A.N. Formozov's work (1929): 

"The distribution of O . pricei is sporadic. It occurs in isolated 
colonies, and in localities which are often far from each other although 
suitable conditions exist in the area between them. This isolation may be 
the reason that traces of extinct colonies are sometimes found in the desert. 



64 



These colonies can come back to life only rarely, because the next colonies 
are too far away. Such traces of extinct colonies are mounds of fine gravel 




FIGURE 39. Price's pika, Ochotona pricei Thos. 



Drawing by A. N, Formozov 



excavated by pikas at the base of rocks covering the burrows. We will 
discuss these structures below, 

"As to the favorite habitats of O . pricei, we can say the following: 
it was most numerous in rocks with deep passages, and in the residual 
outcrop scattered over the desert. These passages produced by water, 
sun and wind, and cracks between structures are most preferred by them. 
They can often be observed lying on flat rocks, basking in the sun. They 
sometimes rest in their niches seeking protection from the wind. They dig 
burrows under such rocks or near them. However, we rarely also found 
burrows in rockless deserts, under peashrub. To our surprise, we 
found some colonies in the valley of the river Dzabhan which had built their 
burrows (each with 5-8 exits) under sandhills overgrown with Stipa 
splendens or on areas covered with stones with isolated peashrubs . 
We also found Daurian pikas near this place. On the Yihe-Bogdo, O. 
pricei is found mainly on higher ground in pastures where C ap r a 
s i b i r i c a graze, and near marmot colonies. The pikas take refuge in thair 
burrows in case of need. 

"The range of this rodent thus reaches from the valleys to altitudes of 

about 8000 ft. 

"Burrows of Price's pika are usually under rocks or bushes. The width 
and shape of the entrances vary considerably. The entrance may be 
either horizontally or vertically elongate, its width 50- 120 mm. There are 



65 



nearly always deep paths through the mound of earth excavated from the 
burrow. O. pricei collects large heaps of rubble (to Im long) consisting 
of small stones (5X5 cm), dry camel and horse dung, and pieces of peashrub 
twigs, near the large flat stones under which the burrows are, and at the 
base of small rocks. These heaps sometimes surround the base of the rock 
and occasionally form small hillocks. The entrances of the burrows were 
blocked with such rubble in some cases. The pikas possibly collect this 
material to block cracks against water and wind. 

"Heaps of grass spread to dry may be found on densely populated rocks 
during the day. Large stores of hay fill large hollows in the rocks, or 
cavities under stones. The hay consisted of the following plants: l) store 
in a niche among rocks on 30 Jul. 1926 in the lower reaches of river Tuin- 
Gol ($ 84) — Ar te m i s i a pectinata, Euratia coratoides, 
Carex ste.nophylla, Tanacetum achillaeoides; 2) store under 
stones on 1 Sept. 1946 between rivers Baydarag Gol and Dzabhan Gol 
Tanacetum achillaeoides, Carex stenophylla, Artemisia 
s p . ; in a third case, grasses and Aster altaicus were found in addition 
to the above plants. 

"When foraging, the pika is very cautious, hurrying to the rocks at the 
first alarm. However, after it reaches its refuge, it does not hide long in 
its burrow, but appears again outside after a short time. The traces in the 
sand indicate that O . pricei does not go farther than a few tens of 
meters from its burrows. It is fond of rocks, and moves rapidly over 
steep surfaces, and has a peculiar way of folding its rather large and broad 
ears. These habits make it easy to distinguish it from the Daurian pika. 
We never heard O . pricei give the prolonged whistle characteristic of 
O. daurica. When it is frightened, it gives a short and sonorous whistle. 
O . pricei apparently has several litters during summer. No. 93='- 5 (l Aug. ) 
was suckling young (milk in the mammary glands) ; 191* $ (iSept. ) had a 
completely developed embryo. We could observe well developed young at 
the same time, which were certainly born in the same summer. " 

G. Allen (1927) reports that some small young pikas were caught on 
31 May and 1 Jun. 40 miles south of Tsetsen Han [Ondor Han]. Other specimens 
of the same size were caught on 21 Jun. near Gun Burte; 25 Jun. near 
Ussuk and 11-24 July near the Artsa-Bogdo Range. 



82 Subspecies of Oc hot ona pricei Thos. 

This species has a well defined subspecies in the Karkaralinsk Region, 
which was described under a separate name by A.I. Argyropulo in his 
monograph in the "Fauna of the U.S.S.R. " (Fauna SSSR) (in litt. ) O. 
sushkini Thos. (vide infra) is certainly identical with O . pricei. 
O . ham i c a Thos. =■=- which was described from Hami [Quomul] south of the 
eastern spurs of the Tien Shan) is also closely related toO. pricei. 



Numbers of the collection of A. N. Formosov. 

O. Thomas, Annals and Magazine of Natural History, ser. 8, 1912, p. 407. 



66 



323. Ochotona (Ochotona) pricei pricei Thomas (1911). 
Mongolian pika or Price's pika 



1867. Ogotoma Pallasii Gray I. E. , Notes on the Skull of Hares (Leporidae) and Pikas (Lagomyidae) 
in the British Museum, Ann, and Mag. of Natural History, Vol, XX, p. 220. 
TYPE LOCALITY AND DEPOSITION: Mountains to the west of Lake Achita-Nur (near 90° E and 

49°30'N), at an altitude of 6700 ft, in the Hobdo Gol Basin, northwestern Mongolia. Type in the 

British Museum, London. 

MATERIAL EXAMINED: 34 pelts with skulls. 



DIAGNOSIS. Color of summer fur on back ocher-rust-gray (generally, 
cinnamon-buff, Ridgway, 1912, pi. XXIX) with some grayer and rustier 
tones (cinnamon and clay color). Winter fur on back beautiful straw-gray 
with a pinkish rust tinge (avellaneous). 

Length of skull 45-49 mm (M.46.8); condylobasal length 41-43.5 mm 
(M.43)i zygomatic width 21-24 mm (M. 23.1); interorbital distance 3.2- 
4.7mm (M.4.2); width of skull at level of auditory meatus 22-24 mm 
(M. 23.2); length of upper tooth row 8.5-9.5 mm (M.9). 

Body length 180-215 mm; hind feet 29-33 mm; ear21-26mm. 

ADDITIONAL DESCRIPTION. Color of summer fur on back ocher- 
rust-gray of varying intensity (generally cinnamon -buff) with some 
grayer and rustier tones (cinnamon, clay color). Fur on top of head of 
the same color as on back; rust-brown-ocher tones more marked on top 
of head and back. Color of flanks lighter than that of back, more 
yellowish ocher. Color of cheeks the same. Eyes surrounded with ash- 
gray fur. Belly whitish, but the lead- gray bases of hair show through the 
white tips. A pale yellowish transverse stripe on the chest between the 
anterior legs. An indistinct pale yellowish rust stripe (pale pinkish-buff) 
runs from the transverse stripe to the middle of the belly. 

Inner side of ear with whitish gray hair; outer side also with such hair, 
but more sparsely. Light margins of ears little marked and undistinct. 

Legs with whitish hair with a dirtier tinge on the soles. Claws horny 
brown. 

The vibrissae consist of a mixture of whitish and brownish black hair; 
tips of the longest vibrissae white. 

General color of winter fur on back straw- gray with a pinkish tinge 
(avellaneous). Top of head grayer, with an ash tone. Flanks with a weak 
light yellowish tinge (pinkish-buff). Belly white, sometimes slightly yellow. 
Color of back uniform without the brown tinge of the gray ground color of 
the winter fur of other forms. A brown tone present only along the midline 
of the back, where there are many hairs with gray-brown tips. 

SYSTEMATIC NOTES. I.E. Gray (1867) described a species with the 
name Ogotoma pallas ii . He wrote as follows "Ogot om a . Skull: 
the orbits very large; space between the orbits narrow; nose narrow, 
bent dov/n". 

"Ogotoma pallasii (Lagomys ogotoma Cuvier, Waterh. , 
Glir. 17. Lepus ogotoma Pallas, Glires, 30, t. 3, 4a, f. 16 a.) 
B.M. ". 

Gray did not indicate the terra typica for his Ogotoma pallasii in 
this fragmentary description. After examining the holotype of O . 
pallasii at the British Museum, B.S. Vinogradov wrote that this species 



67 



84 



is very similar to O. pricei but larger and differing in its narrower 
interorbital space (3 mm). The interorbital space becomes narrower with 
age. It is therefore possible that an old specimen of pricei was used 
as holotype for "O. pallasii". 

As no terra typica was given for "O gotoma pallasii", as there is 
no description of this species, and as Gray partly confused his new species 
with O. daurica Pall. (=Ogot ona Pall.), we think that "Ogot oma 
pallasii" is best considered as a doubtful synonym ofO. pricei. 

The Mongolian pika is closely related toO. daurica, O. 
curzoniae, O. ladacensis and O . rufescens, and forms with 
them a systematic group. It differs slightly from O. daurica in 
characters of its bony palate. 

In Glover Allen's opinion (1927) O . pricei differs from O . daurica 
in the better development of the bare, black, calluses of the toes, and in 
its shorter claws in the hind feet. These characters are not very marked 
and not always constant, but they are certainly correct. 

O. Thomas (1924) (A New Pika from Altai, Ann. and Mag. of Nat. 
Hist. ser. 9, Vol, XIII, pp. 163-164) described anew species from Taldura 
in the southeastern* Altai, west of the Chuya Steppe. He considers O. 
sushkini as closely related toO. pricei, but differing in its richer 
color of the summer fur. Color of fur of O . sushkini according to 
Thomas saturated, rust-ocher-yellowish (cinnamon -buff); this color 
becomes darker on the back because of the dark hair tips; fur on neck and 
lower part of back light and rich. Belly with a strong admixture of rust- 
yellow tones. Top of head grayer than back. 

Length of body and head 200 mm; hind feet (measured on pelt) 28 mm; 
ear 18 mm. 

Skull length 48.3mm; condylo-incisive length 44mm; zygomatic width 
23mm; nasals 15.3X6; interorbital space 4.8mm; width of skull in region 
of bullae osseae 22.4 mm; length of bullae osseae 11.7 mm; length of 
upper row of molars 9.2 mm. 

O. sushkini Thos. undoubtedly is identical with O. pricei, 
according to all characters. 

We are still unable to decide whether Price's pika from the Chuya Steppe 
is a separate subspecies, for lackof material. Winter specimens from river 
Chuya, collected by Romanov in January- February 1841 (in the collection 
of Professor Eversman) do not differ from Mongolian winter specimens of 
O.p. pricei Thos. More material will, possibly, show that the Chuya 
summer specimens ofO. pricei have a richer color than the Mongolian 
ones. In this case, the subspecific name of O . p. sushkini will be 
maintained. 

GEOGRAPHICAL DISTRIBUTION. Given in detail in the general survey. 
This form occurs throughout the wide eastern range ofO. pricei except 
in the western parts of this range, i.e. , the Karkaralinsk Region and 
Sary-Su (vide supra). 



O. Thomas (1924) erroneously recorded this species from the northeastern Altai. Pikas of the Pricei 
type are completely absent in this part of the Altai. 



68 



324. Ochotona (Ochotona) pricei subsp. nov. Price's 
Semipalatinsk pika 

1932. Ochotona pallasii Kuznetsov, B. A. , Gryzuny Semipalatinskogo okruga Kazakhstana (Rodents of 

the Semipalatinsk District, Kazakhstan), Byulleten' Moskovskogo obshchestva ispytatelei prirody, XLI (1-2): 
1 16 [near river Tokradzh (Tokrau), former Kedei subdistrict, former Karkaralinsk County]. 

1932. Ochotona pricei Thomas subsp. nov. Argyropulo A. I. , Materialy po faune gryzunov Srednei 
Azii (Material on the Rodent Fauna of Soviet Central Asia), Trudy Zoologicheskogo instituta Akademii 
Nauk, I, pp. 51-52. 
TYPE LOCALITY AND DEPOSITION. No. 19987 (250, 1929), collection of adults, 21 Jul. 1928, Kedei 

subdistrict, former Karkaralinsk County, former Semipalatinsk Region. Type in the ZMAN collection. 
MATERIAL EXAMINED: 2 pelts and skull fragments, 

DIAGNOSIS. General tone of summer fur on back dull grayish-brown 
with a slight yellowish tinge (general impression: drab). Black with slight 
speckling caused by blackish brown hair tips. Color of top of head like that 
of back; top of head in Mongolian specimens usually more reddish than 
back. Flanks and lower part of back with a more or less marked ocher 
tinge, ground color of flanks gray (smoke -gray). Hair on belly dirty 
whitish with a slight ocher-yellow tinge; bases of hair on belly gray. Winter 
fur''= generally similar to summer fur, but more intensely ocher-brownish 
(buff-brown); dark brown speckles more marked. 
gg SYSTEMATIC NOTES. This subspecies differs very sharply from 
Mongolian specimens of O . pricei. A.I. Argyropulo (1932) mentions 
some cranial differences between Semipalatinsk and Mongolian specimens. 
The skull of the Semipalatinsk specimens is smaller^ the rostrum more 
massive and shorter, tympanic bones larger. These differences are based 
on only one defective skull. It is therefore more than doubtful whether 
they are constant. 

I am not naming the Semipalatinsk form as a subspecies, because this 
has already been done as far as I know by A.I. Argyropulo in the manuscript 
of his work on pikas of the U.S.S.R. in the "Fauna of the U.S.S.R. " 

GEOGRAPHICAL DISTRIBUTION. Discussed above. 



Subgenus C onothoa Lyon 

1904. Conothoa Lyon M. W. Classification of the Hares and their Allies, Smithsonian Miscellaneous 
Collections, Vol.1, parts 3-4, p. 438. 

Skull rather massive, not flattened in the interorbital and frontal regions 
in most species, where the skull is often convex in profile. Frontal 
fenestrae often well developed. Interorbital space broad. Orbits of 
medium size. Triangular fenestrae of rostrum large and angular. 
Structure of palatine and incisor foramina varying: palatine and incisor 
foramina fused and vomer ventrally free inO. roylei, O. macrotis, 
O. wollastoni, O. nubrica; structure of these foramina 
intermediate inO. rutila because they are connected by a narrow slit; 
palatine and incisor foramina separate and vomer almost covered by the 
premaxillaries in O . erythrotis and O . glover i. Color of summer 
fur, and that of winter fur in some specimens, often with well marked 
rust-red tones. Pads of digits well developed. Vibrissae long, to 95 mm. 



Described from specimen No. 17422 in the ZMAN. 

69 



The species inhabits high mountains. 

Type of Subgenus: Ochotona roylei Ogilby. 

This subgenus contains also the following species: Ochotona 
macrotis Giinth. , O. rutila Severtz. , O. erythrotis Biichn. , 
O. gloveri Thos. , O. wollastoni Thos. , O. nubrica Thos, 

GEOGRAPHICAL DISTRIBUTION, Pamir Mountains, Altai, Ferghana, 
Gissar Range, Dzhety-Su Mountains, Ladakh, Himalayas (to 20,000 ft), Tibet 
and Kansu. 

154. Ochotona (Conothoa) macrotis Giinther (1875). Large -eared 
pika or Indian pika. 

1875. Lagomys m aero t is Giinther A. , Description of Some Leporine Mammals from Central Asia, 

Annals and Magazine of Natur. History, Vol. XVI, ser. 4, No. 93, p. 231. 
1914. Ochotona sacana Thomas O. , On small Mammals from Djarkent, Annals and Magazine of 

Natur. History, p. 572 (vicinity of Przheval'sk). 
1925. Ochotona alpinus (partimj), Shnitnikov, V. N. , Dzhety-Su (Dzhety-Su), p. 129. 
1928. Ochotona ruti 1 a Vinogradov B. S. , Mlekopitayushchie (Mammals), Sbomik: "Trudy Pamirskoi 

ekspeditsii" (Reports of the Pamir Expedition), Issue VIII Zoologiya, p. 8 (partim.'). 

1932. Ochotona macrotis Argyropulo A. M. , Materialy po faune gryzunov Srednei Azii, Pishchukhi 
Srednei Azii (Materials on the Rodent Fauna of Soviet Central Asia. Pikas of Soviet Central Asia). 
Trudy Zoologicheskogo instituta Akademii Nauk, II, pp. 42-45. 

1933. Ochotona macrotis Vinogradov B, S. , Mlekopitayushchie SSSR (Mammals of the U. S. S. R. ), 
p. 81. 

1936. Ochotona macrotis Shnitnikov V. N. , Mlekopitayushchie Semirech'ya (Mammals of Dzhety-Su), 

pp. 258-259. 

NAMES: The large-eared pika (English); Langohriger Pfeifhase (German). 

LOCAL NAMES: Tiin-tshkan (Kara-Kazakh, Kara-Kirghiz), 

TYPE LOCALITY AND DEPOSITION: A. Gunther (1875) reports that he described his new species from 2 
specimens. The first (in a very bad condition) is kept at the British Museum without an accurate label since 
1844. The other, which has to be considered as the holotype, was caught by Biddelph on river Doha (Central 
Tibet, 31" N and 87° E). W. T, Blanford (Scientific Results of the Second Yarkand Mission, Mammalia, 
1879, p. 76) stated that the type of macrotis was caught near Duba, while A. Gunther records 
it from Doha. According to Blanford, Duba lies at the northwestern outskirts of the Kunlun Ranges, 
on the way from Yarkand to Kugiar, not far from Yangi-Divan. I find it difficult to judge who is correct 
about the type locality of this species. These two localities are zoogeographically closely related. It is 
more likely that Doba in Central Tibet is the terra typica of macrotis, in our opinion. 

PRINCIPAL FIGURES: 1) Argyropulo A.I. , Materialy po faune gryzunov Srednei Azii (Materials on the 
Rodent Fauna of Soviet Central Asia), Trudy Zoologicheskogo instituta Akademii Nauk, I, 1932, p. 43, fig. 4 
(skull dorsal, ventral and lateral); 2) Vinogradov B. S. , Mlekopitayushchie SSSR (Mammals of the U. S. S. R. ) 
1933, p. 81, fig. 75 (a) (skull dorsal). 

MATERIAL EXAMINED: 19 pelts and skulls. 

DIAGNOSIS. Skull of average size. Length reaching 47 mm. 
Interorbital space flat and broad (5.2-6 mm). Anterior third of nasals little 
widened; lateral margin of posterior third of nasals almost parallel; 
posterior quarter of nasals depressed. Posterior end of nasals rounded. 
Anterior part of frontals with oval elongated frontal fenestrae. Vomer 
free, not covered ventrally by the premaxillaries; which diverge 
anteriorly at an angle. Palatine and incisor foramina fused (Figures 40- 43). 

Vibrissae mixed black and white, length 55-70 mm. Ears rather long, 
to 31 mm. 

Color of summer fur on back of various shades of grayish brown with 
an ocher tinge. Winter fur of back dense, fluffy pale gray with a straw- 
yellow tinge. 



70 



Body length to 200 mm; posterior feet 32-34 mm. 

ADDITIONAL DESCRIPTION. Color of summer fur on back of various 
shades of grayish brown with a more or less marked ocher tinge, general 
tone drab with an admixture of buff-brown, sometimes saccardos -umber 
or tawny-olive. Rusty tones intermediate between pinkish cinnamon and 
cinnamon-buff distinct on head, cheeks and flanks; occiput with a generally 
pale rust-yellow (intermediate between light pinkish cinnamon and pinkish 
buff) indistinctly marked transverse stripe. Belly dirty-whitish, sometimes 
with a slight yellowish tinge. 



87 





FIGURES 40, 41. Skull of large-eared pika, Ochotona macrotls Gunther, dorsal and ventral. 
No. 17950 J, Pamir, Kara-Kul, 14 June, 1928 

Drawing by Yu, A. Kostylev 

88 Winter fur on back dense, fluffy, generally pale, grayish with a straw- 
yellow tinge (avellaneous with pinkish buff and cinnamon-buff tinge, Ridgway, 
1912). Rust tones (cinnamon-buff) better marked on top of head. Cheeks 
with an ash-gray tinge; fur on flanks yellowish ocher (intermediate 
between pinkish buff and cinnamon- buff). Ears whitish. Inner side of 
anterior margin of ears dark ash-gray. Belly pure white; bases of hair 
on belly deep gray. 

SYSTEMATIC NOTES. O. Thomas (1914) described Ochotona 
sac ana from seven specimens from the vicinity of Przhevalsk. V.N. 
Shnitniknov (1936) assumes that these specimens were caught in Terskei 
Ala Tau, perhaps in the valley of the Karakol river near which Przhevalsk 
lies. 



71 




FIGURES 42,43. Lateral view of skull and mandible of large-eared pika, Ochotona 
macrotis Giinther, No. 17950. 

O. Thomas remarks that O. sacanais very closely related to O. 
macrotis. Series of O . sac ana can be easily distinguished, according 
to him, from true macrotis (in winter fur) by the rust (cinnamon) tones 
of the flanks and lower part of the back, and by the more blackish outer 
parts of the ears. The measurements ofO. sacana according to 
Thomas are: length of body and head 200 mm; anterior feet 33 mm; ear 
28mm. 

Maximum skull length 47 mm; condylo-incisive length 44 mm; 
zygomatic width 22.5mm; nasal bones 16X5.5; interorbital space 5.3mm; 
width of skull at level of auditory meatus 18 mm; palatine foramina 
13.5X4.7; width of palatine bridge 1.8 mm; length of upper tooth row 
(alveoli) 9 mm (type caught in December 1913). 

According to Thomas's description and his remark that O. sacana 
and O. macrotis are very closely related (for instance, there are no 
cranial differences between them) we consider that the identity of 
macrotis and sacana as definitely established (cf. A.I. Argyropulo, 
1932). 

We have not enough material to prove that sacana differs from the 
nominal race sufficiently to be considered as a subspecies of macrotis. 
However, this is probably correct if such an expert mammalogist as O, 
Thomas thinks so. 

The skull of Ochotona macrotis differs from that ofO. rutila 
in the following characters: 1) generally smaller size and weight; 2) 
steeper frontal contour; 3) the short rostrum; 4) presence of frontal 
fenestrae; 5) broader connection between the palatine foramina. 



72 



O. macrotis, O. rutila, and O. erythrotis form a group 
well defined by some characters and the general form of skull. O . 
erythrotis is found in Kansu and in the mountains of northeastern Tibet, 
and Burhan-Budda Range. 

The skull ofO. erythrotis is morphologically intermediate between 
that of O . rutila and macrotis; there are frontal fenestrae in 
erythrotis which are absent in rutila. The structure of the palate 
in erythrotis differs rather sharply, erythrotis belongs to the 
subgenus pika (as understood by Thomas) according to the following 
characters: premaxillary processes partly covering the vomer so that it is 
little visible ventrally (see Figures 44,45). 

Finally, it should be mentioned that the Himalaya -Tibet O . roylei 
Ogilby is very small, its color markedly different from that of macrotis 
and rutila. Its skull resembles that of m ac r ot i s (on a small scale); 
the structure of the palate is the same as in macrotis. 

GEOGRAPHICAL DISTRIBUTION. The large -eared pika occurs in 
Russia on Peter the Great Range (2980m) and in eastern Pamir. According 
to recent observations of M. P. Rozanov (1935) O . macrotis occurs 





FIGURES 44,45. Skull of Ochotona erythrotis Buchner (doisal and ventral), No. 1500 d", Burhan- 
Budda Range. N. M. Przheval'skii, ZMAN collection 

Drawing by Yu. A. Kostylev 



73 



throughout eastern Pamir . It was observed near the Pamir station and near 
Kyzyl-Rabat (at an altitude of 4000 m). There are specimens from Lake Range- 
Kul and Kara-Kul in the ZMAN collection. O. macrotis also occurs in 
mountains near Karakol (Pr zhe valsk), on the Ter skei Ala Tau, in the mountain 
region to the south of Naryn and in central Tien Shan. V.N. Shnitnikov 
(1936) found it in the valley of river Kuelyu (Kuilyu), in the Sardzhas Range, 
and also on the Kargalyk Pass connecting the Toguz-Torau with the valley 
of river Alabuga, i.e., not far from the Fergana frontier. A.I. Argyropulo 
(1932) records specimens from the rivers Tandyk and Terek-Su (south 
of Gulcha)in the ZMAN museum. 

O. macrotis is thus sporadically distributed in the mountain region 
of Central Asia in isolated colonies. V.N. Shnitnikov (1936) writes the 
following about Dzhety-Su: "The distribution of O . macrotis (like 
O. rut i la) in this region is rather strange. It lives north of Issyk-Kul, 
then it is found along the Alexander [Kirghiz] Range, then westward and 
^-t.urns south in western Tien Shan, carefully bypassing the boundary of 
Dzhety-Su. Then it turns again east and south, and occurs from the Alai 
and Pamir westward to the Gissar Range. The distribution of O . 
macrotis in the southwestern corner of Dzhety-Su, is wedged into the 
range of O . rutila. 

"The ranges of the two species ought, therefore, to overlap, or at least 
adjoin each other. We should look for such places in the region south of 
Issyk-Kul, in the Dzhumgal or Kabak Mountains, and perhaps also in the 
Susamyr Range. " 

Outside of the U.S.S.R., the large-eared pika occurs on the northeastern 
Kunlun, in the mountains between Yarkand and Kushar, in the mountains 
along Raskam-Daryanear Ley (on the Indus), in Ladakh, near Shushal and 
in Central Tibet (Doha). 

MODE OF LIFE. According to V. N. Shnitnikov (1936) O . macrotis 
lives mainly in screes in spruce forests. It is very numerous in some 
places, for instance in the valley of river Kuelyu. V.N. Shnitnikov found 
it here on screes in the spruce forest. He found many cut leaves of 
Corydalus sp. spread for drying on stones. He found also very many 
cut leaves of A s t r a galus frigidus. V.N. Shnitnikov did not find 
"haystacks", and he thinks that O. macrotis deposits hay in storerooms 
among stones. The pikas were rather cautious and cried rarely. 

M. P. Rozanov (1935) writes that he saw pikas living in crevices among 
stones, in burrows near Kyzyl-Rabat. 

"Near the Kyzyl-Rabat Mountain Pass, we saw a rock with many pika 
excrements and stores of dry grass in the cracks. We did not see the animals 
themselves. We caught one of the animals in the Nimkhan plot, near the 
estuary of river Mats. We saw a large colony in large screes near the 
river Mats dam, and caught some. The pikas live everywhere together 
with the Pamir high-mountain vole (Alticola argentatus) and the gray 
hamster (Cricetulus migratorins)." 

155. Ochotona (Conothoa) rutila Severtzov (1873). Red pika 

1873. Lagomys rutilus Severtsov, N. A. , Vertikal'noe i gorizontal'noe raspredelenie turkestanskikh 

zhlvotnykh (Vertical and Horizontal Distribution of Turkestan Animals): Izvestiya Obshchestva lyubitelei 
estestvoznaniya, VIII (2): 8 3. 



74 



1879. Lagomys rutilus Severtsov N. A. , Zametki o faune pozvonochnykh Pamira (Notes on the Vertebrate 

Fauna of the Pamir), Zapiski Turkestanskogo otdeleniya Geograficheskogo obshchestva, I, p. 64. 
1927. Ochotona ruti 1 a Kashkarov D. N. , Rezul'taty ekspeditsii v raion ozera Sary-Chilek fResults of 

an Expedition to Lake Sary-Chilek), p. 101. 
1930, Ochotona rutila Vinogradov B. S. , Rukovodstvo k opredeleniyu gryzunov Srednei Azii (Key to 

Rodents of Soviet Central Asia), p. 49. 
1932. Ochotona rutila Argyropulo A. I. , Materialy po faune gryzunov Srednei Azii (Materials on the 

Rodent Fauna of Soviet Central Asia), Trudy Zoologicheskogo instituta Akademii Nauk, I, pp. 37-42. 

1935. Ochotona rutila Vinogradov B. S. , Otryad gryzunov (order Rodentia), Sbomik: "Zveri 
Tadzhikistana" (Animals of Tadzhikistan), Izdatel'stvo Akademii Nauk, pp. 250-251. 

91 1935. Ochotona rutila Rozanov M. P. , Mlekopitayushchie Pamira (Mammals of the Pamir), 
Tadzhikskaya kompleksnaya ekspeditsiya, Issue XXXII, p. 55. 

1936. Ochotona rutila Shnitnikov, V. N. , Mlekopitayushchie Semirech'ya (Mammals of Dzhety-Su), 
pp. 254-256. 

NAMES: The red pika (English); Der rote Pfeifhase (German). 

LOCAL NAMES: tyni-tyshkan (Kazakh, Kirghiz). 

TYPE LOCALITY AND DEPOSITION: Type collected in 1867 by N. A. Severtsov in mountains near . 

Vemyi — now Alma-Ata (at an altitude of 7000-8000ft). Type number S. 11,074, kept in the Moscow '^ 

Zoological Museum. 

PRINCIPAL FIGURES: 1) S chaff E. , Ueber Lagomys rutilus, Zoologische Jahrbuch. , Bd. II, 
1887, S. 65-72; 2) Argyropulo A. I. , Materialy po faune gryzunov Srednei Azii (Materials on the Rodent 
Fauna of Soviet Central Asia), Trudy Zoologicheskogo instituta Akademii Nauk, I, 1932, pp. 37-42, fig. 2 
(foramina incisiva); p. 39, Fig. 3 (skull dorsal, ventral and lateral); 3) Vinogradov B. S. , Gryzuny 
(Rodents), Sbomik: "Zveri Tadzhikistana" (Animals of Tadzhikistan), Izdatel'stvo Akademii Nauk, 1935, 
p. 250, fig. 85; 4) Shnitnikov, V.N. , Mlekopitayushchie Semirech'ya (Mammals of Dzhety-Su), 1936, 
p. 254, fig. 67 (map of geographical distribution). 

MATERIAL EXAMINED: 15 pelts and skulls. 

DIAGNOSIS. Skull relatively large; in fully grown and old specimens, 
its length is often more than 48 mm, sometimes 52 mm. Interorbital space 
broad, 5.7-6.9 mm, rather flat, slightly convex in the middle of profile, 
without any lateral crests. Anterior third of nasals with marked spoon- 
shaped widening. Lateral contours of nasals gradually narrowing 
posteriorly; each nasal ends in a separate curve posteriorly. Frontals 
without fenestrae. Premaxillaries in palatine region close together but 
not fused. The vomer is therefore completely visible ventrally, and the 
incisor and palatine foramina are not separated, but connected by a narrow 
space (Figures 46-49), 

Vibrissae long, 80-94 mm; some of them dark brown, the others white; 
the longest vibrissae have dark bases and broad white tips. Soles whitish 
or grayish white. Ears without marked light borders. 

Color of summer fur on back (see pi. I, upper illustration) rich rust- 
reddish (intermediate between cinnamon and sayal-brown, Ridgway, 1912, 
pi. XXIX); flanks more yellow with a cinnamon-buff tinge; a broad 
yellowish white collar behind the ears; belly whitish with a rusty tinge; a 
dull rust-red transverse stripe on the chest. Winter fur ash-gray with blackish 
brown speckling caused by the brown-blackish hair tips. 

Body length 215-230mm; posterior feet 35-38mm; ear 25-28 mm. 

ADDITIONAL DESCRIPTION. The skull of the red pika differs sharply 
from that of O . (Pika) alpina Pall, in the following characters: l) 
skull more convex dorsally in profile (especially in the interorbital space); 
2) longer orbits; 3) anterior part of nasals wider; 4) bullae osseae 
shorter, their form sharply different from that of O . alpina; 5) different 
form of palatine foramina. 

The skull of O . erythrotis Biichn. differs markedly from that of 
O. rutila in the following characters: l) presence of frontal fenestrae; 



75 



92 





FIGURES 46, 47. Skull of red pika, Oc hot on a rutila Severtz. 24 Aug. 1929. Alexander [Kirghiz] 
Range, collection of the Moscow Zoological Museum 

Drawing by Yu. A. Kostylev 




FIGURES 48, 49. Lateral view of 
skull and mandible ofOchotona 
rutila Severtz. (Same specimen 
as in Figures 42, 43) 



76 



93 2) different form of nasals ; 3) different structure of palate. The 
external appearance of O . erythrotis resembles that of rut i la 
(especially the color of the summer fur). These species belong to the 
same subgenus. 

Color of summer fur on top of head and on back intense rust-red 
(intermediate between cinnamon and sayal-brown). Cheeks and region 
around the eyes of the same color. Nose anteriorly more yellowish. 

A pale, indistinct, rust-yellow stripe behind the eyes in the occipital 
region; a well developed collar behind the ears which is narrower, and less 
well marked between the ears. Color of flanks more pale and yellowish, 
intermediate between cinnamon- buff and pinkish buff. A dull rust -red 
stripe on the chest between the forelegs. Chin and top of head white; 
belly whitish with a slight rust-red tinge which is especially well marked 
in the midline of the belly. Inner side of ears with grayish white hair; 
a tuft of yellowish white hair at the anterior inner side of the base of the 
ears. Outer side of ears grayish. Paws with rust-red hair. Feet whitish, 
with a slight rust tinge dorsally, soles whitish gray. 

General tone of winter fur rather pale, gray with blackish brown 
speckling caused by the black-brown tips of hairs. Bases of long winter 
hair deep slate -gray or blackish. Color of flanks like that of back 
(slightly duller and grayer); belly whitish; gray bases of hair visible 
through hair tips on belly. No rust-red stripe on chest, the color of the 
chest is as whitish as that of the belly. 

Color of young on back light ocher with reddish tones on head and flanks. 

GEOGRAPHICAL DISTRIBUTION. N.A. Severtsov (1873), who was the 
first to collect and describe this species, writes as follows: "A young and 
an old specimen were caught in late May 1867 in the mountains near Vernyi 
[Alma-Ata] at an altitude of 7000-8000 ft. An old and molting specimen 
was caught previously, in the ravine of river Kara-Bury to the south of 
Aulie-Ata [Dzhambul]* (in late June 1864) at an altitude of about 6500 ft. " 
A.I. Argyropulo (1932) records the localities where the specimens of the 
ZMAN collection were caught as follows: the Kirghiz (Alexander) Range, 
ravine of the river Tuyuk (Shnitnikov); vicinity of the Alma-Ata (the 
Almaatinskoe Lake); Alai (northern slope of Pengalbai Mountain Pass 
(Molchanov)); the Gandakush landmark, 64 km to the south of Fergana 
(Zhinkin); near the river Kshtut-Dar'ya'^'*; Shaar Mountain at an altitude 
of 7000 feet (Shakhrizyabs); Marguzar-Kul; upper reaches of river 
Shinkt ; the former Samarkand County (N.A. Severtsov); the Gissar 
Range on Iskander-Kul (Russov); Pamir (Russian- German expedition). 
_ . There is a good series from the Kirghiz (Alexander) Range in the 

Moscow Zoological Museum. D.N. Kashkarov (1927) found this species in 
large numbers near Lake Sary-Chilek, at an altitude of 1925 m in the 
northern part of the former Namangan County between spurs of the Chatkal 
Range . 

According to N. P. Rozanov (1935) this species was found in the western 
part of the Pamir, on rocks near the Shid settlement on the Pyandzh. It 
apparently does not occur in the eastern Pamir. According to V.N. 
Shnitnikov (1936) the red pika occurs in the Ala-Tau, on the Alexander 



* On the Kirghiz (Alexander) Range. 
''* A tributary of river Zeravshan. 
f In the Gissar Range, 



77 



(Kirghiz) Range and on the northern slope of the Kungei Ala Tau. 
V.N. Shnitnikov has records of this species from the southern slope 
of this range, but he has no specimens from this locality. The red 
pika does not occur to the south of Lake Issyk-Kul according to him. 
D.N. Kashkarov's record ofO. rutila from the Terskei Ala Tau 
seems erroneous, as this region is inhabited byO. macrotis. The 
range ofO. rutila thus includes the following localities: the Zeravshan 
Mountains; Gissar Range; Alai Range; the wester Pamir; Chatkal Range 
on the Kirghiz (Alexander) Range; Trans-Ili Ala Tau; Kungei Ala Tau. 

V.N. Shnitnikov mentions a species of pika from the Dzungarian Ala 
Tau (valley of river Kora, Kyzyl-Agach). However, no specimens were 
caught in this region, and it is therefore not known which species occurs 
there. 

MODE OF LIFE. There are very few data on the biology of this species. 

A few interesting data are given by V.N. Shnitnikov (1936). The red 
pika generally occurs on high mountains (above 2000m) and screes. Its 
distribution is sporadic. It may be present in one ravine and be absent in 
another, although conditions in both are the same. 

According to V.N. Shnitnikov, this species occurs from the middle of 
the spruce forest belt to the subalpine meadow region. It does not occur 
at higher altitudes in the juniper region (Juniperus sabina, in the 
mountains near Alma-Ata). On the other hand, N. A. Severtsov (1873) 
states that the red pika "lives among juniper". V.N. Shnitnikov's statement 
seems therefore not quite accurate. 

Shnitnikov writes: "Pikas move with great agility among screes and 
heaps of stones, giving the impression of moving shadows. Although it is 
quite natural that not the slightest sound should be heard even when the 
animals move quite close, the absolute silence of their movements seems 
unnatural. If one stands or sits quietly among the stones, the pikas 
approach qmte close. I know of a case in which an animal was killed with a 
whip. " 

The red pika is very quiet. V.N. Shnitnikov did not hear its voice even 
once. 

V.N. Shnitnikov writes: "The pikas are diurnal. Sometimes they can 
be obsei'ved throughout the day. However, they are most active at dusk, 
and feed and carry grass in the morning, after sunrise. However, they 
are less active than in the evening. 

"During the day, they are seen only accidentally, because they generally 
lie motionlessly on a stone like a sphinx. Sometimes they stretch their hind 
legs, and are then difficult to see. They are spotted only when approached 
closely and they run away among the stones". 

The red pika prepares food stores and hay for the winter, which it hides 
under stones. It does not make "haystacks". 

O. rutila occurs sporadically and rarely among stones along the 
coast of Lake Issyk Kul (at an altitude of 1750m); and in the Issyk Kul 
woodland in the Alma-Ata State Reserve, about 45 km east of Alma-Ata. 
The pikas inhabit stone heaps and breccia along the western coast of Lake 
Issyk Kul, and also breccia near the Maloe or Nizhnee Lake. 

O. rutila occurs sporadically also in small numbers in screes along 
the road to the Talgar Mountain Pass at 2200 m in subalpine meadows. 
Juniper and dwarf birch trees grow where pikas were found. 



78 



96 




FIGURE 50. Red pika, Ochotona rutila Severtz. 



Drawing by K. K. Flerov 



I had the opportunity of observing a single pika for a long time on 
29 July 1938, between 1 and 5 p.m. The animal was very active. With 
graceful and silent jumps, it hopped over the stones, climbing on stone 
heaps to collect hay. After a few minutes, it entered its burrow in a 
crevice among stones with a plant it had just cut (usually Epilobium 
or a branch of wild cranesbill). The plants were piled near the entrance 
of the burrow. There were several handfuls of fresh grass under and near 
a stone. 

It was very interesting to watch the graceful movements of the beautiful 
animal. After it reached the burrow, it stopped, examined the people 
sitting near it, moved its long vibrissae and calmly observed the intruders 
with large shining black eyes. It swiftly jumped into a crevice among the 
stones to deposit the newly cut plant. The animal is very trusting, and lets 
itself be approached to 3-4 steps (pi. II). 

Some sporadic colonies, each of a few individuals, were found among 
stones along the coast of the Bolshoe Almatinskoe Lake (in the Trans Ili 
Ala Tau, at 2800m). 

A.I. Argyropulo (1932) thinks that the young are born in the first ten days 
of June (in Lake Iskander-Kul). This seems too late; the young are 
probably born in May. 

We have few data on the molt of this species. In his book "Mammals of 
Dzhety-Su" (Mlekopitayushchie Semirech'ya), 1936, pp. 256-257, V.N. 
Shnitnikov gives contradictory data. In his opinion, "the pika enters the 
winter in its reddish fur". This statement is contradicted by his remark 
that the pikas wear a very long and dense light gray fur* in early spring, 
which is apparently correct. 



* In Shnitnikov's opinion, this is contradicted by the statement of N. A. Severtsovthathe caught a fiery red 
specimen in late May. N. A. Severtsov has not mentioned such a specimen. On the contrary, he mentions 
a specimen caught in late May wearing its old fur and strongly molting. This molting specimen is in the 
collection of the Zoological Museum of the Moscow University. 



79 



97 



This statement is also contradicted by Shnitnikov's remark that "the 
complete molt from reddish fur to gray fur takes place in late autumn". 

If this is true, how do the pikas "enter winter" in a reddish fur? 

A.I. Argyropulo (1932) observed that intense molting on the head (as 
in other species of Ochotona) was observed in specimens caught in May. 
Short-haired reddish patches are clearly visible on the worn, long winter 
fur. The new fur is clearly seen on the head and flanks. Such a specimen 
is in the collection of the Zoological Museum of the Moscow University 
caught on 22 May 1922 on the Kirghiz (Alexander) Range. A specimen 
caught by N. A. Severtsov near Alma-Ata (late May 1867, Old Style) was 
in strong spring molt. Shreds of rust-red fur are clearly seen on top of 
the head, the anterior part of back, chest, flanks and partly on the legs. 
A.I. Argyropulo (1932) observed that the lower part of back molts last as 
a rule. 

The beginning of molt into winter fur could already be seen in specimens 
caught in late August. 

It is difficult to describe the molt from so little material. It apparently 
takes an opposite course to the spring molt, i. e. , the lower part of the back 
molts first. A.I. Argyropulo (l932) observed that patches of winter fur 
appeared all over the body without any regular "molting foci". In my opinion, 
Argyropulo's observations gave negative results for lack of material*. 

We have no data on the winter life of the red pika. 

The ermine is one of the most important enemies of this rodent according 
to V.N. Shnitnikov (1936). The pikas disappeared from Bolshaya Almaatinka 
(near Alma-Ata) because they were hunted by the numerous ermines. After 
the extermination of the pikas in Bolshaya Almaatinka the ermines began 
feeding on the numerous voles (A It i c o la worthingtoni andEvotomys 
f r at e r ). 



Species of the Ochotona pus ilia and 
O. thib etana group 

The species of this group should probably be considered as a separate 
subgenus (vide infra). 

The characters of the group are as follows: 

Skull in interorbital- frontal region slightly convex. Frontal fenestrae 
absent. Interorbital space moderately wide. Orbits small. Rostrum very 
short. 

Palatine and incisor foramina of varying form. They are connected by 
a narrow slit inO. pusilla and inO. thibetana both foramina are 
fused. 

Color of summer fur very peculiar, a rather dark gray-brown, grayer in 
some species, with marked straw speckling on flanks and trunk. Belly 
whitish gray. 

Vibrissae very short, not more than 45 mm long. 

The species inhabit valleys in steppes and plateaus. 

Type species: Ochotona pusilla Pall. 



* Examination of new material collected in 1939 near the Bolshoe Almatinskoe Lake suggested that autumn 
molt begins at the lower part of the back and in small patches in the middle of the back. 



80 






Red pika (Ochotona rutila) carrying grass to its burrow. Trans-Ill Ala Tau, near the Talgar 
Mountain Pass, 29 July 1938 

Photograph by S. I. Ognev 



98 



Species: O. thibetana Milne-Edw. (with subspecies); O. cansa 
Lyon (with subspecies); O. so re 11a Thos. , O. syrinx Thos.; O. 
foresti Thos. apparently belong to this group. 

DISTRIBUTION. From the left bank of the Volga through the steppe 
(non-desert) region of Transvolga and Transural to Omsk, Altai Region and 
Zaisan Basin trough. Some species are also known from Sikkim, Kansu, 
Shansi, Szechwan, Shensi, Yunnan. 

The exact systematic relationshipbetweenthese forms requires critical 
examination of the material. 

156. Ochotona pusilla Pallas (1768). Small pika or chekushka. 

1768. (1769). Lepus pusillus Pallas P. S. , Descriptio Leporis pusilli, Novi Commentarii Academiae 

Petropolitanae, t. XIII, pp. 531-538. 

NAMES: Zwergpfeifhase (German). 

LOCAL NAMES: zemlyanoi zaichik (ground rabbit)-, chivkun; "zhavoronok" ("sky lark"): or chekalka 
(Russian); chokchot, itchishkan (Tatar, imitating voice of animal); timersak, sepsan (Bashkir); kaik 
(Chuvash); kuzla (Kalmyk). 

PRINCIPAL FIGURES: 1) Pallas P. S. , Novi Comm. Academ. Petrop. , 1768(1769), tab. XIV, figs. 1-8 
(entire animal, head, paws, intestine); 2) Pallas P. S. , Novae species Quadrupedum e Glirium ordine, 
1778, pp. 31-45, tab. I (drawing of entire animal); tab, IV, fig. 3 (ear), fig. 4 (skull frontal), fig. 5 (skeleton) ; 
3) Schreber I. Ch. , Die Saugethiere, Erlangen, B. IV, 1792, S. 906-910, tab. CXXXVIl (color plate of entire 
animal); 4) Pallas P. S. , Zoographia Rosso-Asiatica, 1811 (1831), pp. 151-152, tab. XII (color plate of entire 
animal); 5) Simashko, Yu. , Russkaya fauna (Russian Fauna), II, 1851, pp. 790-791, pi. 63, fig. 1 (color 
plate of entire animal); 6) Argyropulo A. I. , Opredelitel' gr^-zunov Ural'skoi oblasti (Key to Rodents of the 
Uralsk Region), Moskva, 1931, p. 8, fig. 33 (skull ventrally); p. 62, fig. 78 (skull laterally); 7) Argyropulo 
A. 1. , Materialy po faune gryzunov Srednei Azii (Materials on the Rodent Fauna of Soviet Central Asia), 
Trudy Zoologicheskogo instituta Akademii Nauk, I, 1932, p. 55, fig. 7 (skull of Ochotona pus i 1 1 a pusilla 
Pall, dorsal, ventral and lateral); p. 56, fig. 8 (skull of O. p. angustifrons Argyrop. dorsal, ventral 
and lateral). 

MATERIAL EXAMINED: 35 pelts with skulls. 

DIAGNOSIS. Skull small, length in fully grown and old specimens not 
longer than 43 mm, usually 40 mm. Interorbital space of moderate breadth 
in comparison with size of skull, 3.6-5 mm. Interorbital space slightly 
arched. There are small facets on both sides of the interorbital space; 
and the middle part of the interorbital space seems therefore depressed. 

Nasals usually only slightly widened anteriorly; posterior parts of 
nasals broad. Frontals without fenestrae. Rostrum very short. Distance 
between inner margins of alveoli of last molars usually as long as the 
upper diastema measured to the alveoli of the second pair of incisors. 
Vomer not completely covered by premaxillaries. The vomer is therefore 
visible ventrally. Palatine and incisor foramina connected by a narrow 
space (Figures 51-54). 

Vibrissae partly short and black-brown and partly white and longer, not 
longer than 45mm. Soles brownish, with dense hair covering the pads of 
the digits unlike in all other species described. Ears with distinct light 
margins. 

Color of summer fur dark grayish brown with more or less marked 
straw speckling on back and flanks; belly dull whitish gray. The winter 
fur slightly differs from the summer fur in its paler color. 

Body length at most 200 mm, usually about 170 mm; hind feet at most 
30mm (usually about 27 mm). 



82 




83 



ADDITIONAL DESCRIPTION. In addition to the cranial characters| 
described, the large size of the strongly swollen bullae osseae should 
be mentioned. They are slightly compressed laterally, and seem therefore 
larger than those of other species, for example O. hyperborea, the skull 
of which is of about the same size as that ofO. pusilla. Orbits small 
and rounded. The brain case seems more elongate than in other species 
because of the short rostrum. 





FIGURES 51, 52. Small pika, or chekushka, Ochotona pusilla angustifrons Argyiop. According 
to type no. 17421, river Dzhancha, former Kedei subdistrict, former Karkaralinsk County, Kazakhstan, 
Filatov, ZMAN collection 

Drawing by Yu. A. Kostylev 

SYSTEMATIC NOTES. The small pika differs sharply from the other 
species of Ochotona by the structure of the skull, color, the very hairy 
soles etc. 

It would be justified to establish a separate subgenus for this species 
because of its separate position among our species. However, the small 
pika is closely related to the Tibet pika Ochotona thibetana Milne - 
Edw. (from Szechwan, Mupin). Ochotona thibetana has the 
following subspecies: O. t. sacraria Thos., southern Szechwan, O.t. 
sikimaria Thos. , Sikkim, O. t. z appe y i Thos. (Tatsienlu [Kangting]), 
The following forms are closely related to thibetana, and therefore 
also to pus i 11a: Ochotona cansa Lyon (Kansu) with the subspecies 
O. cansa morosa Thos. (Shensi); O.c. stewensi Osgood 
(Szechwan); O. sorells Thos. (Shensi, Ningwufu); O. syrinx Thos. 
(southwestern Shensi); O. foresti Thos. (Yunnom, Liliang). 



84 



100 Most of these subspecies would have to be studied to give 

description of a proposed subgenus. However, I know only one form, 
O. thibetana Milne-Edw. well enough (Figures 55-58). 

We therefore establish only the pus ilia- thibe tana group (vide 
supr-a) and postpone the creation of a subgenus until more material 
becontes available. 

GEOGRAPHICAL DISTRIBUTION. The range of this species includes 
the Trans -Volga steppes to about 54° N (not including the Astrakhan 
Desert), the Obshchii Syrt, the Orenburg Territory in the north up to 53°30', 
and the Omsk, Semipalatinsk and Cisaltai regions. Its range reaches the 
Zaisan Depression and the southern slopes of the Tarbagatai in the southeast. 

The first description of the range of this species was given by P.S. Pallas 
(Novae species Quadrupedum, 1778, pp. 31-45). According to Pallas, 
Ochotona pusilla occurs in the southern foothills of the Urals, and is 
common between Kama and Samara. It is also found along the left bank of 
the Volga between Samara and Ural. It is very numerous between the Urals 
and Uem. It is absent in the forest zone and in the deserts in the lower 
reaches of the Volga and river Yaik (Ural). It occurs in the valley of the 
river Ilovlya* on the right bank of the Volga, Much new material has 
accumulated since the classical work of Pallas. 




FIGURES 53, 54. Ochotona pusilla angustifrons Argyiop. After type 
No. 17421. 

O. pusilla was recorded from the former Buguruslan and Buzuluk 
counties in the central Trans -Volga region. According to B. S. Bazhanov 
("Materials on the Study of the Samara [Kuibyshev] Territory" (Materialy 
po izucheniyu Samarskogo Kraya), issue 5, 1928, pp. 19-20) it occurs 15 km 



+ This species does not occur today on the right bank of the Volga. 



85 



north of the middle reaches of the river Buzuluk. It was also observed in 
the upper reaches of river Kamelik (a tributary of river Bolshoi Irgiz) 
north of Obshchii Syrt and along the river Irgiz (Eversman, 1850, 1855). 
A specimen from the vicinity of Spassk is in the ZMAN collection. 
(former collection of Professor Eversman). However, the occurrence of 
O. pusilla at such a northern locality (almost near 55°N) seems doubtful, 
as it was not found in the Tatar Republic in spite of intensive search by 
workers of the Volga-Kama Biological Station. 
101 O. pusilla is common on the Obshchii Syrt. According to N. A. 
Zarudnyi (1897) it is common here in many places, and also in the upper 
reaches of the river Salmysh and the sources of river Chagan. He writes 
(1897) as follows: "East of the northern Obshchii Syrt, in Bashkiria, I 
found this species in very small numbers between the villages Isyangulova 
and Kugarchi. " According to S. V. Kirikov, it is found in the upper 
reaches of river Belaya, near the Irgizly hamlet farm. 

According to N.A. Zarudnyi (1897) it is common in the Orenburg District 
in places with high hillocks along the middle reaches of the Ural river, 
between Orsk and Uralsk. It was found in large numbers in the northern 
Mugodzhary Hills, in the Kyrchaki site, in the Donguz and Vetlyan 
Mountains, and on hillocks near Sulyuk-Kul. B.A. Kuznetsov (1928) 




FIGURES 55, 56. Skull of Tibetan pika, Ochotona thibetana Milne- 
Edw. , No. 157, May 1894, river Danho 

Drawing by Yu. A. Kostylev 

recorded O. pusilla from near Uralsk and near the Saverov farm 
(former Orsk Country). In the ZMAN collection are specimens from 
the vicinity of Brody and the village Klyuchevka, from the former 
Kashira County, from the former Orenburg Province, from the 
vicinity of Orenburg and from river Or. According to A.I. Argyropulo 



86 



(1930), the most northern locality in the southern Urals where O. pusilla 
was caught is the Magnitnaya Railway Station (between 53 and 54°N). 




102 



FIGURES 57, 58. Lateral view of skull and mandibula ofOchotona 
thibetana Milne - Edw. , No. 1577; see Figures 55, 56 

N.P. Sabaneev (1872) wrote that O. pusilla occurred in the former 
Shadrinsk, Ekaterinburg and Chelyabinsk counties, at55°30'N. He claimed 
that the northern boundary of its range in the former Shadrinsk County 
reached 56°N in the east. In "Vertebrates of the Central Urals" 
(Pozvonochnye Srednego Urala), 1874, L. P. Sabaneev does not mention O . 
pusilla . He apparently found that his previous records were erroneous. 
It occurs farther east in the steppes to the south of Omsk according to 
Sotnikov. 

B. A. Kuznetsov (1932) writes that it is widely distributed in the 
Semipalatinsk District and in the Altai. It was first observed between 
Semipalatinsk and Sergiopol [Ayaguz] along the Turkestan-Siberian 
Railroad. It was repeatedly found near Sergiopol [Ayaguz]. It is very 
common in the foothills of the western Tarbagatai, near Znamenka and 
the village Podgornyi. It was observed near Zmeinogorsk (c.f. Pallas, 
Meyer, Finch, A.M. Nikol'skii), and it was caught 90km south of 
Zmeinogorsk (N. F. Kashchenko, 1900). 

According to V. A. Selevin (inlitt. 1935) it is found in the western Altai, 
the southern Kalbinskii Range, along the northern Turkestan-Siberian 
Railroad, in the Karaganda Region, along the hills south of the Spasskii 
Zavod to the border of the desert. A specimen caught by A, P. 
Razorenova in the Krasnoshchekovo District in the western Altai, near the 
village Kamenkovo (east of the river Belaya, a tributary of the Charysh) is 
in the Zoological Museum. It was recorded from the Zaisan depression by 
B.S. Vinogradov (1933). 

It occurs in Kazakhstan in the eastern part of the Kazakh plateau in the 
grass-Artemisia steppe between Karkaralinsk and Kounrad, (A.M. Belyaev, 
1934). Specimens caught along the river Dzhamcha and near the village 
Chelym in the former Karkaralinsk County are in the ZMAN collection. 



87 



If the distribution of this species in the past is compared with its recent 
distribution, it appears that its range is gradually receding. 

Fossil remains of this species were found in the Pleistocene of Germany, 
Austria, France, Belgium, and England (F. Brandt, I. Woldrich, E. L. 
Trouessart, A. Nehring et al, ) and in Quaternary deposits in the Crimea, 
in the former Kiev Province (I. G. Pidoplichka) and in old marmot burrows 
in the former Bobrov County in Voronezh Province (S.I. Obolenskii). 
All this clearly shows that the range ofO. pusilla extended considerably 
further west in the past. Skeletal remains ofO. pusilla found in Western 
Europe seem to be older than those found in the Ukraine and in the Voronezh 
Region. 

O. pusilla still occurred on the right bank of the Volga, in the valley 
of river Ilovlya, a tributary of the Don, in Pallas' time (Novae species 
Glirium, 1778, p. 31. ). 

Various data indicate that this species becomes constantly rarer in the 
Trans -Volga and in western Siberia. 

MODE OF LIFE. B. A. Kuznetsov (1928) gives the following data on the 
habitats ofO. pusilla in the Orsk Region: "It is very difficult to 
determine the favorite habitat of the pika. One can hear the loud cry of 
this animal everywhere in the evening, from a haystack, from wheat fields, 
or from a ravine at dusk. I heard their cry repeatedly in the middle of a 
farm, where they live in weeds near fences. 
103 "The pika is heard everywhere, but it is very difficult to see because 
of its secretive habits. It lives in dense thickets, in wheat fields, wild 
cherry brush or among weeds along fences in a village. " 

Further observations on the habitats of the small pika in the 
Semipalatinsk District are found in "Rodents of the Semipalatinsk District" 
by B. A. Kuznetsov (Gryzuny Semipalatinskogo okruga. — Byulleten' 
Moskovskogo Obshchestva Ispytatelei Prirody, XVI (1-2): 1 14, 1932). 

"The pika lives most frequently in thickets of peashrub, wild rose, 
Filipendula and others in the valleys. It is most numerous in the mixed 
grass belt. It is very rare on the border of the Balkhash Steppes. Pikas 
are also rare on the summits of hills. O. pusilla is very common near 
the village Podgornyi, among shrubs in valleys. They are rarer among the 
weeds in farms or in crevices among rocks. 

"I came across O. pusilla repeatedly on the summits of hills in the 
Glavnyi Range at an altitude of 1500 near Podgornyi. I was surprised 
to hear the voice of this typical steppe animal in meadows in high mountains. 
It was once heard in the Saur Mountains (the Zaisan forestry, near the 
village Chegan-Obo) on the southern slope of a valley high in the mountains. 
I did not find it in other parts of the Semipalatinsk District. " 

As mentioned above, O. pusilla leads a very secretive life, but it 
can be easily spotted by its characteristic cry consisting of a rapidly 
repeated "chiv-chiv-chiv". It cries loudest and most frequently at sunset. 
It can be heard, however, also on cloudy days, and especially in the 
morning. It apparently does not leave its burrow at night. The burrows 
are shallow and have several exits. B.A. Kuznetsov did not find "haystacks" 
of this species near Orsk or in the Semipalatinsk District. 

Our knowledge of the reproduction ofO. pusilla is very fragmentary. 
P. Pallas once obtained a gravid female (30 Apr.). She gave birth to 6 young 
which remained blind for 7 days. They opened their eyes on the 8th day, 
their fur began to grow and they began to crawl around the burrow. 



Three females out of 5 caught by B. A. Kuznetsov in the Orsk District 
/ere gravid. The others were already suckling their young. The number 
jf embryos and their size in several specimens were as follows: 

No. 1 1 1 — 6 embryos of 15 mm length 

No. 144- n " " 35 " 

No, 145- 12 " " 10 " " 

These females were caught in late summer: No. Ill on 25 July; No. 144 
and 145 on 29 July. We can therefore state definitely that O. pus ilia , 
has at least two generations in summer and autumn. 

104 Subspecies of Oc hot ona pusilla 

325. Ochotona pusilla pusilla Pallas (1768). European chekushka 

1768(1769), Lepus pusillus. Pallas P. , Descriptio Leporis pusilli. Novi Commentarii Academiae 

Petropolitanae, t, XIII, pp. 531-538. 
1778. Lepus pusillus. Pallas P. , Novae species Quadmpedum e Glirium ordine, pp. 31-45. 
1811. Lepus pusillus. Pallas P. , Zoographia Rosso-Asiatica, pp. 151-152. 

1850. Lagomys pusillus Eversman E. , Estestvemiaya istoriya mlekopitayushchikh Orenburgskogo kraya 
(Natural History of Mammals of the Orenburg Territory), pp. 207-208. 

1851. Lagomys pusi llu s Simashko Yu. , Russkaya Fauna. Mlekopitayushchie (Russian Fauna. Mammals), 
VoL II, pp. 790-791. 

1891. Lagomus pusillus Nehring A. , Die geographische Verbreitung der S'dugethiere in dem 

Tschernosem-Gebiete. — Ztschr. der Gesellsch fiir Erdkunde, XXVI, No. 4, S 323-326. 
1897, Lagomys pusillus Zarudnyi N. A. , Zametki po faune mlekopitayushchikh Orenburgskogo kraya 

(Notes on the Mammal Fauna of the Orenburg Territory). — Materialy k poznaniyu fauny i flory 

Rossiiskoi Imperii, III, p. 360. 
1928, Ochotona p u s i 1 1 a Kuznetsov B, A. , Zamtki po faune mlekopitayushchikh Ural 'skoi gubemii 

(Notes on the Mammal Fauna of the Uralsk Province). — Trudy po lesnomu opytnomu delu, No. 1, 

p. 58; Auct, cit. , Mlekopitayushchie stepnoi polosy Yuzhnogo Urala (Mammals of the Steppes of the 

Southern Urals), — Byulleten' Moskovskogo obshchestva ispytatelei prirody, Vol. XXXVII, No. 3-4, otdel 

Biologiya, pp. 259-262. 1928. 
1932, Ochotona pusilla pusilla Argyropulo A. I. , Materialy po faune gryzunov Srednei Azii (Data 

on the Rodent Fauna of Soviet Central Asia). — Trudy Zoologicheskogo Instituta Akademii Nauk, I. 

pp. 52-55. 

TYPE LOCALITY AND DEPOSITION: Pallas (1768) mentions in his first description the Volga steppes 
(in campis circa Volgam. . . ). The Samara steppes should be considered as terra typica (e. g. the Buzuluk 
District). It is not known whether the holotype has been preserved. It is not in the ZMAN collection. 

MATERIAL EXAMINED: 12 pelts with skulls. 

DIAGNOSIS. Brown color very prominent on back in summer fur; 
deep brown or black hair tips well marked. 

Length of skull less than 40 mm. Skull of old specimens little elevated 
in the nasofrontal region. 

Length of skull 35-38.7 mm (M.36.9); condylobasal length 32.0-36.7 mm 
(M. 34.4); zygomatic width 18.4- 19 mm (M. 18,7); width of skull at level 
of auditory meatus 18,1- 19.8 mm (M. 18.9); length of upper tooth row 6.9- 
7,3 mm (M, 7,1), Body length 153-182 mm (M, 167.2); hind feet 26-27,5 mm 
(M,26,7), 

ADDITIONAL DESCRIPTION, General color of summer fur on back 
deep chestnut-brown with a slight straw tinge. Color of top of head and 
lower part of back intensively brown because of the well marked black- 
brown hair tips. Straw color most marked on flanks. Underparts dirty- 
gray with a whitish tinge; neck and belly (especially the lower part) with a 
light rust-yellow shade. Inner side of ears with deep brown hair; lower 
part of ears with short gray hair. The hair is more intensely brown 

89 



^ towards the ear margins which are whitish. Legs dark brown; hair on tips 
of digits lighter, whitish gray. Claws horn yellow, covered by the long 
hair of the digits. 

GEOGRAPHICAL DISTRIBUTION. From the right bank of the middle 
reaches of the Volga (54°N) to the southern foothills of the Urals (to 
53°30'N), reaching the basin of the river Or' in the southeast. 

326, Ochotona pusilla angustifrons Argyropulo (1932) 
Siberian chekushka 

1884, Lagomys minutus Nikol'skii A, M. , Puteshestvie v Altaiskie gory (Journey to the Altai 

Mountains). — Trudy Sankt-Peterburgskogo obshchestvi estestvoispytatelei, XI, p. 177. 
1900. Ochotona pusi llus Kashchenko N. F. , Opredelitel' mlekopitayushchikh Tomskogo kraya (Key 

to Mammals of the Tomsk Territory), pi. 30. 
1932. Ochotona pusill a Kuznetsov B. A. , Gryzuny Semipalatinskogo okruga (Rodents of the Semi- 

palatinsk District), — Byulleten' Moskovskogo obshchestva ispytatelei prirody, XII (1-2): 114-115. 
1932, Ochotona pusilla angustifrons Argyropulo A. I. , Materialy po faune gryzunov Srednei Azii 

(Data on the Rodent Faima of Soviet Central Asia). — Trudy Zoologicheskogc instituta Akademii Nauk, 

I, pp. 55-57, 

TYPE LOCALITY AND DEPOSITION: Along river Dzhamcha, former Kedei subdistrict, former 
Karkaralinsk County, Kazakhstan, Type No, 17421 in ZMAN collection (pelt and skull), 

MATERIAL EXAMINED: 23 pelts with skulls. 

DIAGNOSIS, Differs from subspecies in its lighter color with more 
prominent straw -grayish tones which create a distinct mottling caused by 
the black-brown hair tips of the back. 

Skull of old specimens more elevated in nasofrontal region than in 
nominate (Figures 51-54, 59-61). 

Length of skull 39.7-43.2 mm (M. 40.9); condylobasal length 37.2- 
40,2 mm (M, 38,2); zygomatic width 18,9-22,3 mm (M, 20,3); width of 
skull at level of auditory meatus 20-22,2 mm (M. 20.8); length of upper 
tooth row 7. 8-8. 8mm (M.8.3). 

Body length 182-196 mm; hind feet 26-30 mm (M. 28 mm), 

SYSTEMATIC NOTES, The eastern subspecies of O , pusilla differs 
markedly from the nominate race. The differences in color are very 
distinct when series are compared. 

A.I, Argyropulo had only 3 specimens of the Siberian subspecies. He 
selected an old specimen as type, the skull of which is larger than all the 
skulls of the large series collected by B. A. Kuznetsov in the Semipalatinsk 
District, Some cranial characters given by A.I. Argyropulo for O. p. 
in gu s t i f r on s , such as the narrow interorbital space, its elevation in 
profile, the large size of the bullae osseae, the slightly longer rostrum etc. 
are found only in the very old skull mentioned above . This skull has several 
individual characters which distinguish it from the other specimens of the 
series. 

GEOGRAPHICAL DISTRIBUTION. Omsk, Altai, Semipalatinsk and 
Zaisan-Tarbagatai districts. 



90 






FIGURES 59-61. Skull ofOchotona pusilla angustifrons Argyrop. , No. 2350, 8 June 1929. 
Semipalatinsk District of Sergiopol [Ayaguz] Region, Znamenka Village, B. A. Kuznetsov 

Drawing by Yu. A. Kostylev 



107 



Family LEPORIDAE, HARES 

1821. Leporidae Gray]., London Med. Repos. , XV, p. 304. 
1857. Leporina Blasius J. , Naturgesch. der Saugethiere Deutschlands, S. 409. 

1897. Lagidae Schultze E. , Helias, XIII, S. 82; Catalogus Mammalium europaeorum, Ztschr. fur 
Naturwissensch. , Bd. 73, 1900, S. 209. 

Rodents of the suborder Duplicidentata, mainly characterized by the 
long ears and elongate hind legs. Tail short and well developed. Nasals 
not widened anteriorly; postorbital processes well developed; lateral 
surface of maxillaries spongiose; skull relatively high, not much flattened. 
Bullae osseae rounded, of moderate size; with solid, bony, not spongiose 
wall. Clavicles small or rudimentary. Pubic symphysis well developed. 



Dental formula: i — ; c — ; p-; m-=(23)*. 

We consider the family Leporidae as consisting of 11 genera and 8 
subgenera. A revision of these genera and subgenera may make a new 
classification possible. 

At present we recognize the following genera and subgenera: 

I Genus Lepus with the subgenera: 

1) Lepus s. str. , 2) Eulagus, 3) Poecilolagus, 4) Boreolagus,** 

5) Microlagus, 6) Macrotolagus 

II Genus S ylvil agus with subgenera 

7) Sylvilaguss. str. , 8) T a p e t i 

III Genus Brachylagus 

IV Genus Romerolagus 
V Genus Oryctolagus 

VI Genus Limnolagus 

VII Genus Pronolagus 

VIII Genus Nesolagus 

IX Genus Caprolagus 

X Genus Allolagus 

XI Genus Pentalagus 

GEOGRAPHICAL DISTRIBUTION. Eurasia, from the Boreal- Arctic 
zone to the Tropics, North America (to Central Mexico), Japan, Ryukyu 
Islands, Sumatra, Africa. O r y c t o 1 a gu s was introduced into Australia, 
where it has multiplied enormously. 

KEY TO GENERA AND SUBGENERA OF HARES (LEPORIDAE) OF THE U. S. S. R. 

1 (2). Ear (measured from occiput) shorter than head Interparietal well developed both in young, adult 
and old specimens. Postorbital processes narrow, thin, swordlike, not elongate triangular. Fossa 
mesopterygoidea markedly narrower than length of bony palate. 

Genus Or yet ol agus Lilljeb. 

2 (1), Ear as long as head or longer. Interparietal absent in adults. Postorbital processes elongate 
triangular. Genera Lepus L. and Allolagus Ogn. 

3 (4). Ear usually as long as head, rarely a little longer. Hind legs relatively short. When the animal 
stands, the hind legs appear only little higher than the forelegs. Mesopterygoid fossa about as wide as the 
length of the bony palate. 

Genus Allolagus Ognev. 

4 (3). Ear always longer than head, sometimes 11/2 times as long. Hind legs relatively long; when 
the animal stands, they appear much higher than the forelegs. 

1 08 Mesopterygoid fossa 1 1/6 to 3 times as wide as the length of the bony palate. 

Genus Lepus L. 

5 (6). Head relatively large. Ears short, 1/10 - 1/8 longer than head. Tail including terminal hair 
much shorter than hind foot. Tail in summer fur whitish gray, gray dorsallyj tail in winter fur white. Winter 
fur of most forms snow white, as opposed to summer fur. Anterior lower premolar slightly inclined backwards. 

Subgenus Lepus L. s. str. 

6 (5). Head relatively small. Ears markedly longer than head, 1 1/5 - 2 3/4 times longer. Tail 
including terminal hair about as long as hind foot without claws, 80-95 % of length of foot. Tail of summer 
and winter fur white with sharply contrasting, black (rarely brown) dorsal side. 



2 

♦ m — (as in OChotonidae) only in P en t al a gus Lyon. 

♦* M. Degerb^l and F. Wimpffen Braestrup (1934) consider the creation of this subgenus as not justified 
(see below). 



92 



Winter fur never completely white, brown tones always persisting on back. Only slight differences 
between summer and winter fur in western, southern and desert forms. 
Anterior lower premolar sharply inclined backwards. 
Subgenus Eulagos Gray. 

Genus Lepus Linnaeus (1758) 

1758. Lepus Linnaeus Carolus, Systema Naturae, Ed. X, p. 57. 

1829. Chionobates Kaup Jakob, Skizzierte Entwicklungs-Gesch. d. Europ. Thierwelt, I, S. 170 (based 

on Lepus variabilis and L. borealis). 
1867. Eul agos Gray J. E. , Annals and Magaz. of Natur. History, Sept. p. 222 (based on Lepus 

m e diterraneus Wagner, and L. judeae Gray). 
1899. Eulepus Acloque Alex., Faune de France, Mammif.^res, p. 52 (based on L. europaeus and 

L. variabilis). 

Ear measured from occiput longer than head, sometimes 1 l/2 times 
as long. 

Tail dorsally black or brown, markedly different from the white ventral 
side (subgenus Eulagus), or tail in summer whitish gray with well 
marked gray color dorsally; tail in winter white (subgenus Lepus ). 

Hind legs long, appearing much higher than forelegs when the animal 
stands , 

Mesopterygoid fossa (l 1/6 to 2 times as wide as length of bony palate. 

Interparietal of adults reduced. 

Postorbital processes elongate triangular. Nasals in their posterior 
third 11/5-11/3 as wide as the interorbital space behind the postorbital 
processes. 

Nasal measured along their oblique outer side shorter, as long as, or 
longer than, frontal suture measured to its anterior point between the 
nasals. 
109 Width of posterior nasal opening 1 l/5-l 1/3 as wide as length of bony 
palate, width of anterior nasal opening twice, sometimes 3 times, the length 
of the bony palate. 

Width of incisive foramina posteriorly 1 I/8 -1 1/4 (Lepus) or 1 l/3- 
11/2, sometimes twice (Eulagus) the length of bony palate. 

Incisive foramina sometimes little, sometimes markedly shorter than 
upper diastema. 

Genotype: Lepus timidus Linnaeus. 

GEOGRAPHICAL DISTRIBUTION. Eurasia, North America to about 
25°N in the south), Africa (in the southern part). 

Subgenus Eulagus* Gray J. E. (1867) Desert and Tolai hare 

Eulagos Gray J. E., Annals and Magaz. of Natur. History, Sept. p. 222. 

Head relatively small. Ear large, 1 1/5 - 2 3/4 as long as head. 

Tail including terininal hair about as long as hind foot without claws. 
Tail 80-95% of length of foot. 

Tail in summer and winter fur white with well-marked, contrasting 
black (rarely brown) dorsal side*'!^. 



* We use the spelling Eul agus (instead of Eul agos) for consistency. 
** Tail completely white in Lepus oiostolus Hodgson (1 have not seen the type specimen), (W. T. 
Blanford, The Fauna of Brit. India, 1887? p. 453). Tail dorsally rust-red in the Indian species 
Lepus ruficaudatus Geoffr. 1 have not seen the Indian species and I cannot therefore decide 
whether it belongs to the group of common hares. 

93 



Teats 6 (in true common hare and Tolai hare) or 8 (in desert hare). 

Winter fur never completely white, brown shades always persisting on 
back. Winter fur very slightly different from summer fur in southern and 
desert forms. 

Nasals as long as, or markedly longer than, frontal suture. 

Incisive foramina 1 1/3 to twice as wide as length of bony palate. 

Anterior lower premolar sharply inclined backwards. 

Type of subgenus: Lepus mediterraneus Wagner, and also L . 
judeae Gray. 

GEOGRAPHICAL DISTRIBUTION. Europe and Asia, in temperate and 
southern zones. South African hares belong to a separate subgenus, 
Pronolagus Lyon; North African hares belong to the genus 
Or y c t o 1 a gu s *. Hares of Mesopotamia, Palestine and Arabia belong to the 
subgenus Eulagus. Hares of the subgenus Eulagus are absent in 



110 




^^ 




^1 


j i t: 

1 ', 




I 1 


1 




I 



FIGURE 62. Measurements of skull of hare 

I — total length of skull; II — zygomatic 
width; III — length of nasals (along suture); 

IV — length of nasals (along outer side); 

V — width at level of auditory meatus; 

VI — lateral length of facial part of skull 
(anteriorly to alveolar point, but not to the 
end of nasals). 




FIGURE 63. Measurements of skull of hare 

I — basal length; II — condylobasal length; 
III — length of bony palate; IV — width of 
incisive foramina; V— length of incisive 
foramina; VI — length of pygomatic arch: 
VII — upper diastema. 



(Species of Lepus also occur in North Africa. ] 



94 




FIGURE 64. Measurements of skull of hare 

I — length of skull; II — condylobasal length; III — length of zygomatic 
arch; IV — length of row of upper molars; V — upper diastema. 

Japan and in North America. The subgenus M a c r ot o 1 a gu s Mearns in 
North America is closely related to the subgenus Eulagus . 
Macrotolagus is distributed in Mexico and in southwestern parts of the 
United States of America. 

Ill KEY TO SPECIES OF THE SUBGENUS EULAGUS OF THE U. S. S. R. 

1 (2). Large, length of body usually more than 570mm; length of hind foot 135 mm; length of skull 
usually more than 90 mm (100 mm and more). 

Fur coarse, guard hair dense, curled on back in several subspecies. Ratio of length of bony palate to 
greatest width of mesopterygoid fossa 55.8% on the average. 

Lepus (Eulagus) europaeus Pall. 

2 (1). Muchsmaller, length of body usually less than 570mm; hind foot less than 135 mm long, length 
of skull usually less than 90 mm. 

Fur smooth, silky; guard hair smooth, delicate, straight. Ratio of length of bony palate to greatest 
width of mesopterygoid fossa 70.5% on the average (Tolai hare) and 75% (desert hare). 

3 (4). Ear (measured from occiput) 1 1/5 to 1 1/3 longer than skull, i. e. , ear relatively short Teats 6. 
Width of both nasals posteriorly usually considerably exceeding length of postorbital process. 
Mandibular condyle strongly developed, moderately inclined backwards. 

L . ( E . ) tolai Pall. 

4 (3). Ear measured from occiput 1 1/4 to 1 3/4 longer than skull. Teats 8. 
Width of both nasals (posteriorly) usually equal to length of postorbital process*. 
Mandibular condyle weak, strongly inclined backwards. 

L. (E.) tibetanus Waterh. 



KEY TO SUBSPECIES OF LEPUS (EULAGUS) EUROPAEUS PALL. IN THE U.S.S. R. 

1 (4). Fur on back wavy, clay-rust-yellow, without dull earth-gray and black shades. 

2 (3). Length of skull usually exceeding 100 mm; length of hind foot without claws exceeding 150 mm, 
usually about 160- 170mm. Color of summer fur on back light clay-rust- yellow. Winter fur with marked 
gray and white tones. 

Lepus (Eulagus) europaeus hybridus Desm. 



This character is not always true; there are exceptions. 



95 



3 (2). Length of skull usually less than 100 mm; length of hind foot less than ISO mm, usually about 
140mm. Clay tones of summer fur on back more intense and darker than in L. europaeus hybridus. 
Winter fur almost the same as summer fur in color, in length and in density of hair; only lower part of back 
is slightly grayer. 

L. europaeus europaeus Pall. 

4(1). Fur on back not wavy, i.e. hair straight. 

5 (6). Fur on back very light in summer, rust-clay colored with a sandy tinge. Color of winter fur 
duller, sometimes more intense than summer fur with a less well marked rust tinge. Fur of specimens from 

112 southern parts of the range of the Caspian common hare is less gray; the gray color is marked only on the 
lower part of the back. White fur is observed only in specimens from the extreme northeastern range. 

L. e. caspicus Ehrenb. 

6 (5). Summer fur on back either dull brownish yellow gray with an earthy tinge, often with marked 
black tone, or more yellowish clay with an earthy tinge. Grayness of wirter fur varying in degree. 

7 (8). Winter fur markedly more white and gray than summer fur. Size large: hind foot 167-185 mm 
long; length of skull of fully grown specimens usually exceeding 100 mm. Weight 6-6.4 kg. 

L. e. tesquorum Ogn. 

8 (7). Winter fur not becoming white, only grayish in posterior part of back. Size and weight much 
smaller than in L. e. tesquorum Ogn. 

9 (12). Reddish clay tones on chest and flanks marked to various extent. Winter fur without vinaceous 
tinge*. 

10 (11). Reddish clay tones on chest and flanks well marked. Length of hind foot 123-160 mm; length 
of skull about 100 mm, sometimes less, 89.3, or sometimes more than 101 mm. Weight about 3.2 kg (2.9- 

3.9 kg). 

L. e. transsylvanicus Matschie. 

11 (10). Reddish clay tones on chest and flanks weakly marked, replaced by clay-yellow- gray tones. 
Length of skull about 100mm (92-105mm); length of hind foot 140-167mm. Weight about 4kg (3.2-4.4kg). 

L. e. caucasicus Ogn. 

12 (9), Reddish clay tones on chest and flanks absent and replaced by dull straw-brown (in stmamer) or 
dull straw-gray (in winter). Winter fur with characteristic vinaceous tinge. 

Length of hind foot 144-150mm; length of skull usually less than 100mm (91-103 mm). 

L, e. cyrensis Satun. 

157. Lepus (Eulagus) europaeus Pall. (1778) 

1778. Lepus (europaeus) Pallas P. , Novae Species e Glirium ordine, p. 30 (short diagnosis: "Apice 

aurium capite longiorum, caudaque supra atra", without indication of range) 
1811. Lepus timidus Pallas?., Zoographi a Rosso- Asiatic a, p. 148 (Polonia, Lithuania). 

NAMES: Common or brown hare (English); Lampe, Feldhase (German); Le lievre commun (French). 

LOCAL NAMES: rusak, seryak (latter apparently designating steppe rusak, Russian); tumak (hybrid 
between common hare and white hare); according to others, e. g. , M. N. Bogdanov, tumak = steppe rusak; 
zaets sirii, or zvychainyi (Ukrainian); Kuyan (Bashkir); koch' (Komi-Zyryan); toshan (Armenian); urcheli 
(Georgian); tkhakumakikh (Kabardian); kuyan (Balkar); tulai (Kalmyk). 

PRINCIPAL nCURES: 1) Schreber I. , Saugethiere, Erlangen, IV, 1792, Taf. CCXXXIII (Le pus 
timidus = e urop aeus); Taf. CCXXXVC. (Lepus variabilis Pall, hybridus - tumak); 2) Bingley 
W. , Memoirs of British Quadrupeds, 1809, p. 290 (color pi. ) ; 3) Simashko, Yu. , Russkaya fauna. 
Mlekopitayushchie (Russian Fauna. Mammals), II, 1851, pp. 782-785, pi. 62, fig. 2 (color pL of winter 
fur, named Lepus a quilonius); pi. 60, fig. 1; pi. 61, fig. 4 (color pi. and drawing of skull, named 
Lepus europaeus); 4) Middendorf A. Th. , Ueber die als Bastarde angesprochenen Mittelformen zw. 



* Vinaceous buff-avellaneous (Ridgway, 1912). 



96 



L. europaeus und L. variabilis, Bullet, phys. -mathem. , IX, 14, 1851, S. 29 (drawing of skull, 
doisal); 5) Blasius I. , Naturgesch, d. Saugethiere Deutschlands, 1857, S. 412-419, Fig. 226; S. 409 
(entire animal); S. 410, Fig. 227, (skull); S. 412, Fig. 228 (teeth); 6) Meves W. , Holmgren A. E. , 
J 23 Atlas ofver Scandinavian Daggdjur, Stockh. , 1873, pi. XV, fig. 8-9 (entire animal in color); 7) periodical 
"Okhota" No. 6, 1877 (plate of biological scene); 8) Mojsisovics A. , Ueber accessorische Fortsatze am 
Schadel der Leporiden, Sitzb. d. Kaiserl. Akademie d. Wissensch. , math. Class, LXXVl, Bd. I, 1877, 
Taf. Fig. I (skull ventral): 9) Giebel C. G. , Charakter. der Hasenschadel, Ztschr. fur die ges. Naturwissensch., 
LIIl, 1880, Taf. VIIl-X, Fig. 3 (skull lateral and ventral, teeth of Lepus caspius); 10) Fokht K., 
Mlekopitayushchie (Mammals), Sankt-Peterburg, 1885, p. 402 (drawing of entire animal); 11) Flower 
W. H. , Lydekker R. , Anintroduct. to the Study of Mammals, 1891, p. 492, fig. 216 (skull); fig. 217 (entire 
animal); 12) Lydekker R. , A Hand-Book to the British Mammalia, London 1895, pi. XXVII (entire animal, 
color plate); 13) Brehm A. E., Zhizn' zhivotnykh (Animal Life), II, 1893, p. 656 (half tone plate of entire 
animal); 14) Menzbier M. A. , Iz zhizni zverei i ptits (On Life of Animals and Birds), 1887, p. 120 (plate, 
biological scene); 15) Forsyth Major, C.I. On fossil and recent Lagomorpha, Transactions of the Linnean 
Society of London, Vol. VII, Zoology, 1896-1900, pL 36, fig. 25 (teeth); 16) Gaake, V. , Zhivotnyi mir (Animal 
World), Vol. I, 1901, plate 19 (in color); 17) Lyon M. W. , Classification of the hares and their allies, 
Smithson. Miscellaneous Collections, Vol.1, prt. 3-4, 1903(1904), pL LXXIV, fig. 2- 2a (skull ventral and 
dorsal); pi. LXXV, fig. 3 (skull lateral); pi. LXXX and LXXXI, fig. 2 (skull dorsal and ventral); 18) Millais 

I. G. , The Mammals of Great Britain, Vol. Ill, 1906, pi. 49 (color. ), pL , p. 8 (biological scene); pL , p. 16 
(photograph); 19) Lonnberg E. , On Hybrid Hares between Le pus timidus and L. europaeus from 
South Sweden, Proceed. Zoolog. Society London, 1905, pp. 278-287, fig. 53 (teeth); fig. 54 (A. C. ) (part of 
skull); 20)SchaffE. , Jagdtierkunde, 1907, S. 164, fig. 84 (skull lateral); S. 164, fig. 85 (mandibula); 

fig. 86-87 (skull ventral and dorsal); S. 167, fig. 90 (bones of extremities) ; S. 170, fig. 91 (skull with ab- 
normal development of lower incisors); S. 18, fig. 92 (haie with five legs); 21) Hilzheimer M. , Die 
Hasenarten Europas, Jahresbericht d. Vereins fiii Vaterl. Naturk. in Wurtenb. , 1908. S. 393-394, Taf. V, 
Fig. 1, a-c (skull dorsal, teeth— Lepus europaeus typicus): Taf.V, Fig. 4 (skull dorsal — 
L. m, caspius); Taf. V, Fig. 2a-b (skull and mandibula— L. m. aquilonius); 22) Meerwarth 
H. u. SoffelK. , Lebensbilder aus der Tierwelt, II, 1910, S. 321, 322-326, 328-329, 331-332, 335, 337, 
340, 342-345 (photograph of animals, nests, traces etc. ); 23) S chaff E. , Die wildlebenden Saugetiere 
Deutschlands, 1911, S. 117-120, Fig. 34; S. 116 (skull ventral)-; 24) Barrett-Hamilton G. , A History of 
British Mammals, fasc. XI, 1912, p. 230, fig. 44 (I) (skull) ; fig. 45 (I) (skull) ; p. 233, fig. 46 (teeth) ; fig. 
pi, XVI (head); p. 262, fig. 47 (skull lateral); fasc. XII, 1912, pi. XIX (3) (feet); pL XX (7) (feet); fasc. XIV 
1913, pi. XXIII (photograph of hare); 25) Ognev, S. I. , Mlekopitayushchie Moskovskoi gubemii (Mammals 
of the Moscow Province), 1913, pp. 283-298, pi. Ill, figs. 51-52, 54 (photograph of skulls); pi. IV, fig. 55 
(skull); fig. 56 (skull of tumak); fig. 61 (skull of young common hare); 26) Brehm 's Tierleben, Saugetiere, 

II, 1914, Farb. Taf. 82; 27) Ognev S. I. , Mlekopitayushchie Tavricheskoi gubemii (Mammals of the Tauric 
Province), parti, Gryzuny (Rodents), 1916, p. 60, pi. IV, fig. 2 (skull - L. europaeus aquilonius); 

pi. Ill, fig. 1 (skull of L. e. transsyl v anicus); plate IV, fig. 1 (skull - L. e . caspius); 28) Filipchenko, 
Yu. A, ,Izmenchivost' i nasledstvennost' cherepa u mlekopitayushchikh (Variability and Heredity in the 
MammalSkull), Russkii Arkhiv Anatomii, Gistologii i Embriologii, Vol.1, No. 2, Petrograd, 1916, plate 
XVI, fig. 2,4 (photograph of skulls -L. e. aquilonius); 29) Thorbom A., British Mammals, Vol.11, II, 
1921, pi. 33 (in color); 30) Ognev S. I. Fauna pozvonochnykh Voronezhskoi gubemii (Vertebrates of the 
Voronezh Province), 1923, plate 1, fig. 1 (photograph of young L . e . tesquorum); 31) Kaverznev V. N. , 
O zaitse i ego dobyvanii (The Hare and Its Capture), Moskva, 1928, p. 6 (traces); p. 7 (structure of tail, 
excrements); 32) Bobrinskii N, A. , Opredelitel' okhotnich'ikh i promyslovykh zverei (Key to Game and Fur 
Animals), 1928, p. 41, fig. 28 (drawing of tail); 33) Barabash-Nikiforov II., Narysy favny stepovoy 
Naddnipryanshchyny (Description of the Steppe Fauna of the Dnieper Region), 1928, pp. 105-109, fig. 56 
(traces); fig. 57 (lair in the snow)j 34) Shillinger F. F. , Zaitsy rusak, belyak i dikii krolik (Common and 
Blue Hare, and Wild Rabbit), Khar'kov, 1929, p. 9, fig. 2 (hare leaving den); fig. 3 (biological scene); 
fig. 4 (traces) ; fig. 14 (hare in a field); 35) Geptner V. G. , Zaitsy (Hares), Moskva, 1933, p. 15, fig. 2 
(entire animal; excellent drawing by Komarov); 36) Vinogradov P. S. , Mlekopitayushchie SSSR Gryzuny 
(Mammals of the U. S. S. R. Rodents), Leningrad, 1933, plate VIII, fig. XVI (photograph of young hare); 37) 
Bobrinskii N. A. , Opredelitel' okhotnich'ikh i promyslovykh zverei SSSR (Key to Game and Fur Animals 
of the U. S. S. R. ), Moskva, 1935, p. 50, fig. 39 (tail); fig. 40 (figure on right side -entire animal); 38) 
Manteifel' P. A., ZaitsySSSR (Hares of the U. S.S. R. ), in Sbomik "Biologiya zaitsev i belok", Moskva, 
1935, p. 18, fig. 2 (copulation); 39) Gureev A. A. , Kraniologicheskie priznaki zaitsev belyaka i rusaka 
(Cranial Characters of Blue and Common Hare), Trudy Zoologicheskogo Instituta Akademii Nauk, III, 1936, 
pp. 356, 357, 362, figs. 2, 4, 6, 8 (skull and details). 
114 MATERIAL EXAMINED: 249 pelts and 291 skulls. 



97 



114 DIAGNOSIS. Head relatively small. Ear measured from occiput 11/5 
to 1 1/2 times longer than skull. 

Tail including terminal hair about as long as hind foot without claws. 

Tail in summer and winter fur white with dorsal side black, sharply 
contrasting. 

Winter fur never completely white; brown tones always persisting on 
back. Winter fur differs slightly from summer fur in southern and desert 
forms. 

Hair coarse; guard hair dense, curled on back in several subspecies. 

Teats: p: 1-1; a: 2-2; total: 6. 

Posterior third of incisive foramina markedly widened and slightly 
curved. 

Ratio of length of bony palate (measured laterally, from the medium 
anterior process) to greatest width of incisive foramina (in their posterior 
third) 56%. 

Mandibular condyle weak, strongly inclined backwards. 

Posterior ventral plate of mandible from angular process to notch at 
the anterior horizontal mandibular process rather weakly developed. 
Distance from angular process to notch at anterior horizontal mandibular 
process about equal to, sometimes smaller than, distance between anterior 
alveolus of first molar and tip of lower incisor. 

Bullae osseae small and inflated; slightly protruding in profile and 
rounded anteriorly. 

Length of ear along concavity 100- 120 mm; length of ear (measured 
from occiput) 120-171 mm; length of hind foot 124-167 mm. 

ADDITIONAL DESCRIPTION. Skull relatively elongate, rostrum long 
and slender. The specific characters of the skull of the common hare 
are very distinct when compared with the skull of the blue hare. 

This has been carefully studied by various authors^'-, 

115 Compared with the skull ofL. timidus, the skull of the common 
hare is very elongate, narrow, and the nasals are long. Interorbital 
space of the common hare generally narrower than that ofL. timidus. 
Zygomatic arch narrow; in lateral view does not rise so steeply posteriorly 
and does not delimit the lower side of the orbit asinL. timidus. Orbit 
ofL. timidus markedly different from that of common hare. Incisive 
foramina of common hare generally longer than those ofL. timidus. 

Yu.A. Filipchenko (1916) gives the following variations of this character: 
in L, timidus 22-27,5 mm (M. 24.30 mm); in the common hare 24-29 mm 
(M. 26.19mm). 

116 The best distinguishing character is the ratio of the length of the bony 
palate to the width of the incisive foramina and the mesopterygoid fossa 
(see above). 



A. Th. Middendorf, Ueber die als Bastarde angesproch. Mittelformen z. L. europaeus undL. 
variabilis, Bull. phys. -mathem. , IX, No. 14, 1851; I. Blasius, Naturgesch. d. Saugethiere 
Deutschlands, 1857, S. 412-414, 420-421; C. G. Giebel, Charakteristik der Hasenschadel, Ztschr. fur 
die ges. Naturw. , Bd. UII, 1880, S. 318-340; M. Hilzheimer, Die Hasenarten Europas, Jahresb. d. 
Vereins f. vaterl. Naturkunde in Wiirtt. , 1908. S. 384-419; G. S. Miller, Catalogue of the Mammals of 
Western Europe, London, 1912, pp. 498-502, 522-526; S.I. Ognev, Mlekopitayushchie Moskovskoi 
gubemii (Mammals of the Moscow Province), Moskva, 1913, pp. 283-286; Yu. A. Filipchenko, Izmenchi- 
vost' i nasledstvennost' cherepa u mlekopitayushchikh (Variability and Heredity in the Mammalian Skull), 
Russkii arkhiv anatomii, gistologii i embriologii, 1 (2): 329-344. 1916. 



98 



115 





FIGURE 65, Skull of Le pus europaeus caspicus 
Ehrenb., No. M, 3021 <f 27Jan. 1913, Khokhlatskoe, 
Astrakhan Counties, N. L. Chugunov, S.I. Ognev's 
collection 

Drawing by Yu. A. Kostylev 



FIGURE 66. Skull of Le pus europaeus 
caspicus Ehrenb., No. M. 3021 



Comparison of the mandibula of the common hare and ofL. timidus 
116 shows very clear differences. As already stated above, the posteroventral 
' plate of the mandibula from the angular process to the notch at the beginning 

of the horizontal process of the mandibula is weakly developed. In L. 
timidus this plate is strong and broad. The distance from the angular 
process to the notch at the beginning of the horizontal process of the 
mandibula is about equal, sometimes smaller than distance from the 
anterior alveolus of the first molar to the tip of the lower incisor. The 
first measurement is much larger than the second in the blue hare (see 
description ofL. timidus.) 



99 




FIGURE 67. Skull of Lepus europaeus caspicus Ehrenb. , No. M.302I 




FIGURE 68. Mandibula of Le pus europaeus caspicus Ehrenb., No. M.3021 



117 



Yu. A. Filipchenko (1916) carefully analyzed 80 different cranial 
characters. He concluded that only 8 measurements are really characteristic 
for the skull ofL. europaeus and L. timidus.='= Yu . A . 
Filipchenko gives these 8 measurements in form of indices. 



Indices: 

1) Lateral length of brain case (from inion (occiput) to posterior margin 
of orbit and base of upper margin of zygomatic process of temporal bone) 

Lateral length of facial part of skull (from alveolar point to middle of 
anterior or lower corner of orbit) 

2) Length of bony palate 
Width of incisive foramen 
3)Length of tooth row 
Length of diastema 

4) Length of zygomatic arch 
Oblique length of nasal bone 



Lepus timidus Lepus europaeus 
> 80 < 80 



> 65 


<65 


> 65 


<65 


> 95 


<95 



* Yu. A. Filipchenko indicates the most salient characters. However, these 8 characters do not exclude the 
basic differences between these two species (see description and diagnosis of the common hare). 



100 



All these differences are relative, according to Filipchenko; a series of 
indices of one species may exceed the maximum or minimum of the 
measurements of another species, to a certain extent. Filipchenko 
analyzes the cranial differences by statistical methods. He compares 
variation ranges, taking into account the means of each series and the mean 
errors. The latter depend on the quadratic deviation of the series (o) and 
on the number of terms in the series (n). 

In order to calculate the quadratic deviation (provided the mean {M) is 
known), Filipchenko used the formula; 



/?S2. 



in which a - deviation from M, p- number of terms in each class, S — 
summation symbol. After calculating the quadratic deviation, the mean 
error [m) can be determined because 

a 
yn 

If the mean and mean errors of two series are known, it can be 
determined whether there are true differences between the two. The 
difference of the means of one series from the means of the other are 
118 determined, i. e. , the difference M^ — M^. The mean error of this difference 
is then determined, i.e., the square root of the sum of quadratic errors of 
both means, which is equal to the sum of the squares of the errors of both 



means, i.e., ij//n,2 -|- /Oj^-We thus obtain the formula: 



{M,-M,)±i/ml + ml 



Comparison of the two sides of this formula shows whether two series 
differ. 

If the difference between the means is much greater than its mean error, 
the ranges of variation differ. If, on the other hand, the difference between 
the means approximates the average error, there is no difference between 
the series, and the characters recorded are only apparent. 

Filipchenko compares 8 cranial characters by this method. The following 
table shows the results of this analysis. 





Lepus timidus 


Lepus europaeus 






(W, — Af,) * Km,'+m 


2 






m 




m 






Lateral length of brain 














case 


1.1086 


0.24 


1.4130 


0.26 


2.04 ±0. 35 




Lateral length of facial 














part of skull 


1.7500 


0.38 


2.1775 


0.40 


1.77 ±0.55 




Length of bony palate . . 


0.5117 


0.11 


0.6365 


0.12 


1.49±0.15 




Width of incisive foramen 


0.6362 


0.14 


0.8094 


0.15 


1.50±0. 20 




Length of upper tooth row- 


0.9338 


0.20 


0.7728 


0.14 


2. 1 1 ± 0. 24 




Length of upper diastema 


0.9698 


0.21 


1.4172 


0.26 


1.37±0.33 




Oblique length of nasal 














bones 


1.9730 


0.43 


3.1059 


0.57 


3.17±0. 71 




Length of zygomatic arch 


1.7087 


0.37 


1.8874 


0,35 


1.78±0. 51 





101 



These figures prove that the difference between the means of both 
species is 3 to 10 times as large as the mean error. 

There are thus clear cranial differences between the two species. The 
overlapping of the measurements and the relative characters of these 
differences are of no importance. 

In addition, it may be remarked that the upper tooth row of L . 
europaeus is on the average shorter than that ofL. timidus. 

Filipchenko (1916) gives the following measurements of the upper tooth 
row: 

Lepus timidus Lepus europaeus 

17.5 - 21 (M. = 19.28mm) 15.5 - 18.5 (M. = 17.17 mm) 

The color of the common hare varies considerably in different localities. 
The texture of the fur varies with the season. This is especially marked in 
the common hare from the northern and eastern parts of its range. 

A description follows of the color of the summer fur of a central Russian 
common hare, and its geographical variations. 

Anterior part of muzzle, head, forehead and occiput covered with dense, 
short fur which is generally yellowish clay in color with a more or less 
marked grayish brown tinge. Bases of hairs in this region very dark brown; 
subapical parts straw-clay-yellow (intermediate between pinkish cinnamon 
and light ochraceous buff); hair tips blackish. There is also some 
completely black guard hair. 

Color of cheeks and sides of anterior part of muzzle more gray and 
whitish due to some hairs with a broad straw- whitish subapical band. Eyes 
ringed with light whitish ocher hair. 

Middle part of broad inner margin of ear with relatively short black- 
brown hair mixed with hair with rust-straw subapical parts. A sharply 
delinated white stripe on the inner side of the ear, along the inner edge of 
the ear separated from the dark brown hair by an indistinct straw-yellow 
line. 

Inner surface of ears with whitish gray hair. Margin of outer sides with 
an indistinct, narrow brownish gray stripe; base of outer side of ears with 
a narrow white stripe; apex of inner side of ears of dull rust-straw; tip 
of ear with a black-brown edge. Posterior side of ears covered with dirty- 
grayish white hair, with broad (to 55 mm) black-brown tips; clay-yellowish 
fur at the base of posterior side of the ears. The region posterior to the 
ears, upper part of back and flanks covered with long reddish rust-straw 
hairs (intermediate between cinnamon, sayal-brown, Ridgway, 1912, 
pi. XXIX, and wood-brown, pi, XL). 

Upper part of back with wavy, curled hair of a clay-rust-straw tone with 
a black-brown tinge. The hair of this region consists of different types: 
a. guard hair with a gray-black base, broad yellowish rust-straw subapical 
band and black-brown apex; b. longer and denser guard hair with wider 
subapical bands and narrow apices in various parts of the back. There is 
also a predominant number of completely black guard hairs which make the 
fur more flexible and less silky. 

Underfur woolly, dense, light pinkish straw (intermediate between 
pinkish cinnamon and light ochraceous buff). Bases of hairs whitish, their 
tips brownish gray. 



102 



Posterior part of back duller and more grayish than its middle part. 
A clay-rust-reddish stripe runs from flanks to groin (anterior part of hind 
legs), and a similar stripe runs across the chest. This color is 
i particularly marked on the chest; its general tone intermediate between 
pinkish cinnamon and vinaceous cinnamon. Throat white. 

Color of anterior side of foreleg the same reddish rust-clay tone with 
gray streaks caused by gray-black hair. Color of hind legs more yellow, 
inner side whitish. Soles a dirty yellowish gray as the hair is always 
stained by soil. 

Belly and groin white. 

Ventral and lateral surfaces of tail with fluffy white hair; dorsal surface 
black or blackish brown. , 

The distribution of the vibrissae is generally constant, but deviations 
sometimes occur. The longest vibrissae are on the lateral parts of the 
muzzle, between eye and tip of nose. Each group of vibrissae consists 
of at least 28. The color of the longest vibrissae varies, their base is 
usually black and their tip broad and white; they may reach a length of 
130 mm. 

There is one, rarely two, long vibrissae behind the eye, in the lower 
part of the cheek. A group of four to five vibrissae is found above the eye. 
Finally, I found several specimens with two white vibrissae on the chin, 
close to the lower lip. 

The color of the summer fur varies strongly in different localities. 

Specimens from the Chernozem steppes of European Russia (L . e . 
tesquorum) have a duller earth color and less yellowish clay tones; 
those from the Astrakhan semidesert (L. e. caspicus) are a relatively 
light yellowish clay. Specimens from the Kalmyk area differ from 
Astrakhan specimens by their dirtier ocher-clay-yellowish color mixed 
with more gray-yellow tones, etc. 

The winter fur of central Russian hares differs from the summer fur in 
being markedly grayer on the upper side of the body, which is caused by 
long hair with white bases, narrow black-gray subapical bands and very 
broad white tips, especially on the flanks and the lower part of the back. 
These hairs may be 65 mm long. Flanks and lower part of back whitish 
gray with a slight straw tinge. There are many white hairs among the 
ordinary guard hair; and also many hairs with whitish bases, dull straw 
or rust subapical parts and short blackish gray tips. Black-gray hairs 
are rare here. Hairs with rust -clay bands, and pure black guard hairs 
persist in the middle of the back; however, white hair also occurs here, 
giving the fur a more grayish color. Cheeks and area around the eyes 
white; rust and clay colors persist on the forehead and occiput. Only 
small parts in the middle of the chest and on the flanks pale, reddish rust- 
clay; broad white hair tips give a lighter color to the fur in these parts. 
The anterior sides of the forelegs usually retain their typical reddish clay- 
rust color, hind legs usually white. Tail always black dorsally. 

The color of the winter fur varies strongly. As a rule, it becomes paler 
and whiter from west to east. For example, Ryazan, Tambov and Saratov 
hares are much paler and whiter than Moscow hares. Hares from northern 
Kazakhstan (e.g., from the Kustanai District) are very pale and whitish. 
Southern hares (from the Crimea and Transcaucasia), and also hares from 
Kiev and Podolsk become only little paler in winter. A marked gray tone 
appears only on the lower section of the back, and on the cheeks. 



103 



SYSTEMATIC NOTES. There are 7 subspecies of the common hare m the 
the U.S.S.R. 

The following races occur in western Europe; 1) L. e. europaeus 
Pall. , from Denmark and the Atlantic Coast through central and western 
Poland, Germany to central France. This form was introduced recently 
into southern Sweden; 2) L.e.occidentalis de Winton, England and 
Scotland to the Orkney Islands in the north; 3) L.e. pyrenaicus 
Hilzheimer, the Iberian Peninsula, southeastern France; 4) L . e . 
meridiei Hilzheimer, southern and central France, North Italy, Corfu; 
5) L.e.corsicanusde Winton (Corsica, Sicily, Italy; 6) Le pu s e . 
t r a n s s y 1 V an ic u s Matschie - Roumania, the Balkans, Peloponnesus--. 

The systematic relation of the common hares to L.tibetanus, and 
to the small southern European species (L . mediterraneus Wagner.) 
will be discussed in the description ofL. tibetanus. 

GEOGRAPHICAL DISTRIBUTION. The range of the common hare has 
changed markedly during the last century. This species has continuously 
penetrated new localities, mainly in an eastern and northern direction. 

A delineation of the northern boundary of the range ofL. europaeus 
follows. The common hare apparently occurs only rarely in southern 
Finland, south of Savo, Istava (vide J. Valikangas, 1909, D.J. Waden, 
1910). It is rare north of Leningrad. The boundary of the range runs 
to the northeastern coast of Lake Ladoga in the east, turning north of 
Petrozavodsk to the Medvezh'ya Mountain at the northern corner of Lake 
Onega. The boundary extends to Chekuev on the Onega river (its most 
northern point) in the east, reaching Penshinsk in the south, then turning 
south, passing near Plesetskaya*''^ in the east, north of the town of 
Shenkursk on the Vaga river (a tributary of the Northern Dvina). 
122 The boundary turns south from the town of Shenkursk, not reaching 

Veliki Ustyug. It crosses the Sukhona river, including the Nikolsk District, 
It crosses the upper reaches of the Mologa and Vyatka rivers. The 
boundary reaches the Kosa river (a tributary of the Kama river) in the 
Komi-Permyak Territory in the northeast. According to A.G. Tomilin 
(in litt. 1935), the boundary reaches 59°55'N here. It then turns to 
Perm, touches Kungur in the north, makes a large curve near the Ural 
Mountains, south to about 40 km north of Zalair, turns north again at 
Nizhne-Isetskoe Lake (in Central Trans-Ural). It thus makes a curve to 
Shadrinsk, Kurgan and runs east to the Tara river (in the Tobolsk Territory; 
according to B. F. Belyshev, in litt. 1934). This locality is the limit of the 
recent northeastern range of the common hare in West Siberia. The further 
course of the southwestern boundary of the range is not exactly known. 
O. 1. europaeus occurs in the Omsk, Petropavlovsk, Kokchetav, 
Atbasar regions, and the boundary then turns west to Turgai. Its course 
to the northern coast of the Aral Sea is unknown; it then runs east into 
Turkestan to the village Kara-Uzyak (west of Kzyl-Orda-Perovsk). This 
point is the southeastern limit of the range ofL. europaeus (see Map 
II). 

The southern boundary of the range of the common hare runs west of the 
Aral Sea through the Mugodzhary Hills, although not exactly known and then 



♦ G. S. Miller (1912) records L. e. hybridus Desm. from eastern Germany, but with doubt. This race 
does not occur there at all, being replaced by the nominate race. 
** [Now Plesetsk. ] 



104 



south of Temir. It can be traced to the estuary of the Ural and Volga rivers. 
It is distributed in the Kalmyk Region, along the western coast of the Caspian 
Sea to Transcaucasia and Azerbaidzhan. According to Blanford (1876) 
the common hare is numerous between Teheran and Isfahan, where the 
southern boundary of the range in Iran apparently passes. The boundary 
turns northwest and includes Asia Minor (M. Kollmann, 1918). The southern 
boundary in this country has not been determined. Further west the 
boundary passes the Balkan Peninsula (including Greece), the Apennine 
Peninsula (and Sicily, Corsica and Corfu), and finally, it includes 
southern France. A typical hare considered as a separate species, 
Lepus granatensis Rosenhauer, occurs on the Iberian Peninsula and 
in the Balearic Islands. This species has three subspecies on the continent. 
Small hares of the type of the common hare from Crete are also considered 
as a separate species (Lepus creticus Barrett-Hamilton); specimens 
from Sardinia are considered as the species L. mediterraneus 
Wagner. Hares from Parnassus (Liakoura) are considered as a separate 
species, L. parnassius Miller. The exact systematic relations of 
these species to L . europaeus Pall, will become clear only after the 
study of a large amount of material. 

A description of the distribution of the U. S. S. R. follows. In the 
northwest L. europaeus is common in some localities in the Leningrad 
Region. V. A. Bianki (1909) wrote that it was very common in fields and 
meadows near the coast of the former Peterhof [Petrodvorets] County. It 
is common near Novaya Ladoga and Narva. According to A. F. Kessler 
(1868) the former Peterburg Province was the northern boundary of the 
common hare's range in the late 1860's. He remarks: "the common hare 
never enters the former Olonets Province. " In their monograph on the 
common hare, S. S. Folitarek and A. A. Maksimov (1936) give data supplied 
by Sudnik, the Director of the Applied Zoology Section of the Karelian 
Scientific Research Institute. Sudnik wrote that the common hare appeared 
in Karelia between 1880 and 1890, gradually moving north. The 
northward migration of the common hare was apparently especially marked 
near the western coast of Lake Onega, a more densely settled region. 
124 According to Sudnik, the common hare occurs in the following southern 

districts of Karelia: Medvezhegorsk, Kondopoga, Pryazha, Petrozavodsk, 
Sheltozersk, Olonets, Pudozhsk and Onega. It has not yet spread north 
of the Medvezhegorsk District. It is rare in the places mentioned, 
constituting about 1 % of the population ofL. timidus. It becomes 
gradually more numerous further south towards Lodeinoe Pole constituting 2 or 
■ 3 %of the populationofL . timidus . Accordingto S. S. Folitarek and A. A. 

Maksimov (1936) the common hare occurs in small numbers in the 
basin of the Onega river east of the Priozersk District (Kenozero) . It 
appeared here in 1916 or 1917, and had not been recorded previously. 
According to A.M. Korsakov (in litt, 1936) it is found along the Onega 
river to Chekuev and farther east in the Plesetsk District. The number of 
common hares caught in 1935 to 1936 constituted 0.68% of the number of 
L. t imidus caught (94 common hares and 13,557 of L . timidus). 
It is very rare further east, in the Shenkursk District (on the Northern 
Dvina). Accordingto S.S. Folitarek and A. A. Maksimov, only 10 common 
hares were found among 1706 blue hare»examined in this region. 

South of this region, in the Vologda and Novgorod territories, the common 
hare is rare and distributed sporadically, always less numerous than the 



105 



blue hare. However, it is not rare near Cherepovets and Vologda, where 
it occurs in all isolated, more or less open localities. It is rare in the 
Sharya Region. According to A. A. Maksimov (1936) it is more common 
along the Vetluga river. It is, however, completely absent in forests only 
a few kilometers from the river. S. V. Lobachev (1930) reports that in the 
Vyatka ]Kirov] Territory, the common hare keeps to the central, unforested 
parts of the former Vyatka Province. It is most numerous in the southern 
part of the Nizevskoe and the northern Porez forests. It occurs throughout 
the former Nolinsk and Vyatka Counties, in the central parts of the Kotelnich 
County and in the northern parts of the former Yaransk and Urzhum counties. 
According to A. V. Shestakov (1926) it was less common than the blue hare 
around 1860 in the Yaroslavl Territory. It now outnumbers the blue hare on 
the right bank of the Volga. 

According to recent data by A. G. Tomilin (1934) the range of the common 
hare in the Komi-Permyak Territory reaches 59°55'N (near Yukseevo- 
Chazevo, in the Kosa District). It occurs only sporadically in the Kochevo 
District, on a few scattered fields. It disappears completely in the 
northern parts of this district which are covered by continuous forest. It 
becomes much more common in the Yurlinskii District. A. G. Tomilin 
adds that the common hare is in general much less common than the blue 
hare in the region between Vyatka and Kama (in the northern part of the 
former Vyatka Province and the northwestern part of the former Perm 
Province). It constitutes 10-12.5% of the population of the blue hare. 
There is no doubt that Lepus europaeus is spreading northward also 
in this region. L. P. Sabaneev (1874) remarked that the common hare 
was very rare in the former Okha and Osa counties of the Perm Province. 
A. G. Tomilin (1934) recently wrote that the common hare is not rare in 
this region, since as many as 1000 specimens per year were caught in the 
small area mentioned above. 

B.S. Lukash (1929) gives some data on the upper reaches of the Kama 
Region. He says: "The common hare was found in the extreme northeast, 
in Verkhneuralsk, in 1926 only in the Gordino District (near the sources of 
the Kama, north of the Vyatka-Perm Railroad). It constituted 10% of the 
total catch, 90% of which were blue hares. Only the blue hare is found 
in the Kai and Kirs districts, and in the Berezov Clearings. Members of 
the Omutinskoe Hunting Association shot only one common hare in 1928- 
1929 and 1930 blue hares. The zones adjoining the Omutinskoe and 
Sloboda Hunting Regions yielded 20% common hares and 80% blue hares in 
1925-1926, accordingto data of the Glazov Agency of the State Trade Company." 

The common hare does not occur near Cherdyn Kama, which lies 
halfway between Perm and Dedyukhino on the eastern boundary of the range 
of the common hare. According to S.S. Folitarek and A. A. Maksimov 
(1936) the common hare occurs south of Chermosskoe, reaching the 
Kama river. It occurs in small numbers on the right bank, in the Ust 
Garevskii, Selninskii and Shemeshevskii Rural Soviets. It constitutes less 
than 5% of the blue hare population. There are data which indicate that 
the common hare appeared in these places only in 1923-1924. 

Distribution in the southern regions of the lower part of the Kama Valley 
and the central Volga Region. M.N. Bogdanov (1872) gives the following 
data on the distribution of the common hare in this area: "The common 
hare is constantly becoming more numerous in fields of the pine -forest 
region of the tertiary basin. It is not rare any more in fields of the clay 



106 



126 



belts of the Kazan province. This is especially true south of the Volga and 
Kama rivers. The northward migration of the common hare from the 
Kazan Province had already been observed by Eversmann. In a village 
not far from Kazan, a hunter confirmed the above and told me that the 
first common hare was killed in the winter near their village 40 years ago. 
It drew the general attention of the hunters, as they had never seen such 
a hare. It is now distributed throughout the eastern part of the Kazan 
Province to the Ashita river. It also lives in small numbers in the 
southern counties of the Vyatka Province. It is found in fields and 
in the spruce zone in the southwestern corner of the Kazan County. " 

The above shows how far north the common hare spread from its range 
in the former Kazan Province in the 1870's. 

The common hare is widely distributed along the Belaya river although 
in the region of Izmailovo and Dyurtyuli it is much less common than the 
blue hare. 

The common hare has actively spread to the mining regions of the Ural 
in recent years. It reached the Nyazepetrovsk District (in 1927) apparently 
from the west. It now occurs together with L. timidus at a rate of 8 blue 
hares to 2 common hares (out of 10) It also occurs in the N. Sergievskii 
ZavodRegion. Out of 8000-10,000 hares shot here yearly, only 100 
are common hares (S.S. Folitarek and A. A. Maksimov, 1936). 

Thus the boundary of the range of the common hare in the regions near 
the Ural Mountains is not a straight line along the mountains, but an irregular 
curved line the course of which is not very clear. It may be sufficient at 
present to state that the common hare is gradually conquering new regions 
near foothills and uplands . 

According to N. A. Zarudnyi (1897), the common hare abounds in the 
southern part of the Orenburg Territory and in the steppes in the middle 
reaches of the Ural river, between Orsk and Orenburg. It is very numerous 
along the Obshchii Syrt, in the upper reaches of the Salmysh, and near 
the Chagan rivers. The common hare abounds in the former Orsk County, 
and in the former Ural Region, in B.A. Kuznesov's opinion (1928). According 
to M.K. Serebrennikov (1929) it is very rare in the northern Trans-Ural 
fore St -steppe. It is much more numerous in the southern steppes, sometimes 
more common than the blue hare. S. V. Kirikov thinks that the northern 
boundary of its range in the southern foothills of the Urals passes 40 km 
south of Zalair. 

According to S. S. Folitarek and A. A. Maksimov (1936) the common hare 
has recently become more numerous than the blue hare in the Verkhne- 
Uralsk district. According to Professor E.S. Danini (1931. 1933) the 
population of the common hare constitutes 65-75 % of that of the blue hare in 
the Troitsk District. 

There is no doubt that the common hare has recently begun to spread 
eastwards. 

According to A. M. Belyaev (1934) the range of the common hare reaches 
the Aktyubinsk Region in the east, to the Ural and Ilek rivers, i.e. . the 
western outskirts of the Kustanai District. He assumed that the eastern 
boundary of the range is the eastern part of the Ural-Mugodzhary Upland. 

These data are not quite accurate. It has now become known that the 
common hare occurs even farther into western Siberia and Kazakhstan. 
According to S. S. Folitarek and A. A. Maksimov (1936) common hares are 
widely distributed in the Turgai Territory, especially in the Kustanai 



107 



127 



District and near Lake Ubagan-Dengiz. The Kustanai District Bureau 
gives the following data: the common hare was rare in this district until 
1925, and only one common hare was obtained for every 10-15 blue hares; 
the population of common hares increased after 1925. In 1933-1934, 23,577 
specimens of common hares were caught, and only 14,393 blue hares. 

A.M. Korsakov (in litt. 1935) informed me that the common hare occurs 
not only in the Naurzum Reserve but also up to the boundary of the 
Akmolinsk [Tselinograd] Region. It first appeared in the Naurzum Pine 
Forest in 1915 (western coast of Lake Ak-Suat, 51°30'N and 64°29' E). It 
was only found 60-70 km further west previously, in the forest steppe of 
Sypsan-Agach and in the Teresek-Karagai Pine Forest. It was rare at 
first but became common in 1919-1920. 

The range reaches east of Kustanai-Turgai. It appeared in the Atbasar 
District in 1932. In the Petropavlovsk District it appeared in 1933, when 
150,000 blue hares and only 1000 common hares were shot. 

The manager of the Akmolinsk [Tselinograd] Fur Stores informed S. S. 
Folitarek that the common hare appeared in the Akmolinsk District in 
1931. It constitutes 0.1% of the number of blue hares at present. The 
Akmolinsk District should be considered as the extreme limit of the West 
Siberian range in the southeast (70° E. ). B. F. Belyshev's observation 
(in litt. 1934) that common hares were caught near Tara (75°E) is an 
exception. It was recently found near Omsk (A. V. Fedyushin, 1937, in 
litt. ). 

The common hare was observed west of the Kustanai District and south of 
Orsk, near the Dzhurun Station (200km south of Aktyubinsk) (A.N. 
Formozov, 1933). It is common in the Mugodzhary Mountains. 

E. P. Spangenberg obtained the Caspian common hare between the 
Murun-Kudun and Tyulek wells (north of Kazalinsk) in June 1928. He 
definitely saw common hares in the desert steppe 35-40 km from the Chelkar 
Station in summer. He caught his first specimen here among dry bulrushes 
near Lake Dzhumarn-Kul, and the second in the marshes in the hills. 
E. P. Spangenberg found many common hares near Akbulokh and Sagarty 
Stations in May. 

A skull of a specimen caught in 1921 by the expedition of Professor 
K.I Skrjabin near the Kara-Uzyak Station (45°N, 65° E) is in my collection. 

The common hare is widely distributed in eastern Europe. It occurs 
in the Astrakhan semidesert, in the Kalmyk Steppe in the south Russian 
chernozem steppes, the forest steppe and deciduous forests, to the western 
border of the U. S. S. R. The distribution of the common hare outside 
Russia has been described above. It is very common in the Baltic Region 
(K. Greve, 1909) although it was completely unknown in the former Livonia 
and Estland at the beinning of the last century. According to S. Chistovskii 
(1927) the common hare is common in the Pskov Region. 

It is occasionally very numerous in the Odessa Region, the Tauric 
Steppes and in the Crimea. 

It is very widely distributed in the northern Caucasus, from the belt of 
lowland- steppe to the foothills and high mountains. S. S. Turov (1926) 
recorded it from the Georgian Military Road near Kobi, in Siona, along 
the Terek river, near the village of Ozerakan, and in Tepi in the north 
Ossetian ASSR near the village of Zaromag. It is occasionally numerous 
in xerophytic mountain steppes of Dagestan. It occurs along the Black Sea 
Coast in Transcaucasia to the border of Asia Minor. It is widely distributed 
throughout Transcaucasia to the Mugan Steppe (K. A. Satunin, 1912). 

108 



128 



The common hare has thus spread widely during the last century in a 
northern, eastern and southeastern direction. S.S. Folitarek and A. A. 
Maksimov (1936) in their monograph on the common hare show that this 
species increased its range during the last century by approximately 45%, 
over an area of about 180,000 sq.km. 

In western Europe the common hare occurs in Great Britain (to the 
Orkney Islands), in Denmark, Poland, Germany, reaching the Iberian 
Peninsula, Italy, Corsica, Sicily, Roumania, the Balkans and the 
Peloponnesus in the south. 

It was introduced repeatedly into North America (1888 and 1893), in 
the northeastern part of the U.S.A. and East Canada. Several lots from 
Hungary (about 500 specimens ) were introduced into North America in 
1910-1911. 

The common hare has multiplied profusely in North America and become 
acclimatized (James Silver, Journal of Agricultural Research, 1924; C.G. 
Goodwin, The Mammals of Connecticut State, Geological and Natur. 
Survey Bulletin, No. 53, Hartford 1935, pp. 159-162). 

BIOLOGY. HABITATS. The common hare likes open country with 
sparse vegetation. In the central belt it lives in undergrowth and shrubs 
near fields, meadows, river valleys and gardens. It hides among weeds 
and in gullies. In the forest belt (where deciduous trees predominate), 
it avoids large forests, living only along their borders. It becomes 
gradually less common in the north, choosing places near large areas of 
felled forests or open river valleys. The increased exploitation of forests 
in the last ten years has been accompanied by an intensive northward 
migration of the common hare, usually penetrating new territories along 
the raain highways and railways. The hares do not occur in the forest 
itself. They live near villages, threshing floors, barns and gardens in 
the north, to a certain extent living alongside man. 




"^ 



FIGURE 69. 



Common hare, Lepus europaeus Pall 

Drawing by A. N. Komarov 



109 



129 



130 



The common hare is mainly found in ravines with oak shrubs (relicts of 
ancient forests) in the steppes of the central Chernozem Region, where, 
high grass and weeds grow in abundance. They prefer hard soil in the open 
steppe, where they are very common in fall. Such soil does not become 
muddy in the rain and does not stick to its hairy soles. It is very fond of 
weeds (such as in the Voronezh Steppe), mainly of Verbascum 
lychnitis and V. orientale, and also of Artemisia. A resting hare 
may be closely approached, and sometimes jumps out under the feet of 
passing people. In the vicinity of the Dokuchaev Experimental Station in 
the Kamennaya Steppe of the central Chernozem Region (former Bobrov 
County of the former Voronezh Province) the common hare usually keeps 
to the open steppe in mild winters, rarely hiding in the bushes or forests 
characteristic of the steppe near the Dokuchaev Station. It takes shelter 
either among bushes or within the forest in severe winters. They also 
frequently live near villages in winter, to feed on threshing floors at night. 

The typical habitat of the common hare in southern, semidesert steppes 
such as the Kalmyk Steppe is the hilly steppe (E.I. Orlov and B.K. Fenyuk 
1927). It preferably lives here among weeds, and hillocks overgrown with 
Tamarix, Elymus arenarius, Xanthium spinosum etc. It 
is probably attracted by the easy shelter, and makes its shelters among 
shrubs and in hollows. Hares also occur in meadows near lakes. Orlov 
and Fenyuk often startled hares from heaps of last year's thistle (Salsola 
kali ) in the Berovskie Hills. According to K. N. Rossikov (1887) the 
favorite habitats of the common hare in the northern Caucasus are 
brushwood, arable fields, pastures, weeds in the steppe, plains, river 
valleys, near forests, among shrub and loose thickets of reeds: and 
finally gardens and orchards near villages. K.N. Rossikov (1882) found the 
common hare in the barren sandy steppe inhabited by the Kara-Nogai 
nomads, southeast of the Kura. It occurs in the foothills and also in the 
high mountains. K.N. Rossikov (1887) writes: "One of my hunters killed 
an old hare which was a typical Lepus timidus Pall, while hunting 
Capra caucasica Giild. in the high mountains. " 

I found the common hare in the former Kizlyar Department, in the Terek 
Valley in the northern Caucasus. The hares were very numerous here in 
thickets of Elaeagnus , Hares are common in the foothills and in the 
xerophytic upland steppes of Dagestan. They also occur in the high 
mountain zone of the Caucasus, as recorded by K.N. Rossikov, in which it 
displays unusual dexterity in climbing steep cliffs although considered a 
plains dweller. 

The above survey of habitats demonstrates the great adaptability of the 
species to different environments. 

GENERAL DESCRIPTION AND HABITS. The hare is an active and 
graceful animal. Its sense of hearing is excellent, its sight is weak, but 
its sense of smell is apparently well developed. It runs rapidly. The 
steppe hares (L. e. tesquorum), usually called "feather- grass' hares" 
by the hunters, are particularly swift. Their speed may sometimes be 
considerable. According to I.I. Almazov ("Okhotnich'ya gazeta" (Hunter's 
Gazette) No. 2, 1928, p. 5) a hare ran in front of a train for 2km. The 
speed of the train was 55-60 km. This hare had been first chased by a dog, 
but when the dog fell behind, it was "chased" by the train. 

The track of the common hare differs markedly from that of the blue 
hare. Its feet are narrower, and the toes are not separated as widely as 



110 



in the blue hare, which is more adapted to run over loose snow. The track 
of the common hare is therefore narrower, and the traces are more 
elongate. 




FIGURE 70, A common hare running. (Lepus europaeus Pall.) 

Drawing by A. N. Komarov 



The space between the footprints is also narrower than in the blue 
hare. The imprints of the forefeet (in normal slow gait) are more widely- 
separated than in the blue hare. The whole group of imprints of the common 
hare in normal gait is more elongate*. 

The fecal pellets of the common hare differ slightly from those of the 
blue hare; they are elongate with a sharp tip. They are about the size of 
a hazel nut. The pellets of the white hare are more oval, with broadly 
rounded tips and slightly compressed laterally. 

The common hare is a good swimmer. I.S. Slesarev informed me that 
he saw a frightened hare jumping into a broad stream and swimming across. 
The ears were slightly inclined backwards. Its swimming capacity is, 
oj however, limited because its paws are adapted to running, not to swimming. 
Many hares therefore die during the spring and summer floods after 
prolonged rains. 

S.S. Folitarek and A. A. Maksimov (1936) write: "Many hares, 
especially common hares, die during the high floods. The sharp decrease of 
their numbers after the floods has a marked effect on the results of hunting 
in fall. 

"We had the opportunity to see the great influence of spring floods on the 
hare population in 1933. in the lower reaches of the Tsna river, from Sasovo 
to the inflow of the Tsna river into the Moksha river, and from Kadym to 
the mouth of the river. There was a great flood in 1932, 10-15 km wide 



* The differences between the track of the hare in slow and rapid progression, and the description of turns 
and loops are given in the chapter on the biology of the blue hare. 



on the Tsna and Moksha rivers. There were nearly no hares in these 
places in 1933, although conditions in general were favorable. 



132 




FIGURE 71. Tracks of the common haie in the snow 

Drawing by A. N. Formozov 

"We found shreds of fur and the bones of hares in two places in osier- 
beds near the water where drowned hares apparently became lodged. We 
were informed that hares are widely hunted in spring in this area. Hunters 
in boats in the flooded forest searched the wooded ridges and frightened 
hares on them. The animals jumped into the water and were killed with 
sticks. " 

I know of many cases in which hares died in the floods, unable to swim 
long enough. 

DAILY LIFE. Although the hare is occasionally active during the day, 
it is mainly a crepuscular animal. It comes out to feed after sunset, and 
remains active until dark. It continues feeding and moving about at night 
in winter, leaving its tracks in the snow. Early in the morning, before and 
after sunrise, before the dew disappears and the heat begins, hares can be 
seen moving about and feeding. It usually rests during the day in its 
shelter among vegetation. In the steppe it is found in thickets of weeds and 
shrubs. 

It is interesting to watch the hare feeding on the steppe and to observe 
the habits of this animal the life of which is full of perils. 

I wrote in my diary of the Voronezh expedition (1922): " I watched 
a hare feeding in the evening between patches of forest in the Kamennaya 
Steppe. I was trying to approach it when something unexpected and rather 
funny happened. Two swallows appeared. Apparently mistaking the hare 
for a cat, they attacked it with loud cries and with the same vigour with 
which they attack cats. The hare ran away as fast as it could. It stopped 
for a moment from time to time, and when the swallows repeated their attack 
it ran away again. " 

I saw a fox chasing a hare in the Kamennaya Steppe on 15 July 1922 in 
the evening. The fox soon gave up. The hare stopped for a moment and 
resumed its rush. It then stopped near a marmot burrow from which a 



112 




* ' 



■-■~. . -«i<- 



FIGURE 72. Common hare in summer 



Photograph by S. I. Ogne\ 



113 



large old marmot emerged. The hare sat down 5 steps from the marmot 
and raised its ears and forefeet. Both animals eyed each other in dumb 
surprise for some time and then the hare dashed into the open steppe, 
fleeing from the dangerous spot. " 

BURROWS AND LAIRS. As has been said, the lairs of the common 
hare are usually in "secure" places, under bushes, in thickets of high grass 
and weeds, overgrown wheat fields, underbrush in ravines, tamarisk thickets 
in the semidesert etc It thus has generally no permanent refuge. It 
often burrows in the snow under a shrub and rests there during the day. 
In sandy and semidesert areas, it sometimes deviates sharply from its 
typical habits. For instance, A.N. Formozov found southern common hares 
living in burrows dug in the sand on Dzharylgach Island in the Black Sea 
between the Crimea and the Dnieper estuary. I.I. Barabash-Nikiforov 
(1928) writes that in the Sivash Steppes hares are frequently attacked by 
sea eagles (H aliaet us albicilla). They therefore often dig burrows 
in the snow which may be 1.5 m long. In one case, a burrow had 2 lairs 
together at the end of the tunnel. 

A chased or wounded hare sometimes finds shelter in an old marmot 
burrow in the steppe. Hares may hide in fox burrows in the central forest. 

According to V.N. Kaverznev (1928), "hares sometimes use abandoned 
burrows of foxes. I saw a wounded hare hiding in a burrow so that I could 
not reach it. " 

FOOD. P. A. Manteifel (1927) writes: "The food of the common hare 
is more nourishing than that of the blue hare in the forest. As it lives near 
settlements, it visits gardens, digs up vegetables in winter, enters hay barns, 
feeds on haystacks, gnaws cabbage stumps in gardens, and in the 
February-March frosts it walks on the hard snow and enters orchards where 
it gnaws the bark of fruit trees, often causing great damage. " 



i ' 




FIGURE 73. Common hare hiding in winter 

Drawing by A. N. Formozov 



114 



A.I. Yurin (in litt. 1934) informed me that the common hare is very- 
fond ofHelianthus tuberosus. 

He wrote: "I found a new method of catching hares. Blinds are erected in 
Helianthus plantations, or near stalks ofHelianthus placed for this 
purpose. Helianthus left in the field in winter is one of the favorite foods 
of hares. " 

A small hare which I kept accepted sage (S a Ivi a s i Ive s t r i s), plantain, 
wintercress (Barbarea vulgaris arcuata), peas, potatoes, and 
knotgrass. It also liked bread moistened with milk. When eating a plant, 
the hare does not hold it with its forelegs as other rodents (e.g., marmots) 
but seizes the food with its mouth. 

REPRODUCTION, YOUNG AND THEIR DEVELOPMENT. Our knowledge 
of the reproduction of the common hare is even more incomplete than of that 
of the blue hare . 

The female produces 3 or 4 litters from spring to fall; each litter usually 
consist of 3-5, rarely of 7 young. 

The first litter is normally born in March in the Moscow Region, and the 
last litter in September (rarely in October). But these dates vary greatly 
with the climate. Thus, I.S. Slesarev (in litt.) informed me that he shot a 
gravid female near El'nya (in the former Smolensk Province) in the mild 
winter of 1925 on 15 January. 




FIGURE 74. Common hare sitting 

Photograph by S. I. Ognev 

E. V. Sharleman (1915) observed rutting hares near Kiev in late 
February. He saw several males chasing a female at midday on 20 
February 1912, completely ignoring people. In Sharleman's opinion, 
copulation may take place even earlier, because he had already seen 
newborn hares in March. He found young in March 1913 which were half 
the size of adults. He found newborn hares in the same year, on 28 March 
and 15 April. K. F. Kessler usually found the first newborn hares in the 



115 



Kiev Province in late February, when the ground was still covered with 
snow. E. V. Zverezomb-Zubovskii (1923) found the last litters in late 
September near Rostov-on-Don. K.N. Rossikov (1887) saw newborn litters 
in 1882, 1883, and 1884 in North Caucasus near the Cossack village of 
Prokhladnaya in early April, early June and late August to early 
September. -■= 

The exact period of gestation of the common hare is unknown ** 
According to E. V. Zverezomb-Zubovskii (1923) the young may be nursed 
for 8 weeks. 

P. A, Manteifel' (1935) gives the following data on the reproduction of the 
common hare: 

"The blue hare brings its young to open places where they scatter in all 
directions. On the other hand, the common hare in the open country digs a 
shallow pit in which the young hide from raptors. 

"The common hare feeds its young more frequently and protects them 
more energetically, especially when they cry. I found that the presence of 




FIGURE 75. Young common steppe hare, Lepus europaeus 
tesquorum Ogn. , Kamennaya Steppe, fonner Voronezh Province 

Photograph by S. I. Ognev 



♦ We converted Rossikov's dates to the New Style. 
+* E. A. Manteifel' informed me that the gestation period is 44-46 days, i. e. , shorter than in the blue hare, 
according to observations in the Moscow Zoo. 



16 



young of previous litters is fatal for newborn hares. The older specimens 
136 consume all the milk of the females. The females let the young feed, 
without touching them or driving them off. Common hares apparently 
change places for different broods, to prevent starvation of the newborn. 
Mass multiplication of hares usually gives rise to epidemics of infectious 
diseases. " 





nCURE 76. 



Postures of females of common hare (left) and the blue hare (right) during copulation. 

After P. A. Manteifel' 



MOLT. The fall molt usually starts in early October and ends in late 
November in the Moscow region. The spring molt is more rapid, lasting 
from April to early May. 

MIGRATION, DISPERSAL AND FLUCTUATION OF POPULATION. 
We described the wide dispersal to the north, east and southeast in the 
survey of the geographical distribution. S.S. Folitarek and A. A. Maksimov 
(in litt. 1936) showed that the range of the common hare increased by 
approximately 45% during the last century, as it occupied an area of about 
180,000 sq. km. 

The fluctuations in populations are discussed in detail in the description 
of the biology of the blue hare. At present, it should only be noted that these 
fluctuations show a certain periodicity. 

In the southern and southwestern regions of Russia, population 
fluctuations take place at intervals of about 5 years. This corresponds to 
the observations of Biger (1928) in Germany, in forests in Mecklenburg- 
Schwerin, 

Fluctuations of populations of the common hare are not as great as those 
of the blue hare. According to A.N. Formozov (1935) common hare 
populations fluctuated between one and ten in the Crimea, between one and 
seven in the Leningrad Region, between one and six in the Ivanovo Region 
and Bashkiria, one and three in the Gorki (former Nizhni Novgorod) 
Territory and Tataria etc. A.N. Formozov gives the following explanation 
for the smaller fluctuation of the populations of the common hare: l) a 
more uniform and regular exploitation of the population by hunters; 2) 
the relative immunity of the common hare to diseases because it lives in 
drier country than the blue hare. The second factor is of greater 
. „„ importance in our opinion. 

There are many factors which regulate the populations of the common 
hare. It is difficult to define them and to evaluate their relative importance. 

The following factors regulate the population of hares: 



17 



l) Strong floods, which will be discussed in detail in the description 
of the blue hare. It should be stressed that the common hare prefers 
river valleys in the forest belt and spreads north along them. However, 
this preference for river valleys is often fatal for them during strong spring 
floods. 




FIGURE 77. Young Crimean common hare, Lepus europaeus 
transsylvanicus Matschie 

Drawing by A. N. Formozov 







FIGURE 78. Nest of common hare with young 



Photograph by S. I. Ognev 



1254 



138 



2) As the common hare lives in open country, it is less endangered by 
forest fires than the blue hare. 

3) Cold spells in spring after the onset of warm weather are often fatal 
for the young of the common hare . 

S.S. Folitarek and A. A. Maksimov (1936) state: "The common hare is 
of southern origin. After having moved north, it retains the reproduction 
rhythm of more southern latitudes. The northern summer is short, and 
the hares reproduce in haste. We often found litters which were born too 
early or too late, and which thus had to die. This happened most frequently 
in spring when the onset of hot weather is followed by sudden cold snaps 
(often with snow). The first litter of the common hare often dies during 
these short periods of cold. This also happens during long cold springs. 
There are records of mass mortality of hares in the spring of 1905, 1927- 
1928 and 1931. Warm April weather was followed by severe cold in the 
first days of May in 1933 near the Moksha river in the Moscow Region. 
Water was iced over in the morning and snow fell on 6 May. A local' hunter 
found four newborn hares in a meadow near the village of Koshebeevo at that 
time. Two of them were already dead, and the others were frozen. The 
hunter took them home, warmed and revived them. " 

4) S.S. Folitarek and A. A. Maksimov (1936) skeptically quote stories 
of mass mortality of hares during severe winters as a result of blizzards, 
snow and starvation. This is correct, but rare. The influence of the 
climate is increased by indirect factors. For instance hunting by 
man is intensified as exhausted and starving hares enter settlements and 
villages. B.K. Fortunatov (1929) gives very interesting illustrations of 
this phenomenon. 

He writes: "The year 1927 was very favorable for the hares throughout 
the Ukraine. Their population increased considerably in fall. 

"The first half of the 1927-1928 winter was relatively mild and there was 
little snow. However, on the night of 3rd February a savage blizzard 
broke out in Ascania-Nova which covered buildings and parks with 
snowdrifts. 

"We concentrated all our efforts in the first two days to dig out all zoo 
animals which had been buried by snow. It was impossible to go into the 
steppe, as this meant certain death. However, the wind subsided and snow 
ceased to fall in the night of 4-5 February. 

"The night was clear. The full moon illuminated the steppe which was 
covered with thick snow. I stood on a mound of snow under which the sheds 
of the American bisons were buried. We noticed black chains moving 
towards us from all directions one hour after sunset. We realized that they 
were hares fleeing from starvation. The courtyard of Ascania-Nova was 
thronged with thousands of hares in half an hour. 

"As a result of the shock suffered in the steppe, or due to hunger, the 
hares had almost completely lost their fear of men. They ran at our feet, 
fell greedily upon every scrap of fodder, and hundreds huddled near haystacks 
at the borders of the estate. During the same night, we carried several 
haystacks to the steppe, and thousands of hares were thus saved. 

"The wind calmed down the next day, and we found many dead hares upon 
examination of the steppe. Not all of them had died of hunger. Even a few 
days later, dead hares were still found in Ascania-Nova where there was 
sufficient food. Such a severe blizzard was apparently sufficient to kill 
even the best adapted wild animals. 



19 



"Ascania-Nova attracted and saved only a relatively small part of the 
hares. Most of them headed for the hamlets and haystacks near them. 
They were all killed by the peasants, as the bright moonlight enabled them 
to shoot as easily as during the day. An even greater number of hungry 
139 and desperate animals were killed with sticks. Hunters carried away 
hundreds of hares caught during the night. 

"The February events could be summed up a month later. Several 
thousand hares obstinately remained near Ascania-Nova, not moving more 
than one kilometer away. No hares were found beyond this line. I saw 
tracks of hares only twice in late February, on the 70 km between 
Ascania-Nova and the Novaya Alekseevka Station. " 

5) Epizootics decimate hare populations, both of common and blue hare. 
We will discuss the magnitude and characteristics of these epizootics in 
the description of the biology of the blue hare. 

The mortality of common hares as a result of epizootics is intensified 
by climatic factors. Mortality is increased in winter by: l)deep snow 
which hampers the movements of the narrow-footed common hare, and 
exhausts it; 2) snowstorms (see above); 3) severe frosts; 4) iced-over 
ground. Common hares contract diseases more easily during very rainy 
summers. 

DAMAGE. Damage to crops caused by hares is negligible in summer 
because of the large size of the fields, but when deep snow falls in the steppe 
and buries the vegetation, the hares migrate to the villages. If they feed on 
straw and hay, the damage is small, but if they enter orchards and gnaw 
bark of trees and the young tree tops in plantations, the damage becomes 
more severe They cause great damage to fruit orchards and vineyards in 
the Crimea and the Caucasus. 

The common hare is less numerous than the blue hare in the forest and 
is less widely distributed there than in the steppe. The damage it causes 
is therefore negligible. V.N. Kaverznev (1928) correctly states that the 
common hare can be made harmless by rational methods. This can be 
attained by supplementary feeding, putting out boxes with fodder, stumps, 
and rotten cabbage leaves. Such supplementary feeding would avert its 
harmful activity in severe winter and would prevent its destruction as a 
pest. 

ENEMIES. It is usually assumed that the fox is one of the main enemies 
of the common hare. However, the hare does not occupy first place in the 
diet of the fox. I.I. Zharkov, V. P. Teplov and V. I. Tikhvinskii (1932) 
examined the excrement of 508 foxes in Tataria. Remains of hares were 
found only in 12.6%, but remains of small rodents (mice and voles) were 
found in 49.2%; hamsters in 28.9%, of susliks in 17.5%, of birds in 11.2%, 
of insects in 17.7%, carrion in 2.6%, and amphibians in 0.8%. 

The common hare is hunted by wolves. It may be chased by the lynx 
,40 in the forest zone, which it occasionally enters, but the lynx feeds mainly 
on blue hares. Stray and shepherd dogs cause great damage to the common 
hare. 

Goshawks, golden eagles, spotted eagles, A qu i la clanga, andrarely 
sea eagles and eagle owls prey on hares. I.G. Pidoplichka (l930) found 
only remains of one hare among remains of 22,000 animals in excrement 
of owls and came to the conclusion that owls do generally not prey upon 
hares. 



120 



COMMERCIAL VALUE. We shall discuss the great commercial value 
of hares for the fur trade of the U. S. S. R. and as food in the description of 
the blue hare. It should only be stated here that the supply of fur varies 
considerably in different years. According to B. A. Kuznetsov (1932) the 
fluctuations compared with 1924-1925 (taken as 100%) ranged between 
100-335.5%. Pelts of the common hare constitute 1.6-5.6% of the total 
Russian fur trade. 



Subspecies of Le pus europaeus 

327. Lepus (Eulagus) europaeus europaeus Pall, (1778). 
European comraon hare 

1756. Le Lievre. Daubenton L. J. M. , in E. L. le Clerc, Comte de Buffon. Histoire Naturelle, VI, pp. 264- 
299 (described from a specimen from Burgundy, France). 

1777. Lepus timidus (nee Linnaeus.'). Erxleben J. C. R., Systema Regni Animalis, p. 325 (partim.'). 

1778. Lepus (europaeus) Pallas P. S, , Novae Species Quadrupedum e Glirium ordine, p. 30 (terra 
typica not stated). 

1801. L[epus] t [imidus] albus Bechstein Job. Matth. , Gemeinniitz. Naturgesch. Deutschlands nach 

alien 3 Reichen, I. Editio 2, S. 1096 (Thiiringen, Germany). 
1801, L [epus] t [imidus] flavus Bechstein Job. Mattb. , Op. cit. , S. 1096 (Tbiiringen, Germany). 
1801. L [epus] t [imidus] n i g e r Bechstein Job. Mattb., Op. cit., S. 1097 (Thiiringen, Germany). 
1811. Lepus ti mi dus (nee Linnaeus.'). Pallas P. S. , Zoograpbia Rosso-Asiatica, pp. 148-149 (Poland 

and Lithuania). 
1857. Lepus timidus (nee Linnaeus.'). Blasius J. , Naturgeschicbte der Saugetbiere Deutschlands, S. 

412, 417 (Mittel-europaische Form L. tim i dus auct., L. c a mpi col a Schimp. ). 
1859. Lepus campicola Gervais P. , Zool. et Paleontol. fran^aise, Editio II, p. 47 (Nomen nudum.' It 

describes a hare in France). 
1867. Lepus timidus: coronatus, rufus, cinereus, nigricans, maculatus, albus et 

niger. Fitzinger L. , Sitzungl. d, Kaiserl. Akademie der Wissenscb. , Wien, Mathem. Naturwissensch. 

Klasse, LVI, S. 161 (Nomina nuda.' Hares from Austria are described). 
1910. Lepus europaeus Pall. Trouessart E. L. , Faune des Mammiferes d'Europe. Berl. , S. 219. 
1912. Lepus europaeus europaeus Miller G. S. , Catalogue of the Mammals of Western Europe, 

pp. 502-504. 

TYPE LOCALITY: G. S. Miller (1912) proposed to consider Burgundy (France) as the terra typica of 
L. e. europaeus for the following reasons: 1) in Pallas "Novae species glirium", p. 2, reference is 
made to the description of Daubenton in "Histoire Naturelle" of Buffon; 2) in Miller's opinion (op. cit. p. 5) 
Pallas had another form in mind when he discussed the distribution of the bare in Poland; 3) Pallas does 
not mention the distribution of the common bare in the very short description on p. 30. 

Pallas quite casually mentions a hare measured by Daubenton, on p. 2. He mentions— doubtfully— a 
hybrid between L. variabilis and L. europaeus found in Poland and Lithuania on p. 5 ("... banc, 
mibi vix dubium est, varietatem vere hybridum esse productum, ex adulterio L. variabilis cum L. 
europaeo, australiorem Poloniam atque Pannoniam occupante, progenitam et in istis regionibus 
multiplicatam. . . "). 

In his "Zoograpbia Rosso-Asiatica", 1811, p. 148 Pallas gives the range of the common hare (Lepus 
timidus) precisely and clearly: "In Rossiae australibus, ultra 55 tum latitudinis gradum, haec sola species 
occurrit, ut et in Polonia et Lithuania. . . ". 

E. L. Trouessart (1910) was correct, in our opinion, in stating that the terra typica of the common bare 
is Poland. 

We consider southwestern Poland as the type locality of the nominate race, where hares with well 
marked characteristics of the type are found. 

MATERIAL EXAMINED: 7 pelts and 4 skulls. 

DIAGNOSIS. Fur on back wavy, slightly curly, clay-rusty-yellowish, 
without a dull earth- gray and black tinge. Color of winter fur hardly 
different from summer fur; only the lower part of the back is slightly 
grayer. 

121 



142 



Length of skull usually less than 100 mm; hind feet less than 150 mm 
(usually about 140 mm). 

ADDITIONAL DESCRIPTION AND SYSTEMATIC NOTES. Color, 
waviness and curliness of summer fur of the typical common hare does 
almost not differ from that of central Russian hares (L.e. hybridus). 

The winter fur of the western common hare differs sharply from that 
of L.e. hybridus. The color of the winter fur of L.e. europaeus 
hardly differs from that of the summer fur, the former is, of course, 
denser, longer and softer. The winter fur of L.e. europaeus lacks 
gray hair; there is occasionally a slight gray tinge on the lower part of the 
back, near the tail. 

I gave a detailed critique of the color characteristics of the nominate and 
central Russian races given by Hilzheimer and Trouessart in "Mammals 
of the Moscow Province" (Mlekopitayushchie Moskovskoi gubernii), 1913, 
p. 292-293. I confirmed, provisionally, the following data given by Hilzheimer 
(1908) on the basis of the small material at my disposal: 1) shoulders of 
the German common hare compared with that of the central Russian race 
much darker reddish brown with a marked clay tinge; 2) anterior sides of 
forelegs of German hare duller, with more brownish tones and with 
numerous black hairs. Since then I re-examined many common hares and 
found that black hairs on the anterior parts of the forelegs also occur in 
L.e. hybridus. On the other hand, I saw typical common hares from 
Germany without such black hair. 

L.e. europaeus is clearly distinguished from L.e. hybridus by 
the small size of body and skull, and characteristic color of its winter fur, 
L. A. Obrecht informed me that of a large number of central German hares 
examined none weighed more than 3,5-5.8 kg. This is much less than the 
we ight of L.e. hybridus. 

I am inclined to consider hares from the vicinity of Novaya Aleksandriya 
[Pulawy] and Kiev (vicinity of Pochapenets, Zvenigorodka District) as 
intermediate between L.e. europaeus and L.e. hybridus. These 
specimens resemble L.e. europaeus in the small difference in color 
of winter and summer fur, and in the slight gray color of the lower part of 
back and cheeks. Their measurements are identical with those of L . e . 
hybridus . 

The characters of L. e. europaeus are particularly well marked 
in specimens from its western range. Thus, German common hares are 
smaller than Polish, and French specimens are smaller than German 
common hares (E. L. Trouessart, Faune des Mammif^res d' Europe, 1910, 
p. 219). 

Measurements of L. europaeus europaeus: length of skull 89 - 
99 mm (M. 97.7 mm); condylobasal length 84-87.7 mm (M. 86.4 mm); 
zygomatic width 44-48 mm (M. 46.1 mm); length of nasals 44-47 mm 
(M. 44.4 mm); width of nasals 16-17 mm (M. 16.9 mm); length of bony 
palate 6.1-6.4 mm (M. 6.2 mm); width of incisive foramina 11-12 mm (M. 
11.4 mm); length of upper tooth row 17.0-17.6 mm (M. 17.4 mm). 

G.S. Miller (1912) gives the following cranial measurements for the 
typical subspecies: occipitonasal length 94-102.2 mm; condylobasal 
length 85. 2-92. 2mm; zygomatic width 45.2-50.2 mm; interorbital width 
18. 4-24. 8mm; postorbital width 11. 4-15. 2mm; width of brain case 31- 
34.6 mm; length of nasals (along oblique outer margin) 42.2-46.2 mm; 
maximum width of both nasals 20.6-24.6 mm; diastema 26. 8-31. 8mm; 



122 



length of upper tooth row 17. 2-19. 2mm; length of lower tooth row 18.6- 
20.2mm. 

Measurements of specimens, l) From Hasley, Zealand (Denmark) 
2 males ad. and female ad. : length of body 660, 680 and 640 mm; tail 80, 
82, and 74mm; hind feet 136, 147 and 138 mm; ears (measured from 
occiput of live specimen) 138, 131 and 125 mm. 2) Rheinhessen, Ingelheim 
(Germany) male ad. : length of body with head 571 mm; tail 98 mm; hind 
foot 135mm; ear (measured from occiput of dried skin) 120 mm. 3) 
Rheintal, St. Gallen (Switzerland) 2 males ad. : length of body with head 
570 and 540 mm; tail 100 and 85 mm; hind foot 136 and 137 mm; ear (measured) 
from occiput on pelt) 117 and 115 mm. 4) Verdenberg, St. Gallen 
(Switzerland) 2 males ad. : length of body with head 600 and 605 mm; tail 
90 and 95 mm; hind feet 145 and 144 mm; ear (measured from occiput) 
114 and 118 mm. 

GEOGRAPHICAL DISTRIBUTION. The nominate subspecies occurs in 
central Europe from the Atlantic Coast of Denmark to Central France, and 
also throughout Germany, Austria, and Switzerland. It is also found in 
western and central Poland (E.K, Lubicz-Niezabitowski, 1934). 

According to G. S. Miller (1912) this form has been recently introduced 
into southern Sweden. 

328, Lepus (Eulagus) europaeus hybridus Pallas (l81l). 
Central Russian common hare 

1811, Lepus variabilis var. hybrida Pallas P. , Zoographia Rosso- Asiatica, pp. 147-148. 

1820. Lepus hybridus Desmarest' M. A. , Mammalogie ou description des especes de Mammiferes, I. Paris, 

p. 349. 
1842. Lepus aquilonius Blasius I. , Amtl. Bericht XIX Versamml. Naturf. und Aertzte, Braunschweig, 

p. 89 (Central Russia). 

1850. L. t[imidus] var. hyemalis Tumac, Eversman, E. , Estestvennaya istoriya Orenburgskogo 
kraya (Natural History of the Orenburg Territory), part II, Kazan, p. 201. 

1851. Lepus medius Middendorf A. Th. , Ueber die als Bastarde Angesproch. Mittelformen zu, L. 
europaeus undL. variabilis, Bull. phys. -mathem. , IX, No. 14, S. 13-25 (- a qui 1 on 1 us 
Blasius). 

1851. Lepus a qu ilonius Simashko Yu. , Russkaya Fauna. Mlekopitayushchie (Russian Fauna. Mammals) 

II, pp. 782-785, pL62, fig. 2. 
1871. Lepus timidus P. hyemalis (=hybri dus) Bogdanov, M. N. , Ptitsy i zveri chernozemnoi 

polosy Povolzh'y^ (Birds and Mammals of the Chernozem Belt of the Volga Region), pp. 175-176. 
1889. Lepus timidus var. tu m ak Tikhomirov A. A. and Korchagin A, N. , Spiski i opisanie predmetov, 

nakhodyashchikhsya v zoologicheskom muzee Moskovskogo universiteta (Catalogue and description of 

Specimens in the Zoological Museum of Moscow University), Izvestiya Obshchestva lyubitelei 

estestvoznaniya, LVI (4):31 (named from a hybrid between blue hare and common hare). 
1908. Lepus medius aquilonius Hilzheimer M. , Die Hasenarten Europas, Jahrh. d. Vereins f. 

Vaterl. Naturk in Wiirttenb. , p. 388. 
1910. Lepus medius aquilonius Trouessart E. L. , Faune des Mammiferes d'Europe. p. 217 (Central 

Russia). 

1912. Lepus europaeus hybridus Miller G. S. , Catalogue of the Mammals of Western Europe, 
pp. 508-509. 

1913. Lepus (Eulagos) europaeus aquilonius Ognev S. I. , Mlekopitayushchie Moskovskoi 
gubemii (Mammals of the Moscow Province), pp. 283-298. 

1929. Lepus (Eulagos) europaeus hybridus Ognev S. I. , Zur Systematik der russischen Hasen, 

Zoologischer Anzeiger, Bd. 84, H. 1/4 p. 74. 

TYPE LOCALITY AND POSITION. Former Moscow Province (for details, see systematic notes). Type 
lost. 

MATERIAL EXAMINED. 46 pelts and 61 skulls. 



123 



144 



DIAGNOSIS. Fur on back wavy, in summer rich clay- rust -yellow 
without a dull earth-gray tinge and blackness. 

The color of winter fur differs sharply from that of summer fur because 
of the marked gray and whitish tones. 

Length of skull usually more than 100 mm; length of hind feet more than 
150mm (usually 160-I70mm). 

ADDITIONAL DESCRIPTION. Weight ofL.e. hybridus about 
4-4.5 kg. G.I. Polyakov reported that the heaviest of 40 common hares 
killed in the former Bogorodsk County of the Moscow Province weighed 
5.2 kg. Larger weights are very rare. 

G,I. Polyakov sent me a document signed by three people who testified 
that A. A. Kustarev killed a female hare (not pregnant) on December 24, 
1915 in the former Pokrov County of the Vladimir Province which weighed 
10.2 kg. Such specimens are of course exceptional. 

A detailed description of summer and winter fur of adult L . e . 
hybridus has been given above . 

The general color of the fur of head and back of very young hares about 
half the size of adults is straw-yellow-sandy speckled chestnut-brown. 
This color is caused by isolated hairs of the underfur. Basal parts of 
underfur dull gray (hair-brown, Ridgway, 1912); their apical parts more 
straw colored (drab). Color of giaard hair as follows: base brown-gray; 
middle part blackish brown; subapical part pale, straw-sandy; apex 
blackish brown. 

A broad, pale straw-gray stripe across the chest. Legs of the same 
color. 

There is often a white spot on a straw-brown background on the forehead. 

Young L.europaeus hybridus of this age are of a lighter and 
duller color than young L. e. tesquorum. 

The color of young L.e. hybridus (about half the size of adults) differs 
generally only little from that of very young specimens. 

The general color of the fur on head and back varies between shades of 
yellowish straw-gray with a more or less pronounced sandy tinge. Rust- 
gray shades on chest and flanks, and on a broad transverse stripe on the 
chest varying between wood-brown (brownish gray with pinkish straw) to 
grayer shades than pinkish cinnamon (rust-ocher). 

Measurements of L . europaeus hybridus: length of skull 99. 3- 
106.5 mm (M. 101.9 mm); condylobasal length 89.1-97 mm (M. 91.9 mm); 
zygomatic width 46.9-51.2 mm (M. 48.5 mm); length of nasal bones 46.1- 
49.5mm (M. 47.5mm); width of nasals 15.8-21.1 mm (M. 19.2mm): 
length of bony palate 5.3-8.1 mm (M. 6.6 mm); width of incisive foramina 
11 -13. 5 mm (M. 11.8 mm); length of upper tooth row 17. 1-20. 2 mm (M. 
18.6 mm); hind feet 164- 175 mm; ear including terminal hair (measured 
from forehead) 110- 129 mm. 

SYSTEMATIC NOTES. Pallas (1811) noted the variety Lepus 
variabilis var. hybrida. He gave the following description: 
"Hyeme non plane alba, quam Rossi sequentis nonnine Russak apellere 
Solent". He does not give the range of "var. hybrida". 

M. A. Desmarest (1820) in "Mammalogie ou description des especes des 
Mammiferes", mentions "Le p u s hybridus Pall., russak des Russes". 
He gives the following range for it: "Habite certains cantons de la Siberie 
et de la Russie". 



124 



PLATE III 




Central Russian common hare (Le pus europaeus h ybri dus Desm. ) (foreground). Steppe common 
hare(Lepus e. tesquorum Ogn. ) (background) 



125 



I. Blasius (1842) mentions a hare from central Russia under the name 
Lepus aquilonius. 

Blasius (1857) in "Naturgeschichte der Saugethiere Deutschlands", 
p. 412, 417 makes L . aquilonius a synonym of L . timidus 
(= e u r o p ae u s ). On later pages Blasius separates a "northeastern form" 
(nordostliche Form), to which he ascribes L. caspicus, L. 
aquilonius, L. medius and L. variabilis var . h y b r i d u s . 

E. Eversmann (1850) mentioned the variety: hyemalis Tumac, with 
the following description: "Albicans, vultu, mediodorso, colli lateribus 
paginaque antica antipedum griseorufis". He writes in "Natural History of 
the Orenburg Territory" (Estestvennaya istoriya Orenburgskogo kraya) 
p. 204, "This race of the common hare named by Tumac turns white in 
winter. " Later, Professor Eversmann suggests doubtfully that the Tumac 
hare (var . hyemalis) may be a hybrid between the common and the 
blue hare . 

The central Russian common hare described by us was described without 
a doubt by Academician Middendorf (1851) under the name Lepus 
medius . 

G. S. Miller (1912) mentions this form under the name Lepus 
europaeus hybridus Desmarest, and considers central Russia as its 
.J- terra typica. The range of this form should be defined more accuratelv, 
in my opinion, and the former Moscow Province should be considered as 
the type locality of L . e . hybridus (see S. Ognev, Zoologischer 
Anzeiger, 1929, S. 74). 

I have discussed the so-called Tumac hare in sufficient detail ("Mammals 
of the Moscow Province" (Mlekopitayushchie Moskovskoi gubernii) 1913, 
p. 294-295). There is confusion in the old hunting literature about the 
identity of the Tumac hare. Some zoologists also confused the Tumac 
hare (a hybrid) with the central Russian common hare (a subspecies). The 
winter fur of the latter becomes much lighter than the clay-yellowish 
brown summer fur of the Tumac hare. M.N. Bogdanov (1871) named 
the Tumac hare "steppe common hare", but did not give its exact range 
and gave only a short, inexact diagnosis. Bogdanov also gave the Tumac 
hare (or, according to him, the steppe common hare) a special name, 
Lepus campestris Bogd. 

Pallas (Zoographia Rosso-Asiatica, 1811, p. 147) mentions, in his 
description of the blue hare, that: "varietas forte hybrida, hyeme non 
plane alba, quam Rossi sequentis nomine (Russak) apellere solent. . . ". 
He doubts whether this variety should be considered as a hybrid between 
two species. 

According to the hunting literature, the Tumac hare (hybrid) is more 
agile and runs faster than the blue and common hares. 

It seems quite clear that there are 3 separate forms: l) a race of the 
common hare L.e. hybridus, the winter fur of which is much lighter' •. 
2) a steppe form called Tumac hare by M.N. Bogdanov; 3) hybrid between 
the common and blue hare, which is correctly called Tumac hare (this 
name is thus used only for the hybrid form). 

I already pointed out in the past that the characteristic features of the 
blue hare usually predominate over those of the common hare in the Tumac 



♦ The Turgai common hare L.e. caspius natio turgaicus Ognev, the description of which follows 
below, becomes even more white in winter. 



126 



hare. But there appear new specific characters as well, such as the rust 
color of the base of winter hair, which do not occur in the common and 
blue hares. The structure of the skull of the Tumac hare is in many- 
respects intermediate between that of the two parents. This was pointeed 
out by E. Lonnberg (Proceed. Zool. Soc. London 1905, pp. 278-287). 
Skull No. S. 2399 (collection of the Moscow Zoological Museum) from the 
former Saratov Province is very similar to the skull of the blue hare. 
However, it resembles that of the common hare in the following characters: 
l) premaxillary more slanting and longer than in the blue hare; 2) first 
lower molar inclined backwards. Orbit delimited by the thick zygomatic 
bone which is elevated posteriorly at a considerable angle and generally 
resembles that of the blue hare. 

Specimens of the central Russian common hare, which are very closely 
related toL.e. tesquorum (see below) occasionally occur among 
typical L . e . hybridus. Such is, for instance, specimen No. S. 4976 
(collection of the Moscow Zoological Museum) obtained near the Kudinovo 
Station (near Moscow) by E. P. Spangenberg on September 22, 1922. The 
hair of this specimen lacks the characteristic waviness of hybridus, and 
its color is duller, more earth-colored than that of typical central Russian 
hares. 

There are two pelts in the collection of the Moscow Zoological Museum 
obtained by S. A. Buturlin near Wesenberg [Rakvere] Estonia (former 
Estland). The color of these pelts is very peculiar. 

General color of the back intense clay-rust-red (intermediate between 
tawny and orange -cinnamon, Ridgway, 1912). Flanks, and especially chest, 
with a very prominent reddish brown tinge. General color of chest 
intermediate between orange -cinnamon and cinnamon. Anterior sides of 
forelegs very richly colored. 

The specimens from Wesenberg are also characterized as follows: l) 
underfur pure white; 2) middle part of hair broadly black; 3) apical 
part of hair deeply colored, causing the typical reddish clay tinge. The 
subapical part of the hairs of the back appears intermediate between pinkish 
cinnamon and ochraceous buff under a magnifying glass. The dark color 
of the fur is caused by black hair tips and long guard hair with black apical 
part and without clay-yellowish rust subapical part. 

Specimen No. S.3998 in the collection of the Moscow Zoological Museum 
has no date of capture. It was probably caught in fall before the onset of the 
molt, according to its fur. 

The second specimen (No. S. 4980) was caught near Wesenberg in mid- 
October 1903 (New Style). The lower part of the back is slightly grayer in 
this specimen, indicating the beginning of molt. 

The Wesenberg [Rakvere] hares are systematically closely related to 
L. e. occidentalis de Winton, described from England (Herefordshire). 
A series would have to be compared, to solve the question. Hares living 
along the Baltic Coast are very closely related to, and probably identical 
with, common hares in Great Britain. 

L.e. hybridus is usually considered by furriers to belong to the 
"central" strain. They also ascribe to this strain hares from the Central 
Chernozem Area (L.e. tesquorum). L.e. hybridus penetrates 
the region of the "Menzelinsk Strain" at the eastern boundary of its range. 
Common hares transitional to the well differentiated eastern race (see 
below) occur in this region. 



127 



The Turgai-Orenburg Steppe is populated by the well differentiated 
natio L.e. caspicus natio turgaicus Ogn. (see below). 

GEOGRAPHICAL DISTRIBUTION. This subspecies is widely distributed 

in the central part of eastern Europe. Intermediate forms between L.e. 

europaeus and L.e. hybridus, in which characters of the typical 

subspecies predominate, occur in the west from the vicinity of Novaya 

Aleksandriya [Pulawy] (from where I obtained specimens). The same 

.„ intermediate forms occur in the Kiev District and in Zvenigorodka. L. e. 
1 47 

hybridus with more pronounced characters of the race occurs farther 

east. Hybridus also occurs in northeastern Europe (see below) and in 

the south (about 53-54°N). Hares from near Novosil are intermediate 

between h yb r i du s and t e s quo r u m . Most of the hares from Skopin 

are intermediate between tesquorum and hybridus, with strong 

resemblance to the latter. Hares from Lipetsk and Kozlov definitely are 

L.e. tesquorum. Hares caught near Kuznetsk belong to L . e . 

hybridus but their winter fur is much whiter than that of Moscow 

specimens. 

It is difficult to determine the boundary of the range of L.e. hybridus 

in the central Trans-Volga region. Hares from Kazan and specimens from 

the middle reaches of the Belaya river are apparently an intermediate 

type. They have characters in common with the eastern race L.e. 

caspicus natio turgaicus, namely the markedly white winter fur . 

Hares from the following localities are characterized by intensely white 

winter fur and large size: Vyatka, Kotelnich and Nolinsk districts; Votish 

Region; Vologda Region. In order to determine to which subspecies they 

belong, their summer fur should be compared with that of hybridus and 

their skulls measured. They certainly differ from the central Russian 

subspecies. This question should be further studied. 

329. Lepus (Eulagus) europaeus tesquorum Ogn. (1923). 
Steppe common hare 

1853. Lepus timidus Chernai A. , Fauna Khar'kovskoi gubernii (Fauna of the Kharkov Province), p. 14. 
1871. L [epus] campestris Bogdanov M. N. , Ptitsy i zveri chernozemnoi polosy Povolzh'ya (Birds 

and Mammals of the Chernozem Belt of the Volga Region), p. 175-176. 
1894. Lepus timidus Silant'ev, A. A. , Fauna Padov (Fauna of the Pady'), St. Petersburg, p. 79. 
1910. Lepus europaeus Ognev S. 1. and Gorbachev. S. N. , Mlekopitayushchie Yugo-vostoka Orlovskoi 

gubernii (Mammals of Southeastern Orel Province), p. 28. 
1923. Lepus europaeus Zverezomb-Zubovskii E. V., K poznaniyu fauny mlekopitayushchikh Donskoi oblasti 

(The Mammal Fauna of the Don Region), p. 29. 
1923. Lepus europaeus tesquorum Ognev S. I. , Fauna nazemnykh pozvonochnykh Voronezhskoi 

gubernii (Terrestrial Vertebrates of the Voronezh Province), pp. 115-121. 

1927. Lepus europaeus tesquorum Migulin, A. A. , Obzor gryzunov Ukrainy (Survey of the 
Ukrainian Rodents), Protection of Plants in Ukraine, No. 3-4, p. 73. 

1928. Lepus europaeus Gavrilenko, M., Opyt sistematicheskogo kataloga zverei Poltavshchiny (A 
Systematic Catalogue of the Mammals of the Poltava Region), p. 13. 

TYPE LOCALITY AND DEPOSITION. Type No. M 2986, male, June 17, 1922, Dokuchaev Experimental 
Station, Bobrov County, Voronezh Province, collected by A. I. Yurin. S. I. Ognev's collection. 
MATERIAL EXAMINED: 44 pelts and 60 skulls. 

DIAGNOSIS. Fur on back not wavy. Summer fur dull, brown-yellowish 
gray with an earth shade, often with marked blackness. Chest and lateral 
parts of inguinal region dull, straw-grayish rust colored. Winter fur 



["Pady" - depressions in the steppes of European Russia. ] 

128 



148 



149 



markedly whiter and grayer than summer fur. Earthy rust -brown color 
persisting in winter fur only in middle of back as a round patch, on top 
of head, on ears and neck. Size large: length of skull in full-grown 
specimens usually more than 100 mm; length of hind feet 167- 185 mm. 

Average weight about 6-6,4 kg*. 

ADDITIONAL DESCRIPTION. Anterior part of muzzle dull, straw-gray 
(close to drab, Ridg-way, 1912); a strong admixture of blackish hair on the 
forehead between the eyes, where a dull straw tinge (drab) appears only in 
spots on a black-gray background. 

Eyes surrounded by gray- white ring with a slight straw tinge. Blackish 
hairs mixed with straw-gray ones on the cheeks under the eyes. Hair on 
neck anteriorly dull, earth- straw colored (close to drab but with a slight 
rust tinge). Back earth- gray with a brownish and straw tinge. Hair of 
back without the typical waviness of northern hares. Basal part of guard 
hair whitish gray; subapical parts intensely black; distal parts of guard 
hairs pale straw-gray; tip often blackish. Posterior part of back 
w^ith a slight gray tinge. Flanks dull, straw-earth- gray (intermediate 
between drab and wood-brown) with a slight reddish rust tinge. Lower 
part of throat, chest and sides of inguinal region rust-straw-gray 
(intermediate between wood-brown and avellaneous). Anterior sides of feet 
earth- gray with a straw tinge. Inner margins of ears with blackish hair, 
and with a broad pale straw-earth band on both sides. Insides of ears 
whitish. A distinct oblong blackish spot on outer side of ears. Posterior 
parts of ears gray- straw- whitish; tips black. 

There are many variations of this typical color either towards an even 
duller color, or towards richer and more intense colors. However, the 
color ofL.e. tesquorum is never as rich as that ofL.e. hybridus. 

Winter fur much longer and fluffier than summer fur. The brownish 
earth color persists in winter fur only as a small rounded patch in the 
middle of the back. Flanks, lateral and posterior parts of back white, with 
straw-gray tinge. Whitish gray tones cover the anterior part of the 
muzzle, cheek, throat and chest. Rust -straw colors persist only weakly 
on the chest. 

The color of young L.e. tesquorum differs sharply from that of 
L.e. hybridus. Specimens of this subspecies are more easily 
distinguished when young. 

In young hares about 150 mm in length the head and back are very dark, 
blackish brown with a slight pale straw speckling intermediate between 
avellaneous and pinkish buff. A straw tinge is more pronounced on flanks. 
Chest and neck dull brown-gray except the whitish gray chin. Lighter, pale 
straw colored hair tips are also found in these ares. Belly a dirty whitish 
gray. Many specimens with a small white spot on forehead. Feet with dark 
brown hair, pale str^w at tips. Tail dorsally blackish brown and white 
ventrally. Outsides of ears vs*iitish with a straw tinge. Inner margin of 
ears with dark brown hair with straw tips. Insides of ears whitish straw. 
Inner margin of inner half of each ear with an elongated blackish brown spot. 



According to N. V, Kireevskii ("Forty years of Hunting" (40 let postoyannoi okhoty), Moscow, 1875, p. 145) 
the largest hare of 12,903 common and blue hares killed from 1815-1856 weighed about 20 pounds (i. e. , 
8 kg). This probably applies to L. e . tesquorum, as N. V. Kireevskii hunted chiefly in the Kursk and 
Orel provinces. 



129 



Color of individual hairs on the back as follows: bases of hair brownish 
gray; a dark black-brown band in the middle; subapical part narrow, pale 
straw colored; tip blackish brown. Hair of underfur with brownish gray 
bases and more brownish tips. 

This coloration varies, as the straw color of the hairs is developed to 
a various extent in different specimens. 

In young specimens, half the length of adults, the fur becomes smoother 
and hair straighter than in the soft, fluffy and slightly wavy fur of the very 
young. The straw-yellow subapical parts of the hair on the back become 
better marked and the fur therefore appears lighter. Color of head dark, 
from grayish black-brown to brownish. White spot on forehead usually 
well marked. 

The white area on the chin becomes broader and better marked, and 
covers the upper part of the neck. A broad stripe which varies between 
pale rust- straw to dull rust color (from wood-brown to grayer than 
cinnamon) is clearly marked across chest and neck. Belly white. Inguinal 
region with pale straw-rust tones. 

Measurements of L . europaeus tesquorum: length of skull 
97-108. 5mm (M. 102.8mmO; condylobasal length 87-97. 5mm (M. 92.2mm); 
zygomatic width 46 -54 mm (M. 49.2 mm); length of nasal bones 43.2 - 
50.9 mm (M. 47.8 mm); breadth of nasals 17.2-23.6 mm (M. 20.1 mm); 
length of bony palate 5.6-7.9 mm (M. 6.8 mm); breadth of incisive foramina 
10-15.2 mm (M. 12.2 mm); length of upper tooth row 16.8-20.7 mm 
(M. 18.2mm). 

Length of body with head 595-610 mm; hind feet 150-185 mm; ear 
(measured along concavity) 103-140mm. 

SYSTEMATIC NOTES. Various hunters have written on the differences 
between the steppe common hare and the common hare of the central 
forest belt of the European part of the U.S. S.R. M.N. Bogdanov (187 1 ) 
named the steppe common hare 'Tumac', and wrote: "The Tumac hare 
(L. campestris nob.) is not a hybrid of the common and blue hare, 
but a steppe variety of the common hare (L . t i m i d u s ), differing from 
it in the following characters: l) grayer color; 2) short broad body; 
3) rapid running. The Tumac hare is rare and sporadically distributed 
in stony parts of the steppe. " 

It is, in my opinion, more correct to call the steppe common hare 
L. e. tesquorum for the following reasons: 1 ) Bogdanov wrongly 
named the steppe common hare 'Tumac'; 2) the form has no description, 
as the incorrect indication of its "gray color" can hardly be called a 
description; 3) the geographical range of the Tumac is not given. 
150 Like all subspecies with transgressive characters, the steppe common 
hare is clearly distinguished from its northern relations in series. Some 
specimens of the steppe common hare, especially from places on the border 
of the range ofL.e. hybridus, are almost indistinguishable from L . e . 
hybridus. One of the clearest and most constant differences, besides 
color, is the absence of wavy hair on the back, a typical character of L . e . 
hybridus . 

GEOGRAPHICAL DISTRIBUTION. Russian Chernozem steppes. The 
northern boundary of its range has not yet been defined. This subspecies 
apparently occurs from Nezhin in the east to Novgorod-Seversk, 
Trubchevsk, Orel, Novosil, Skopin, Morshansk, and Penza up to the 
Samara Bend. I have not examined material from the Trans-Volga region. 



130 



I cannot, therefore, state definitely which subspecies occurs in the Samara 
Steppes and on the Obshchii Syrt. L.e. tesquorum probably occurs 
along the Greater and Lesser Irgiz rivers, to judge from material from 
near Volsk. L.e. tesquorum possibly also occurs between Bugulma, 
Buguruslan and Buzuluk, but this region may be inhabited by forms 
intermediate to the eastern race, which become very white in winter. The 
Dnieper is the western boundary of the range of this subspecies. A. A. 
Migulin (1927) was apparently correct in stating that the range of 
tesquorum includes the forest steppe of the left bank Ukraine [territory 
east of the Dnieper]. 

It is difficult to state how far south the range ofL. e. tesquorum 
extends. L. e. transsylvanicus occurs in the southern Dnieper and 
southern Tauric Steppes, but not in the Crimea. N. F. Nikolaev sent me 
a specimen from Berdyansk. The large skull of this specimen suggests 
that it belongs to L. e. tesquorum. However, its summer fur is a 
lighter color than that of the typical tesquorum . Such a hare was sent 
to me by V.V. Raevskii from the southern Don Steppes. The range of 
L.e. tesquorum in the northern Caucasus reaches that of L . e . 
caucasicus. It is difficult, or impossible, to determine the exact 
boundary of these subspecies. At any rate, the Kalmyk, Nogai and Kuban 
hares belong to L. e. caucasicus. 

330. Lepus (Eulagus) europaeus transsylvanicus Matchie 
(1901). Roumanian or Crimean common hare 

1889 Lepus timidus var. (=L. m e di te rraneus) Tikhomirov A A and Korchagin A N , Spiski i 
opisanie predmetov, nakhodyashchikhsya v Zoologicheskom muzee Moskovskogo universiteta (Lists and 
Description of Specimens in the Zoological Museum of the Moscow University), Izvestiya Obshchestva 
lyubitelei estestvoznaniya, antropologii i etnografii, VI (IV), Mlekopitayushchie, p. 31 (from aspecimen 
from the vicinity of Sevastopol). 

1891. Lepus europaeus (Lepus m e dit errane us)Nikol'skii A. N. , Pozvonochnye zhivotnye Kryma 
(Vertebrates of the Crimea), appendix to Vol. XVIIl, "Zapiski Akademii Nauk", No. 4, pp. 77-80 (the 
Crimean common hares belong to the "form" mediterraneus in the author's opinion). 

1901. Lepus transsylvanicus Matschie P. , Rumanische Saugetiere, Sitzb. der Gesellsch. naturf. Freunde 
z. Berlin, S. 236. 

1908. Lepus europaeus transsylvanicus Hilzheimer M. , Die Hasenarten Europas, Jahresb. des 
Vereins fur Vaterl. Naturk. in WUrttemb. , S. 394. 
151 1912. Lepus europaeus transsylvanicus Miller G. S. , Catalogue of Mammals of Western Europe, 
p. 509. 

1916. Lepus (Eulagos) europaeus transsylvanicus Ognev S. I. Mlekopitayushchie Tavricheskoi 
gubernii (Mammals of the Taurian Province), parti, Gryzuny (Rodents), pp. 55-63. 
TYPE LOCALITY AND DEPOSITION. Tasleu, northeastern Roumania. Type in the collection of the 

Berlin Museum. 

MATERIAL EXAMINED: 30 pelts and 32 skulls. 

DIAGNOSIS. Fur not curly. Summer fur yellowish brown-gray with an 
earth-clay tinge, often with a distinct black tinge. 

Chest and sides of inguinal region rich clay-red. 

Winter fur usually darker than summer fur, with more marked blackness. 
Gray color very little marked; gray hair found only in lower part of back, 
and to a small extent on flanks and cheeks. 



131 



152 



Size small: length of skull usually less than 100mm (89.3-101 mm, 
M. 94.2mm); length of hind foot 123- 160 mm (M. 147 mm). Average weight 
3.2kg (2.9kg-3.9kg). 

ADDITIONAL DESCRIPTION. Winter fur. Entire forehead and occiput 
brownish gray, mottled with yellowish straw hair tips. The general color of 
these parts is therefore relatively light. Color of hair as follows: the well 
marked bases of the hair earth- gray-brown, followed by well marked black 
bands; straw-yellow subapical parts; black tips. Upper part of head 
frequently with small patches of white hair. General color of upper part of 
muzzle lighter as a result of the greater admixture of yellowish straw hair 
tips which become white on the ventral side of the muzzle. A diffuse, 
indistinct whitish stripe of hairs with gray basis and whitish apex from the 
eye to the tip of the nose. Ring around the eyes whitish with indistinct 
ocher tinge. A narrow whitish stripe from the eye to the ear. A broad 
rust-brown stripe ventral to the whitish ring around the eye. Cheeks 
grayish white with a slight ocher tinge and an admixture of blackish hair. 
Anterior margin of ears with dense, short hair with marked black bases and 
straw-yellow tips. Outer margin of anterior edge of ears with long, 
straw-whitish hair. Inner side of ears densely covered with short straw- 
whitish hair with a diffuse elongate spot on the outer side. Proximal part 
of outside of ear with dirty-yellow fur; distal part of outside of ear with a 
well marked black spot extending 40mm along the outer edge. The area 
behind the ears covered with reddish rust-brown fur. Back with dense 
fur which is generally rather dark earth-straw with a strong black tinge. 
Color of individual hairs on back as follows: 1) base of guard hair 
blackish; 2) middle part pale straw-rust colored, sometimes more 
whitish; 3) tip blackish. Completely black guard hair occurs sometimes 
among guard hair of the color described. Underfur wavy, whitish. Tips 
of underfur hairs dull blackish brown. 

Many white hairs or white hair with short blackish tips occur on the 
lower part of the back. The admixture of white hair imparts a general 
grayish white color to the lower part of the back. 

Rust-ocher tones well marked on sides of back, best marked on the 
shoulders and the inner side of haunches; the same colors form a broad 
belt across the chest and the lower part of the neck. The long hair in this 
region has a dull rust-red-ocher tinge (intermediate between wood-brown 
and cinnamon). White hairs also occur in these parts. 

Isolated very long hairs with black base and a broad white tip on the 
sides. Anterior sides of forelegs with light ocher-straw-earth colored 
fur. Soles of hind legs whitish, mixed with pale ocher hair. Tail white, 
blackish brown or black dorsally. 

Color of summer fur more rust-colored, lacking black tone typical for 
the winter fur. Top of head with marked rust-straw tones. Gray tones 
absent on cheeks and lower part of back. Color of flanks, a transverse 
stripe on the chest, and legs more saturated clay-rust colored than in 
winter. 

Color of young Crimean hares relatively light, similar to that of Moscow 
hares though perhaps a little lighter, sharply different from the color of 
young L.e.tesquorum. 

Measurements of Lepus europaeus transsylvanicus. Length 
of skull 89.3-101 mm (M. 94.9 mm); condylobasal length 78.2-89 mm 
(M. 83.6mm); zygomatic width 43.2-48.2 mm (M. 46.1 mm); length of 



132 



nasals 37.2-46.5 mm (M. 43.1 mm); width of nasals 16.8-21.1 mm 
(M. 18.6 mm); length of bony palate 5-7.8 mm (M. 6,2 mm); width of incisive 
foramina 9.6-13.3 mm (M. 11.1 mm); length of upper tooth row 15.3-18.1 mm 
(M. 16.9mm). 

Length of body and head 550-600 mm (M. 564mm); hind feet 124-160mm 
(M. 147 mm); ear (measured along concavity) 102-118 mm (M. 107.2 mm). 

SYSTEMATIC NOTES. The Crimean common hare resembles L . e . 
tesquorum externally, but differs from it in the following characters: 
1) smaller size and weight; 2) richer clay- rust color of chest and flanks 
in summer fur; 3) winter fur darker, with black hair more developed 
than in the summer fur, not lighter, more whitish or grayish as in 
tesquorum. 

The Crimean hares of my collection are identical with an Albanian 
specimen sent to me by the British Museum identified as Lepus 
transsylvanicus . 

According to the classification of furriers, the Crimean common hares 
belong partly to the southeastern strain of the common hare. 

GEOGRAPHICAL DISTRIBUTION. Throughout the Balkans, in Roumania, 
Bessarabia (at least in its southern part), between Dubossary, Olgopol 
and Nikopol, and in the southern Tauric Steppes in Crimea. More eastward, 
near Osipenko, [former Berdyansk], occur hares closely resembling 
tesquorum in their skull measurements. These specimens are 
apparently of an intermediate type, and they resemble hares from the 
southern Don Steppes (see above). 

331. Lepus (Eulagus) europaeus caucasicus Ognev (1929). 
Caucasian common hare 

1887. Lepus timidus Rossikov K. N. , Obzor mlekopitayushchikh doliny reki Malki (Survey of the 
Mammals of the Malka River Valley), pp. 74-75. 

1924. Lepus europaeus transsylvanicus Ognev S. I. , Gryzuny Sevemogo Kavkaza (Rodents of the 
Northern Caucasus), pp. 7, 8. 

1925. Lepus europaeus transsylvanicus Berne L, Rezul'taty omitologicheskoi ekspeditsii v 
Kizlyarskom okruge Dagestana (Results of an Ornithological Expedition to the Kizlyar District (Dagestan)), 
p. 24. 

1926. Lepus europaeus transsylvanicus Radishchev A. M. , Materialy k poznaniyu fauny gryzunov 
Kabardino-Balkarskoi Avtonomnoi oblasti (Data on the Rodent Fauna of the Kabardino-Balkarian 
Autonomous Region), "Izvestiya Severokavkazskoi kraevoi stantsii zashchity rastenii", No. 2, p. 84. 

1926. Lepus europaeus tr anssyl vani c us Turov S. S. , Opyt systematicheskogo obzora 

mlekopitayushchikh Osetii (A Systematic Survey of the Mammals of Ossetia), Uchenye zapiski 

Severokavkazvskogo instituta kraevedeniya, I. p. 321. 
1929, Lepus europaeus caucasicus Ognev S. I., Zur Systematik der russischen Hasen, Zoologisch. 

Anzeiger, Bd. 84, H. 1/4, S. 75. 

TYPE LOCALITY AND DEPOSITION. M. 2973 female, 30 November, 1923, vicinity of Vladikavkaz 
[Ordzhonikidze]. Type in S. I. Ognev's collection 

MATERIAL EXAMINED: 38 pelts and 47 skulls. 

DIAGNOSIS. Fur on back not wavy. Summer fur of most specimens 
rather dull, brownish yellow -gray with an earth tinge. Summer fur often 
with slight black tinge, sometimes more clay-yellow with an earthy tinge. 

Clay-red tones on chest and flanks little marked in specimens found in 
summer, replaced by stronger yellow -clay-gray tones. 

Color of winter fur almost like that of summer fur, not darker as in L , e , 
transylvanicus . Only the lower part of back has a gray color , A 



133 



154 



light gray tinge also on the flanks. Whitish gray colors on cheeks and 
sides of muzzle. 

Measurements larger than in L. e. transsylvanicus. Length of 
skull about 100 mm (92-105 mm; M. 99.2 mm); length of hind feet 140- 
167 mm (M. 151.4mm). Average weight 4 kg (3.2-4.4 kg). 

SYSTEMATIC NOTES AND ADDITIONAL DESCRIPTION. The color of 
North Caucasian hares varies little. General tone of summer fur earth- 
colored, only slightly different from that of L . e . tesquorum in its 
richer color and stronger ocher tone on chest and anterior sides of 
forefeet. Color of summer fur intermediate between that of the Crimean 
subspecies and ofL.e. tesquorum. 

Specimens with light clay tones and a slight sandy tinge occur together 
with specimens of ordinary color. Specimen No. M. 2946 in my collection 
(14 October 1923, near Vladikavkaz [Ordzhonikidze]) may be considered 
as representative of this clay-colored form. This specimen was caught 
at the beginning of the molt. 

As mentioned above, the winter fur ofL.e.caucasicus is 
generally almost indistinguishable from the summer fur. The winter fur 
of Caucasian hares is lighter than that of Crimean hares, in which it 
shows a marked blackness on the back. 

During examination of the collection of the Caucasian Museum, my 
attention was drawn to specimens from the Bogos Range. The color of 
their winter fur is transitional to that ofL.e. cyrensis. Specimen 
No. 2-09, 1907 resembles this subspecies particularly closely. 






FIGURES 79-81. Variation of nasals in Le pus europaeus caucasicus Ogn. 79 — Specimen from 
Dagestan, near Karabudakhkent, No. M. 2959, S.I. Ognev's collection. 80 — Specimen from the same 
locality, caught in 1923. No. M. 2972, S.I. Ognev's collection. 81 — Specimen from the same locality, 1923. 
No. M.2969, S. 1. Ognev's collection 

Photograph by S. I. Ognev 



134 



155 



Hares from the Kizlyar Valley and from Kara-Nogai Steppes are much 
lighter in color than L. e. caucasicus. They are transitional to L . e . 
caspicus . 

There are specimens amongthe Kizlyar hares which are indistinguishable 
from L . e . caucasicus from the mountains. The old specimen No. 
M. 2966, caught on 4 June 1923 near Malaya Areshovka, in the Kizlyar 
District belongs to this group. Its color closely resembles that of a hare 
shot in the xerophytic mountain steppes of Dagestan (near the village of 
Karabudakhkent ) . 

Color of summer fur of specimens from the Caucasian Coast of the Black 
Sea (near Beta) differs from that of specimens from the Central Caucasian 
Range in the stronger development of clay tones, especially on chest and 
flanks. The color of the winter fur on specimens from Batum and Sukhumi 
differs markedly from that of typical L.e. caucasicus in the more 
brownish clay tones on the back, the brownish rust-clay transverse stripe 
on the chest, and the similar color of the flanks. 

The winter fur of hares from the Black Sea Coast differs clearly from 
that of L.e. caucasicus, but the summer fur of both is similar. We 
observed the same relationship between L.e. caucasicus and L.e. 
cyrensis . 

I classified this Black Sea hare as a separate natio, L.e. caucasicus 
natio ponticus Ogn. (type No. 4915 female, 29 July 1928, Beta Village, 
Caucasian Black Sea Coast, B. V. Obraztsov, collection of the Moscow 
State Zoological Museum) (Zoologischer Anzeiger, Bd. 84, S. 75). 

Measurements of L e p u s europaeus caucasicus: Length of skull 
92-105 mm (M.99.2); condylobasal length 83-94.2 mm (M. 88.2 mm); 
zygomatic width 43. 6-49. 2mm (M. 46.6mm); length of nasals 41. 1-48. 8mm 
(M. 45.7mm); width of nasals 16.3-22.2 mm (M. 19.4 mm); length of bony 
palate 5-7.7 mm (M. 6.2 mm); width of incisive foramina 10- 12.6 mm 
(M. 11.3mm); length of upper tooth row 15. 2-18. 9mm (M. 16.9mm). 

Length of body with head 560-680 mm; hind feet 140- 167 mm 
(M. 151.4mm); ear (measured along concavity) 104-115mm (M. 108mm). 

GEOGRAPHICAL DISTRIBUTION. This subspecies occurs on the 
Greater Caucasus and in the adjacent northern areas. The northern 
boundary of the range is not exactly known. It probably runs from the 
eastern coast of the Sea of Azov through Stavropol - Mozdok - Makhachkala 
(Port Petrovsk). The southern boundary is also yet to be determined. It 
runs approximately along 42° N. Specimens intermediate between L.e. 
caucasicus and L.e. cyrensis (see above) occur in Dagestan on the 
Bogos Range. In the Central Caucasus, the boundary betweenthe ranges of 
these two forms passes along the Dushet-Kutaisi line. It is not known how 
far south along the Black Sea Coast the range ofL. e. caucasicus 
extends. Natio ponticus of this subspecies occurs south of Batum. As 
to the northern range of L.e. caucasicus, hares from the Kizlyar 
Valley are to some extent intermediate between L.e. caspicus and 
L.e. caucasicus. 

332. Lepus (Eulagus) europaeus cyrensis Satun. (1905). 
Transcaucasian common hare 

1832. Lepus timidus M^netries E. , Catalogue Raisonn^ des Objets de Zoologie, p. 23 (partim.'). 
1876. Lepus caspicus Blanford W. T. , Eastern Persia, II, Zoology, pp. 82-83 (Teheran - Isfahan). 



135 



1896. Lepus europaeus var. Satunin K. , Voriaufige Mitteilvmgen fiber die Saugethierfauna der 

Kaukasusiander, Zoolog. JahrbUcher, Bd. 9, S. 308. 
1903. Lepus sp. Satunin K. A. , Obzor issledovaniya mlekopitayushchikh Kavkazskogo kraya (A Study of 

the Mammals of the Caucasus), Zapadno-Kavkazskoe otdelenie Geograficheskogo obshchestva, XXIV, p. 61. 
1905. Lepus cyrensis Satunin K. A., Novye i naaloizvestnye mlekopitayushchie Kavkaza i Zakaspiiskoi 

oblasti (New and Little Known Mammals of the Caucasus and the Transcaspian Region), Izvestiya 

Kavkazskogo muzeya, II, pp, 60-63. 
1913. Lepus eyre ns is Satunin, K. A. , Fauna Chemomorskogo poberezh'ya (Fauna of the Black Sea Coast), 

p. 41, 
1918, Lepus cyrensis Kollmann M. , Note sur les mammiferes rapport^s d'Asie Mineure par M. Gadeau de 

Kerville, Bull. Mus. , Paris, pp. 201-204. (Asian Minor). 
1929. Lepus europaeus cyrensis Ognev S. I. , Zur Systematik der russischen Hasen, Zoologisch. 

Anzeiger, Bd. 84, H. 1/4, S. 76. 
TYPE LOCALITY AND DEPOSITION. No. 129 1/k male, December 1904*, post office of the Barda 
Dzhevanishir County of the Qisavetpol (Kirovabad] Province. Deposited in the Georgian Museum (former 
Caucasian Museum). 

MATERL^L EXAMINED: 54 skulls and pelts. 

DIAGNOSIS. Fur straight, neither wavy nor curly. Color of summer 
fur rather dull, yellowish brown- gray with an earth tinge. 

Color of chest and flanks dull, brownish straw (intermediate between 
avellaneous and pinkish cinnamon), usually without rust tones. Winter 
fur differs clearly from summer fur in its general paleness and straw- 
rust-gray tones with a typical vinaceous tinge (vinaceous-buff and 
avellaneous). Slight gray tones on flanks, lower part of back near tail, 
and on cheeks. 




.O^ 



FIGURES 82, 83. Transcaucasian common hare, Lepus europaeus cyrensis Satun. According to type 
No, 129 1 Jan, female, Barda, Dzhevanshir County, collection of the Georgian Museum 

Photograph by S. 1. Ognev 



* The date September 1905 is given in K. A. Satunin's work (loc. cit. , 1905). The number of the specimen 
"129, k" is the same. 



136 



157 



Length of skull usually less than 100mm (91-103 mm; M. 96.6mm); 
length of hind feet 144-150mm. 

ADDITIONAL DESCRIPTION. Color of summer fur of L. e . cyrensis 
very similar to that ofL. e. caucasicus. Ground color of back of 
Transcaucasian common hare slightly lighter. Color of flanks and 
transverse stripe of chest duller than in L.e. caucasicus in most 
specimens of L.e. cyrensis. This difference is clearly seen in series . 
However, there are specimens of L.e. cyrensis in which the color 
is practically identical with that of L.e. caucasicus. 

Winter fur of L.e. cyrensis much paler than that of caucasicus. 
Pale, straw-rust-gray tone with a typical vinaceous tinge, close to 
vinaceous-buff and avellaneous, intermediate between avellaneous and 
pinkish cinnamon, predominates in its color. Color of transverse stripe on 
chest dull straw-brown. 

Anterior sides of forelegs and flanks of Transcaucasian hare paler than 
those of caucasicus, without the rust tinge of the latter. Anterior side 
of feet especially dull, being whitish straw-gray with a very slight, hardly 
perceptible rust tinge. 

Thus, the top of the head of L.e. cyrensis in winter fur is lighter, 
paler and grayer than in caucasicus. Cheeks paler, pale straw-gray 
with a vinaceous tinge (intermediate between vinaceous and light pinkish 
cinnamon). 

Color of young specimens pale yellowish straw- sand colored with earth- 
rust -brown mottling. Color of transverse stripe on chest and of flanks 
yellowish straw- gray. General color of fur of young c yr e n s i s similar 
to that of young L.e. hybridus and L. e. transsylvanicus. 

SYSTEMATIC NOTES. The Transcaucasian common hare is very 
closely related to L . e . caucasicus. It differs from it in its smaller 
size, paler color of winter fur, and some variable color characters of the 
summer fur. 

An excellent series obtained in winter 1932 (collected by Yu. A. Isakov) 
in the vicinity of Kalinovka, 35 km south of Lenkoran, is kept in the 
Moscow State Zoological Museum. The small Lenkoran hares are 
identical in their skull structure with L.e. cyrensis Satun. 
(specimens from Yerevan*). 

Length of skull of Lenkoran hares varies between 93-97 mm; 
condylobasal length 81-86 mm; zygomatic width 43.5-48mm; length of 
nasals 41-47 mm; width of skull at level of auditory meatus 34.2-36.3 mm; 
length of row of upper molars 15.3-17.2 mm. 

Measurements of body small, generally like those of L.e. cyrensis: 
length of body with head 490-563 mm; hind feet 130- 150 mm; ear 110- 
115mm. Weight 2. 5-3. 3kg. 

The color of winter fur differs sharply from that of specimens caught 
near Yerevan and along the Kura river. Lenkoran hares are much darker 
because of the very strong development of black hair tips on the back. The 
dark color on the back forms a wavy, black pattern varied by the clay- 
yellow tone of the subapical parts of the hairs. In addition to hair with 
black tips, the color of the winter fur on the back is greatly influenced by a 



* The skulls of the Lenkoran hares in this series are smaller than those of the Yerevan hares, and the rostra 
and nasals are narrower. 



137 



158 



large number of black guard hairs. The lower part of the back almost 
never becomes gray in winter fur. Top of head with a marked black tinge, 
speckled with a lighter yellowish clay tone. Anterior and lateral parts 
of muzzle with a grayish tinge. A marked rust -brown spot under the eye. 
Cheeks rust-gray. Anterior margins of inner side of ears with a marked 
blackish gray tinge. Edges of anterior parts of inner side of ears whitish 
yellow. Inner sides of ears whitish gray with an admixture of blackish 
brown hair along the margins, forming a broad, long marginal patch with 
a yellowish edge. Outer parts of ears whitish gray. Tips of ears with 
black fringes especially well marked on posterior side of ears. Flanks 
with a strong reddish rust tinge. An intense reddish rust (pinkish 
cinnamon) transverse stripe on the chest strongly darkened by black-gray 
hair tips in many specimens. In specimen No. S. 7120 (female), 29 January 
1932, Kizyl-Agach (35 km south of Lenkoran, Yu. A. Isakov), the color of the 
the entire belly is reddish rust-white (light vinaceous-cinnamon). Belly 
pure white in all others. 

The Lenkoran hare can certainly be considered as a separate natio. 
We designate specimen No. 8.7119^ female, 24 January 1932, of Kizyl- 
Agach, 35 km south of Lenkoran, as holotype for L.e. cyrensis natio 
lencoranicus nova. We also examined 18 paratypes. 

Not all specimens of the Lenkoran series are identical. The color 
of most is dark, but some of them are transitional to the lighter form 
L.e. cyrensis which occur farther northwest . The specimen obtained 
by A.N. Karamzin on 20 November 1903, 20km north of Dzhevat is 
intermediate between the Yerevan and Lenkoran hares. 

The collection of the Moscow State Zoological Museum includes a stuffed 
hare caught in winter in the Mugan Steppe. The light color of this specimen 
is identical with that of the Yerevan specimens. 

It would be very interesting to establish, from adequate material, the 
type of color of hares in northwestern Iran. We have no material from this 
interesting region at present. 

GEOGRAPHICAL DISTRIBUTION. Caucasia, south of 42°N Asia Minor, 
northwest Iran (Teheran, perhaps Isfahan). 

333. Lepus (Eulagus) europaeus caspicus Ehrenberg (1830). 
Caspian common hare 

1830. Lepus caspicus Ehrenberg Chr. G. , Symbolae Physicae seu icones et descriptiones Mammalium. 

Decas secunda (without pagination). 
1875. Lepus caspicus Karelin G. S. , Razbor stat'i Ryabinina i. t. d. (A Critique of Ryabinin's Article, etc.), 

Trudy Sankt-Peterburgskogo obshchestva estestvoispytatelei, p. 78. 
1908. Lepus medius caspicus Hihheimer M. , Die Hasenarten Europas, Jahresber. d. Vereins fUr 

Vaterl. Naturk. in Wurttemb. , S. 388, Taf. V, Fig. 4. 
1910. Lepus medius caspicus Trouessart E. L. , Faune des Mammlferes d'Europe, p. 217. 
1916. Lepus europaeus caspicus Ognev S. I. , Mlekopitayushchie Tavricheskoi gubernii (Mammals 

of the Taurian Province), parti, Gryzuny (Rodents), pp. 58-59, pi. IV, p. 1. 

1927. Lepus europaeus caspicus Orlov E. I. and B. K. Fenyuk, Materialy k poznaniyu fauny 
nazemnykh pozvonochnykh primorskoi polosy Kalmytskoi oblasti (Material on the Fauna of Land 
Vertebrates of the Maritime Belt of the Kalmyk Region), p. 83. 

1928. Lepus europaeus Kuznetsov, B. A. , Zametki po faune mlekopityushchikh Ural'skoi gubernii 
(Notes on the Mammalian Fauna of the Uralsk Province), Trudy po lesnomu opytnomu delu, Tsentral'naya 
lesnaya opytnaya stantsiya, issue I, pp. 58-59. 

1929. L [epus] e[uropaeus] caspicus Ognev S. I. , Zur Systematik der russischen Hasen, Zoolog. 
Anzeiger, Bd. 84, S. 76-77. 



138 



159 TYPE LOCALITY AND DEPOSITION. Vicinity of Astrakhan. Holotype apparently lost. 
MATERIAL EXAMINED: 30 pelts and 33 skulls. 

DIAGNOSIS. Fur on back neither wavy nor curly but straight. 

Summer fur very light, rust-clay colored with a sandy-yellow tinge. 

Clay-yellow tones on chest and flanks well pronounced in summer fur. 
The same tones, duller with pinkish and white tinges in winter fur. 

The winter fur differs little from the summer fur in specimens from the 
southern parts of the range of the Caspian common hare. In specimens from 
the northern border of the range, the winter fur is lighter and whiter. 

In specimens of caspicus from Astrakhan, the winter fur is duller 
but occasionally even more intensely colored than the summer fur, with 
a less marked rust tinge. 

As already stated, the winter fur of specimens from the southern parts 
of the range is only slightly grayer than the summer fur and this grayer 
color appears only in the lower part the back. The fur does not become 
whiter in Astrakhan specimens. 

Length of skull 95.3-103.1 mm (M. 100mm); length of hind feet 137- 
153 mm (M. 152.2 mm). Weight 3.2-4. kg. 

ADDITIONAL DESCRIPTION. Winter fur. Entire forehead and 
occiput of Caspian common hare clay-yellow. Color of individual hairs from 
these parts as follows: l) base of hair with broad gray-earth-yellow 
band; 2) a narrow black medial band; 3) broad straw-yellow tip. A 
well-marked black apex in some hairs. A few completely black hairs 
among the hairs described above. Color of anterior-dorsal part of muzzle 
much richer and more yellow. A broad straw- white stripe from the 
anterior margin of the eye to the nose. Ring around the eye whitish ocher. 
A stripe of whitish hair from the eye to the ear. Cheeks light, clay-yellow, 
sometimes with a marked whitish tinge. 

Anterior margin of ears with short hair which has marked black roots 
and clay-yellow tips. Upper fringe of anterior margin with longer, yellow- 
white fur. Inner side with dense clay-white hair, with an indistinct dark 
spot on the side. Outer side with whitish fur. Only the proximal part of the 
outer side with a clay tone. Tip of ear with a black fringe about 35mm long. 

The whole area behind the ears, neck and occiput with a reddish pink 
dull fur, generally of fawn color. Fur on back dense, even, generally of 
light yellowish clay color. Posterior part of back with many whitish gray 
hairs. Flanks with rich yellowish clay-red fur (cinnamon) which is 
strongly marked on the haunches, shoulders and on the transverse white 
stripe of the chest. Anterior and posterior sides of forelegs with rich 
yellow-clay-colored hair. Hind legs with light yellowish straw fur. 

160 Chin, upper part of throat, all underparts (except the chest), belly and 
groin with long white fur. 

Summer fur, except for the underfur, of a rich rust-clay tone with a 
general pinkish cinnamon tinge on neck, flanks and chest (Ridgway, 
pi. XXIX). 

Color of head and cheeks with marked rust -clay tones. Summer fur of 
Caspian hares generally not darker, often lighter and rustier than winter 
fur. 

Young Caspian hares have a duller and paler yellowish gray-rust color. 
Neck with duller, vinaceous-yellow-rust tones (related to avellaneous, 
Ridgway, 1912). Flanks more rust-colored, internnediate between 
avellaneous and pinkish cinnamon. 

139 



161 



SYSTEMATIC NOTES. The Caspian common hare is a distinct 
geographical form. Its color is generally pale, in adaptation to its habitat, 
the semidesert. 

In the northern outskirts of its range, for instance in the vicinity of 
Uralsk and Temir, the hare becomes markedly lighter towards winter, and 
transitional to the more northern and eastern natio turgaicus . 

Specimens from the Kalmyk steppes differ in some characters from the 
type. The summer fur of Kalmyk common hares is not so pure ocher-clay- 
yellow. It is mixed with gray tones in many specimens, sometimes also 
with more intense brownish clay tones. Flanks and the stripe across the 
chest in Kalmyk hares have a more intense clay-rust color (cinnamon). 

The clay-yellow color of the winter fur of Kalmyk hares is less intense; 
the flanks and the stripe across the chest are darker, rust-red clay; the 
color of the stripe on the chest is intermediate between wood-brown and 
cinnamon. 

I described this color variation under the name Lepus europaeus 
caspicus natio k a Im y ko r u m ("Zoologischer Anzeiger", 1929, Bd. 
84, S. 77). The type locality is Ikitsokhura, Akima sands, Kalmyk Region, 
(E.I.Orlov). We also examined 10 paratypes from the type locality. 

Hares from the Kizlyar valley are closely related to L . e . c . natio 
kalmykorum, but paler . Kizlyar hares are lighter than L . e . 
caucasicus in series. They are intermediate between the Caucasian 
and Kalmyk hares. They are closer to the Caucasian form, but not 
identical with it. It is difficult to name this form, and I therefore only 
give its typical characters. 

Hares in the Turgai-Orenburg steppe are a well characterized natio of 
the Caspian common hare. 

Their summer fur is yellow, not rust-clay-yellow, on flanks and chest. 
The winter fur becomes almost completely white. Hairs with blackish 
gray roots, pale rust middle parts, and white tips are present only in the 
middle of the back in a small, oval spot which is made indistinct by the 
white tips of the guard hairs. The feet become completely white. A 
rust-ocher shade persists weakly only on the anterior side of the forelegs, 
and also at the roots of hairs on the flanks and chest. The white hairtips on 
the chest are so intensely developed that they give a general white color to 
this area. The transverse rust stripe on the chest disappears completely. 
Hind feet completely white. A rust-yellow tinge is retained on the top of 
the head and on the ears, especially on their inner margins. 

The Turgai common hare constitutes a well differentiated natio, L . e . 
caspicus natio turgaicus nova. I designate the specimen caught 
by A.M. Korsakov on 15 October 1934 in the Sypsyn Forest (Naurzum 
Reserve), south of Kustanai, asholotype. We exannined 5 specimens. 

It would be important to establish the range of variation of the 
measurements of the Turgai hare from adequate material. The labels of 
the pelts have the following data: length of body with head 525, 550, 580 mm; 
hind feet 150 mm; ear 120, 128,135mm. Weight 3.855-4 kg. 

Length of skull 102.5mm; condylobasal length 90mm; zygomatic width 
50 mm; length of upper tooth row 18 mm. 

GEOGRAPHICAL DISTRIBUTION. Astrakhan and Ural steppes. 
Northern boundary of range of type subspecies not certain. In the Astrakhan 
steppe, the boundary apparently follows 49.5°N rising on the Ural up to 
Uralsk. It turns to 49° N in the east reaching the vicinity of Temir. It 



140 



occurs along the Emba river farther south, in the region of the Bol'shie 
Barsuki, in the Kara-Kum as far as Kazalinsk, Chelkar and Kara-Uzyak. 

Lepus e.c. natio kalm y ko r um occurs in the Kalmyk steppes. 
It is not known how far north it occurs. 

The exact boundary of the range of L . e . c . natio turgaicusis 
also not certain, but it occurs north of Uralsk. Specimens from near 
Uralsk are apparently intermediate between the true caspicus and its 
northeastern natio. 

L . e . c . natio turgaicus occurs in the Orenburg, Orsk, and 
Kustanai steppes. 

In the north of the Nizhni Novgorod Territory, in the Vologda District, 
Votskii Region, Bashkiria, a form occurs which rarely becomes white in 
winter. The relationship between this form and L. e . c . natio turgaicus 
should be determined. The furriers distinguish a very large Menzelinsk 
strain in this area. To judge by our limited material, the Turgai hare is 
not very large. It is closely related to the Caspian common hare in this 
respect. The east Vologda hares probably form a separate natio of the 
subspecies hybridus . 

The northwestern boundary of L . e . c . natio turgaicus is thus, 
so far, unknown. 

158. Lepus (Eulagus) tolai tolai Pall. (1778). Tolai hare 

1778. Lepus tolai Pallas P. S. , Novae species Quadrupedum e Glirium Ordine, pp. 17-28. 
262 1811. Lepus tolai Pallas P.S., Zoographia Rosso- Asiatica, p. 149. 

1859. Lepus tolai Maak R. , Puteshestvie na Amur (A Journey to the Amur), Sankt Peterburg, p. 108. 

1862. Lepus tolai Radde G. , Reisen im Siiden von Ost-Sibirien, S. 211-215. 

1910. Lepus tol ai Kashchenko N. F. O kollektsii mlekopitayush chikh iz Zabaikal'ya (A Collection of 

Mammals from Transbaikalia), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, p. 272:273; 

KashtchenkoN. F. , Novye issledovaniya po mammologii Zabaikal'ya (New Studies of the Mammals 

of Transbaikalia), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, XVII, pp. 396-397. 
1927. Lepus tolai tolai Allen Gl. , Lagomorphs Collected by the Asiatic Exped. , Americ. Mus. 

Novit., No. 284, pp. 6-7. 
1929. Lepus tolai Formozov A. N. , Mlekopitayushchie severnoi Mongolii (Mammals of Northern 

Mongolia), Izdatel'stvo Akademii Nauk, pp. 95-96. 

1929. Lepus tolai Kuznetsov B. A. , Zverovye promysly vostochnogo Zabaikal'ya (Animal Trade in 
Eastern Transbaikalia), Trudy po lesnomu opytnomu delu, Tsentral'naya lesnaya opytnaya stajntsiya. 
Issue VI, p. 93. 

1930. Lepus tolai Vinogradov B. S. , Rukovodstvo k opredeleniyu gryzunov Srednei Azii (Key to Rodents 
of Central Asia), p. 50 (partim.'). (The author considers the tolai hare as identical with the desert hare,' ). 

1933. Lepus tolai tolai Lukashkin A. S. , Zveri stepnoi Bargi v ikh biologicheskikh osobennostyakh 

(Biology of the Mammals of the Barga Steppe), p. 78. 

NAMES: The Tolai Hare (English); Tolai Hase (German). 

LOCAL NAIVlES: "Kamennyi zayats" (stone hare), tolai, tolui, tulai (Transbaikalia); ranchvo (Tangut); 
tun-tu-ula (according to A.Cherkasov, in Transbaikalia). 

TYPE LOCALITY AND DEPOSITION. The valley of the Selenga river should be considered as type 
locality of the Tolai hare. At the beginning of the description of Lepus tolai, Pallas ("Novae Species 
Qudrupedum e Clirium Ordine", 1778, p. 18) writes: "Caeterum in montibus apricis campisque rupestribus 
vel arenosis circa Selengam ubi Lepus rupestris (Kamennoi Saiz) inter Russos audit, uti per omnem quoque 
Mongolian atque Davurlam, husquam deest, frequentior ibi Lepori variabili; ad Septentrionem contra 
Baikalis non observabilis". 

Glover Allen (loc. cit, 1927) considers the Gobi hare (described by K. A. Satunin) as a synonym of the 
Tolai hare, and states that the Gobi Desert should be considered as terra typica of the Tolai hare. 

This is unacceptable. The first exact geographical reference is usually considered as terra typica. As 
mentioned above, Pallas indicates the Selenga Valley (p. 18). He mentiones in the same book (p. 20)- that 
the Tolai hare definitely occurs in the Great Gobi Desert as far as Tibet ("Certe in deserto magno GobeSnsi 
ubique ad Tybetum usque vivit. . . "). 

141 



163 



These supplementary data at the end of the description of the Tolai hare cannot be considered as 
indication of its terra typica. 

Principal figures: l)SchreberJ. Ch. , Die SsLugetiere, IV, 1792, S. 878-881, Tab. CCXXXIV (color 
plate of entire animal); 2) Dvigubskii I, Opyt estestvennoi istorii vsekh zhivotnykh Rossiiskoi imperii 
(Natural History of the Animals of the Russian Empire), 1829, p. 43, pi. 25 (entire animal); 3) Simashko, Yu. 
Russkaya fauna (Russian Fauna), II, 1851, p. 778, pi. 61, fig. 1 (color plate of entire animal); 4) Radde G. , 
Reisen im Siiden von Ost-Sibirien, 1862, S. 211-215, Taf.VlIl, Fig. 2a-c (black pattern of tips of ears); 
5) Brehm, Tierleben. Saugetiere, II, 1914, Taf. Ill, Fig. 1 (photograph of entire animal). 

MATERIAL EXAMINED: 15 pelts and skulls. 

DIAGNOSIS. Head relatively large. Ear when stretched (measured 
from occiput) 1 l/5-l 1/3 as long as head, i.e., much shorter than ear 
of desert hare. 

Tail, including terminal hair, about 85-95% of length of hind foot 
(without bones), i.e., relatively long. 

Tail in summer and winter fur white, with contrasting black or black- 
brown dorsal side. 

The fur does not become much whiter in winter, but is much fluffier, 
denser and softer than summer fur. Color of winter fur varies. In some 
specimens it is yellower than the summer fur, and the lower part of the 
back is much grayer. In other specimens, the gray color covers a larger 
part of the back. 

Winter fur very soft, silky; guard hairs rather thin, straight. Summer 
fur much coarser than in desert hares, but more delicate than in common 
hares. 

Curliness not observed. 

Teats 6. 

Length of nasals (measured along line from inner anterior margin to 
outer posterior margin) either slightly longer, or as long as, frontal 
suture. Maximal width of both nasals (measured posteriorly) usually 
much exceeding length of the postorbital process. 

The lateral contours of the posterior third of the incisive foramina are 
usually markedly widened in Transbaikal Tolai hares, often with slightly 
curved lateral projections; posterior third of incisive foramina of 
Manchurian Tolai hares only slightly widened. 

The lower anterior molar is sharply inclined backwards in profile. 

Mandibular condyle compressed, moderately inclined backwards. 

Lower posterior plate of mandible (from angular process to notch at 
beginning of horizontal process) broad. 

Distance between angular process to notch at beginning of horizontal 
mandibular process much greater than that between anterior alveolus of 
first molar and tip of lower incisor. 

Bullae osseae large, markedly inflated. Bullae projecting anteriorly in 
profile, rounded. 

Length of skull 87-95.2 mm (M. 90.6 mm); zygomatic width 42.3 - 
47.8 mm (M. 44.6 mm); length of nasals 36-41 mm (M.38mm); width of 
nasals 15. 3-19. 3mm (M.17mm). 

Length of body with head 430-580 mm (M. 492 mm); hind feet 110-135 mm 
(M. 123.6mm); ear (along concavity) 85-115mm (M. 94.9mm); tail (without 
terminal hair) 75-llOmm (M. 92.6 mm). 

ADDITIONAL DESCRIPTION. The skull of the Transbaikal Tolai hare 
closely resembles that of the common hare. It is a transitional form 
between the desert hare and the common hare, but it has some special 
characters. The skull of the Transbaikal Tolai hare differs from that of 



142 



the desert hare as follows: l) larger size; 2) greater thickness and width 
(especially at the base of the nasals, which in most Tolai hares are broader 
than in desert hares); 3) ratio of length of nasals to that of frontal suture 
usually different from that of the desert hare; 4) shorter bony palate; 
5) broader incisive foramina (Figures 84-87). 

The Transbaikal Tolai hare resembles the common hare in these 
characters. A further resemblance between them is that, in both species, 
the lateral contours of the posterior third of the incisive foramina have 
rounded protuberances. 



164 




FIGURE 84. Tolai hare, Lepus tolai tolai 
Pall. No. M.2323, 16 January 1928. Vicinity of 
Troitskosavsk [Kyakhta] , P. S. Mikhno, S.I, 
Ognev's collection, skull dorsal 



FIGURE 85. Tolai hare, Lepus tolai tolai 
Pall. , skull ventral 



16 5 Rows of molars of Tolai hare longer than those of desert hare; diastema 
longer. 

In addition to the above characters which resemble those of the common 
hare, the skull of the Tolai hare has some specific characters. Posterior 
plate of mandible large (as in the blue hare); mandibular condyle more 



143 



massive than in the desert hare or even the common hare; condyle 
resembling that of the blue hare to some extent; the posterior part of the 
zygomatic bone which delimits the orbit ventrally, is obliquely elevated, 
the orbit thus becomes narrower than in the blue hare. 



164 




FIGURE 86. Tolai hare. Lepus tolai tolai Pall., skull lateral 

165 COLOR. Summer fur. General color of back grayish straw-earth with 
a slight rust tinge. A yellowish straw -clay tone on flanks. 




nCURE 87. 



Mandible of L e p u s tolai tolai, specimen No. M. 2323 

Drawing by Yu. A. Kostylev 



Color of top of head similar to that of back but occasionally darker 
because of the stronger admixture of black hairs. Color of anterior part 
of head (near muzzle), cheeks, and sides of neck dull, gray-brown-straw- 
colored (intermediate between avellaneous and light pinkish cinnamon). 

Neck dull, straw-rust with a vinaceous tinge (intermediate between 
avellaneous -vinaceous -buff and pinkish cinnamon). 

Chin and upper part of throat white. 

A straw-gray stripe of varying intensity across the chest. Its color is 
either; l) a straw tinge mixed with gray (avellaneous); 2) a stronger rusty 



144 



tinge (light cinnamon); or 3) a well marked gray shade (intermediate 
between mouse-gray and light mouse-gray), because of the gray hair roots 
in this region. 

Anterior margin of inner side of ears brownish earth-colored with a 
fine speckling of straw color. 
166 A broad whitish straw stripe along the outer side of the anterior inner 
margin. Inner side whitish with a slight gray tone. A pale straw-rust- 
colored fringe on the distal part of the inner margin. Outer surface 
whitish with a marked grayish or straw tinge. 

Anterior sides of forelegs dull straw-yellow with a slight clay tinge. 
The relative purity of this tone is changed in some specimens by the 
grayish brown hair tips, and in some specimens by some grayish brown 
guard hairs. 

Soles of hind feet white with a dirty straw-gray tinge. 




FIGURE 88. Tolai hare, Lepus tolai tolaiPall. 

Drawing by A, N. Kamarov 

The general color of the Tolai hare resembles that of several southern 
forms of the common hare, e.g., the Caucasian common hare. There are 
also rare specimens with a lighter grayish color. To this group belongs, 
for instance, specimen No. M. 3311, 1 July 1927, collected between Lake 
Tsagangol and Lake Barun-Tsaidan, Transbaikal, S.I. Ognev's collection. 

The general tone of the back of this specimen is dull, grayish straw with 
well marked black speckling especially in the middle of the back, where the 
light hair tips are worn off. Color of top of head almost like that of back. 



145 



Nose and cheeks grayer and lighter (intermediate between light drab and 
vinaceous-buff). 

Color of neck dull straw-gray with a slight rust tinge. 

Anterior inner parts of ear dull straw- gray with black speckling. A 
broad white stripe along the outer edge of the anterior inner margin. Inner 
sides whitish with a marked gray tone. A whitish stripe along the outer 
margin. Outer side dirty-whitish gray. Tips of ears with a narrow blackish 
fringe. 
167 Color of anterior side of forelegs straw-gray with a rust tone. Hind legs 
whitish with a straw tinge on the dorsal part of the feet. 

Winter fur. Ground color of back pale, straw-yellow with a dark 
rust-vinaceous tinge intermediate between avellaneous and light pinkish 
cinnamon, strongly mottled in a blackish-brown pattern. 

Color of top of head similar to that of back. Color of sides of head 
without the rust tones characteristic of common hares. Ground color dull 
straw-gray with a vinaceous tinge (intermediate between vinaceous-buff 
and light pinkish cinnamon). This ground color is mottled with a 
characteristic blackish pattern which is especially marked on the cheeks 
under the eyes and in the postorbital region. Streaks of blackish hair on 
anterior part of muzzle and near the roots of vibrissae. 

Neck dull, straw-rust-vinaceous. Color of roots of hair intermediate 
between avellaneous and pinkish cinnamon, but hair appears paler because 
of the whitish hair tips. 

Chin and throat white with a light straw tinge. 

Color of stripe across chest pale straw-yellow (intermediate between 
light pinkish cinnamon, vinaceous-buff and pinkish cinnamon). 

Anterior inner margin of ear of blackish color overshadowing the 
straw ground color. Outer fringe of inner margin pale straw. Inner side 
covered with white hair with a rich straw tinge, especially marked on the 
tips. Color of inner margins of posterior side straw-rust colored (light 
pinkish cinnamon). A large elongate white spot on the outer margin of the 
outer side. Tip of ear with a black stripe. 

Color of forelegs pale straw-rust. Belly and groin pure white. 
The pale straw-rust color of the flanks also enters the inguinal region, 
forming two irregular, indistinct stripes. 

There is a very light winter specimen of the Tolai hare in my collection 
(No. M. 3310 male, 19 January 1928, vicinity of Troitskosavsk [Kyakhta]). 
General color of this specimen whitish gray with a slight straw cast and a 
blackish tinge in the middle of the back, which is caused by black guard 
hairs and the blackish tips of some hairs. Flanks whitish with slight gray 
tinge. Head whitish straw-gray with more grayish tinge on the cheeks. 
Straw-gray stripe hardly perceptible across the chest. Whole ventral part 
(except the chest) white with a slight straw tinge. 

Specimen No. M. 3772 female ad. 30 November 1929, vicinity of village 
Mirolyubovka, left bank of Irtysh (Samarskoye District) is also very light 
colored. The general color of the back of this specimen is whitish with a 
rust-straw tinge (pinkish cinnamon) caused by pale straw-rust subapical 
parts of the guard hair, which is mixed with some completely black or 
brownish black hair. Anterior part of head, forehead and cheeks white. 
Flanks whitish with a slight straw-rust tinge. Straw-rust stripe across 
chest indistinct, almost invisible on the generally whitish background. 
Belly pure white. Yellowish straw ears with a strong white tinge. 



146 



168 



Color of summer fur of young dull gray with a straw tinge and blackish 
brown speckling. Color of the stripe across the chest dull straw-gray 
(intermediate between light-drab and vinaceous-buff). The color of the 
Tolai hare is grayer, without rust tones, compared with young specimens 
of Central Russian common hares of the same age. Fur of young L . 
tibetanus lehmanni much lighter than that of Tolai hare, with 
predominating straw-gray tones. 

SYSTEMATIC NOTES. I stated in the article: "Zur Systematik der 
russischen Hasen" that the Tolai hare should be considered as a subspecies 
of the common hare. 

Having examined extensive material, I now consider the Tolai hare as 
a separate species. It is true that it has much in common with the small 
subspecies of the common hare such as L. e. cyrensis and L . e . 
caspicus . However, the following characters justify the consideration 
ofL. tolai as a species closely related to the common hare, but not 
as a subspecies: 1) some cranial differences; 2) the soft fur; 3) the 
relative shortness of the ears, etc. 

It does not seem justified to include the Tolai hare in the group of 
Central Asian hares. The Tolai hare shows clear, constant cranial 
differences. It seems to constitute a transition from tibetanus to 
europaeus in some respects, but it also has special morphological 
characters such as the form of the orbit, the structure of the mandibular 
condyle etc. (see above). 

As stated above. Glover Allen (1927) considers L. gobicus as 
identical with the Tolai hare, and makes it a synonym of this (see above). 

However, as the terra typica of the Tolai hare is not the Gobi Desert, 
but the valley of the Selenga river, L. gobicus cannot be synonymous 
with the Tolai hare. L. gobicus is closely related to the Tolai hare. 
The winter fur ofL. gobicus is characterized by a clay color which 
differs from that of the winter fur of the Tolai hare. 

After comparing the Gobi hares with L. centrasiaticus of the 
tibetanus group, I found that they are very similar. The Gobi hare is 
possibly a natio or a subspecies closely related to centrasiaticus . 

A. S. Lukashkin sent me several skulls of Tolai hares from northern 
Manchuria, along the eastern side of the Great Khingan. Apart from these 
skulls, I have only one pelt obtained 13 November 1934 in the valley of the 
Ivangol river, a right tributary of the Nonni [Nun Kiang] river, near the 
Santien locality, 30 km north of the Chalantun Station of the East 
China Railway. This pelt resembles that of the Transbaikal hare, but 
it is much smaller: length of body with head 480 mm; tail 112 mm (without 
terminal hair); ear 85 mm; hind feet (without claws) 115mm. Weight 
2.5kg. 

The skull of Tolai hares from north Manchuria is smaller than that of 
specimens from Transbaikalia. 

Measurements of skull of north Manchurian Tolai hare: length of skull 
84-88.2 mm; condylobasal length 76.2-78 mm; zygomatic width 40.5- 
43.6mm; length of nasals 36-40.3 mm; widthof nasals 17.2-18.7 mm; width 
of skull at level of auditory apertures 33.2-34.6 mm; length of bony palate 
6.2-7.3 mm; width of incisive foramina 8.1-9.5 mm; length of row of upper 
molars 15- 15.5 mm. 



147 



170 It would thus be possible to consider the Manchurian Tolai hare as a 
separate subspecies. However, I am waiting for more material to make this 
description more accurate. It is especially necessary to determine the 
relationship between the Manchurian Tolai hare and the Gobi Desert hare. 

In spite of the limited material, it can be stated that the following 
cranial characters of the Transbaikal Tolai hare are also present in the 
Manchurian form: 

l) short postorbital processes; 2) width of nasals in their posterior third 
(longer than postorbital process); 3) short nasals (as long or slightly 
longer than frontal suture); 4) mandibular condyle massive, etc. 

Lepus quercerus from the Chuya Steppe and its systematic position 
will be discussed in the chapter on the desert hares. 

GEOGRAPHICAL DISTRIBUTION. The Tolai hare is widely distributed 
throughout southern Transbaikalia, where it occurs near Troitskosavsk along 
the Selenga river, near Verkhneudinsk [Ulan-Ude] in the Aga Steppe, the 
Steppes of Dauria, Onon river valley, near Kulusutai. It occurs near Lake 
Tarei-Nor, Dalai (Hulun)-Nor, Buir-Nor (G. Radde, 1862) near Lake Koso 
Gol, in the upper reaches of the Irkut river (Radde, 1862). A pelt and 
skull of a Tolai hare from the eastern Sayans, near the village of Mondy, 
25 km north of Lake Koso Gol is in my collection. I also consider as a 
Tolai hare a specimen with an aberrant coloration obtained in the 
Samarskoye District, on the left bank of the Irtysh. The boundary of the 
range of the Tolai hare is apparently very close to that of the desert hare 
(L . t . le hm an ni ) in this area. 

The American Expedition found Tolai hares in northern Mongolia, 30 
miles south of Ulan-Bator (Urga), and also south of Uliassutai [Jibhalanta] 
near Tsagan-Nor, Argabogdo, Ula-Usa. 

A.N. Formozov (1929) caught Tolai hares near Lamen-tJegen, in the 
lower reaches of Tuin-Gol, on the northern coast of Lake tJreg Nur, in 
Yihe-Bogdo (Gobi Altai), along the river Aidarik, in the valley of river 
Dzabhan Gol. 

The Tolai hare is thus apparently widely distributed in northwestern 
and Central Mongolia. K. A. Satunin (1907) described the Gobi hare from 
northern Gobi, apparently a subspecies oftibetanus closely related to 
Lepus tibetanus centrasiaticus Sat. 

Towards the east, R. Maak (1851) recorded the Tolai hare along the 
Argun [Ehrkuna Ho]. He thinks that it occurs south of Aihun, on the way 
to Tsitsihar. According to A.S. Lukashkin (1933) it occurs in the Barga 
[Hulun-Buir] District* of the Heilungkiang Province, Manchuria. This 
region is an eastward continuation of Russian Transbaikalia in its general 
character. According to A.S. Lukashkin, the Tolai hare occurs in large 
numbers between the Manchouli and Hailar stations, near the Station 
Heierhungteh. ■■''■'■' 

171 MODE OF LIFE. Our knowledge of the mode of life of the Tolai hare 
is unfortunately very incomplete. B.A. Kuznetsov (1929) states that it 
lives among Arctic dwarf birch in ravines and among shrubs in tidal 
marshes of Transbaikalia. 

A.N. Formozov writes: "The favorite habitats of the Tolai hare in 
Mongolia are two connpletely different biotopes. The hare is most numerous 



♦ [Now in the Inner Mongolian Autonomous Region. ] 
** As stated above, local Tolai hares differ sharply from Transbaikal specimens. They should be carefully 
described from extensive material. 

148 



in river valleys (Tuin-Gol, Dzabhan Gol, Murin-Gol rivers, etc, ) and lake 
basins (tJreg-Nur). One can find three or four during half an hour in 
these places, in thickets of tall L yc iu m halimifolium Mill., shrubs 
ofCaragana spinosa, Nitraria schoberi and willows. The 
animal is rapid and hides quickly behind mounds and vegetation when seen 
by men. 

"We were surprised to find them high in the mountains (on the Yihe- 
Bogdo, to an altitude of 8000 feet), in meadows with many marmot burrows. 
Tolai hares calmly sit here on marmot mounds. They let the hunter 
approach closely, and then hide in the burrow. It is useless to wait for 
their reappearance. They were found near marmot burrows also in Laman- 
Gegen. There were only a few in meadows with numerous marmots and 
susliks. Only once did we find them in rocky desert covered with pea- 
shrubs. The diet of this hare is unknown. On the tJreg-Nur, the Tolai 
hare fed on Nitraria berries. Tolai hares apparently have at least two 
litters during summer. We found at the same time large young hares and 
suckling females. " 

A.S. Lukashkin (1933) reports that the Tolai hare lives in river valleys 
and in the sandy Barga Steppe overgrown with willow thickets, in the 
Heilungkiang Province of Manchuria*. He also reports that in Barga the 
Tolai hare has only two litters per year. Litters in September are 
very rare. 

Tolai hares occasionally occur near human settlements. 

A.S. Lukashkin writes that in the vicinity of Hailar, the Tolai hare is 
often hunted close to the villages, near gardens of the Chinese which it 
enters from the open steppe. He found the lair of a hare 30-40 steps from 
the Chinese huts. The dogs of the Chinese sit on the roof of the huts for 
days, barking. However, the Tolai hare lies calmly and fearlessly under a 
willow or in a hollow on the ground. 

The Tolai hare had no commercial value in Manchuria until recently, as 
its fur is of inferior quality. Only recently did an English food-export 
company begin to manufacture canned and frozen meat products for export 
to England. 

This firm exported about 6000 dressed hares from the Heierhungteh 
Station in December 1927, The firm hired eight hunters, who killed SO- 
SO hares daily in spite of the severe cold at the time. These figures show 
that the Tolai hare is very numerous and widely distributed in Barga. 
172 According to A.S. Lukashkin ("Okhotnichii i pushnoi promysel na 

Bol'shom Khingane" (Hunting and Fur Trade on the Great Khingan), "Vestnik 
Man'chzhurii" No. 5, 1934, p. 140) the Tolai hare occasionally produces 
hybrids with small local blue hares. These hybrids are called Tumacs, 
like the hybrids of the common and blue hare. 

159. Lepus (Eulagus) tibetanus Waterhouse (1841). Desert hare 

1841. Lepus tibetanus Waterhouse G, R. , Proceedings Zoological Society of London, pp. 7-8 (Little Tibet). 

NAMES: The Afghan hare, the desert hare (English) ; der Hase von Westtibet, der Wustenhase (German). 

LOCAL NAMES: kuyan, or koion (Kazakh, Kirghiz); taushan (Turkmenian); khar-gush ("Donkey's 
eais" - Persian). 



* [Now in the Inner Mongolian Autonomous Region. ] 



149 



PRINCIPAL FIGURES: 1) BlanfordW. T., Eastern Persia II, Zoology 1876, pp. 80-82, pi. VIII (color plate of 
entire animal under the name Le pus craspedotis); 2) Blanford W. T. , Scientific Results of the Second 
Jarkand Mission, Mammalia, 1879, p. 63, pi. IV, fig. 2 - Lepus tibetanus (entire animal in color); 
3) Blanford W. T. , I.e., 1879, pp. 67-68, pi. V, fig. 1 (entire animal in color); pi. Va, fig. 1-lb (skull) - 
Lepus pamirensis; 4) Blanford W. T. , I.e., 1879, pp. 68-71, pi. V, fig. 2 (entire animal in color): 
pi. Va, fig. 2-2b (skull) - Le pus s t ol ic zk anus ; 5) BogdanovM. N., Ocherki prirody Khivinskogo 
oazisa (A Survey of the Nature of the Khiva Oasi^, 1882, p. 68 (half tone plate of entire animal under the 
name Lepus kessleri); 6) Marcus Word Layon, Classification of the Hares and their Allies, 
Smithsonian Miscellaneous Collections, Vol.1, parts 3 and 4, 1904, pi. LXXXIV, LXXXV, fig. 4 
(photographof skull dorsal and ventral -Lepus tibetanus); 7) Kashkarov D.N. , Zhivotnye Turkestana 
(Animals of Turkestan), 1932, p. 196, fig. 140 (under the name Le pus tolai lehmanni); 8) Heptner 
V. G. , Zaitsy (Hares), Moscow, 1933, p. 23, fig. 3 (entire animal); 9) Vinogradov B, S. , Pavlovskii E. N. , 
Flerov K. K. , Zveri Tadzhikistana, ikh zhizn' i znachenie dlya cheloveka (The Life of the Animals of 
Tadzhikistan and their Importance to Man), 1932, p. 252, fig. 86 (in summer fur); p. 254, fig. 87 (in winter 
fur, under the name Le pus tolai Pall.). 

MATERIAL EXAMINED: 74 specimens. 

DIAGNOSIS. Head relatively small. 

Length of ear (measured from occiput) 1.25-1.75 times as long as head. 

Length of tail with terminal hair about 4/5 of length of hind foot (without 
claws). 

Tail white with contrasting black of black-brown dorsal side in summer 
and winter fur. 

Hair not becoming white in winter. Winter fur only lighter (changing its 
tone) and slightly different from summer fur in being fluffier and denser. 

Fur very soft, silky. Guard hairs soft, delicate and straight. Fur not 
curly. 

Teats 8. Nasals measured along line connecting inner anterior corner 
wath outer posterior corner usually longer than frontal suture. 

Maximal width of both nasals in their posterior part about equal to 
length of postorbital process, rarely exceeding it. 

Posterior third of incisive foramina widened, without protuberances, 
and with straight margins. 

Anterior lower molar sharply inclined backwards in profile. 
173 Mandibular condyle weak, strongly inclined backwards. 

Lower posterior plate of mandible (from corner of angular process to 
notch at beginning of mandibular horizontal process) weak. Distance 
between angular process to notch at beginning of mandibular horizontal 
process much greater than that between anterior alveolus of first premolar 
to tip of lower incisor. The lower posterior mandibular plate of the 
common hare is also rather weakly developed. However, the skull of 
the common hare differs from that of the desert hare as follows: l) the 
ratio between the two measurements mentioned above is the opposite of 
that of the desert hare; 2) the lower diastema of the desert hare is 
much shorter, and the whole mandible is thus little elongate anteriorly 
and relatively short. This distinguishes the desert hare from the common 
hare. 

Bullae osseae larger, strongly inflated. The rounded bullae osseae 
protrude markedly anteriorly in side view (Figures 89-92). 

Length of ear from occiput 100-145mm; hind feet 106-130mm. 

ADDITIONAL DESCRIPTION. The skull of the desert hare is very 
similar to that of the common hare; in some respects it is a miniature 
of it. However, there are the following differences: l) bony palate of 
desert hare relatively wider than that of common hare; 2) incisive 
foramina narrower, sides of their posterior third usually without 



1254 150 



174 










^ 


r—> 





M 


o 


00 


> 




01 







<M 


tV) 


Ol 


O 


<T| 




00 


HH 


£2 


(/) 


« 


^ 


n 


0) 


o 


^ 



151 



173 protuberances; 3) ratio between width of incisive foramina and length of 
bony palate different from that of common hare; 4) mesopterygoid fossa of 
desert hare narrower than that of common hare, and ratio between length 
of mesopterygoid fossa and of bony palate also different; 5) length of 
lower diastema and of mandibular process smaller than in the common hare; 
this affects the ratio between the length of the row of lower molars and the 
lower diastema. 

COLOR. The color of the back of the desert hare varies strongly 
according to the locality. Mountain forms are as a rule darker than valley 
forms. According to Blanford, the color of the back varies from light 
gray='= to rust-brown in the nominate subspecies, Lepus tibetanus 
tibetanus. Specimens of these two color types have a distinct black 
tinge. The lower part of the back is often sand-gray. The brownish gray 
ground color of the inner sides of the ears is mixed with black"'*. 
175 A specimen of Le pus tibetanus tibetanus No. 12393 female 
adult, 25 May 1879, collected by J. Scully in Gilgit is in the ZMAN 
collection. This locality is only a little north of the terra typica of L . 
tibetanus, adjoining the Pamir Plateau, but separated from it by the 
Hindu Kush and Karakorum massifs. 

General color of back of this specimen dull grayish yellow mixed with 
blackish hair tips. Color of top of head like that of back, i.e., ground 
color straw -gray with fine blackish mottling. Anterior inner margins of 
ears of the same color. Black-brown hair on tip of ears gives a darker 
color to the apex. Broad black-brown fringes on the outer apical 
part of the ears. Flanks paler than back, with a dull straw-gray tone. 
Chest with a transverse dull straw-gray stripe (paler and grayer than 
avellaneous, Ridgway, pi. XL). Lower part of back grayer. Cheeks 
whitish gray, with blackish speckling. Eyes surrounded by a whitish 
ring. 

The general color of this hare closely resembles that of the Pamir 
subspecies, and is almost identical with that of darker specimens of L . t . 
lehmanni. 

Color of Lepus tibetanus subsp. from Kopet-Dagh similar to that 
described above; general color of back straw-gray with well marked 
blackish mottling. Inner margin of ears with a distinct blackish tinge on 
a straw-gray background. 

The color ofL.t. lehmanni also resembles that ofL. tibetanus 
subsp. from Kopet-Dagh. 

The dull tones of color with the blackish tinge are typical of a group 
of desert hares which could be called "relatively dark forms". This 
group also includes L. t. craspedotis. 

A group of light-colored desert hares can also be distinguished. The 
color of the palest members of this group is very dull. For example, 
L.t. desertorum has an unusually pale straw- sand color in summer 
fur with a hardly perceptible blackish tinge. In winter specimens, the back 



The description of the type of L. t i b e t a nus by G. R. Waterhouse (1841) differs sharply from the more 
recent description in Blanford's work, Waterhouse described the type from a specimen from Ngari (caught 
by Vigue near Skardu). Blanford described specimens from the valley of the Nutra river in Ladakh. 
The color of this hare is generally pale, according to Waterhouse, without any yellowish or rust tones. 
Hair on chest pale gray with slightly rust-colored hair tips. Ears very large, their inner margins pale gray. 
It is possible that specimens of the second color type from Ladakh, which were identified by Blanford as 
tibetanus, belong to another form. This can only be determined by comparison of material. 



152 



has an irregular blackish mottling.on a general light straw-yellow 
background. In some subspecies, the back has a rust-red tinge on a sand- 
straw colored ground (e.g., inL.t.buchariensis). The color of 
L.t. pamirensis and L.t.zaisanicus is more intense , In the 
latter, the color of the back is r^ather pale, straw-gray with a blackish tinge 
in summer fur. 

As stated above, the color of the summer fur differs little from that 
of the winter fur, which is often darker than the summer fur with a slight 
black tone caused by the blackish brown tips of the long guard hairs or 
by black-brown parts of some hairs on the back. 

General habitus of desert hare light and graceful. Its ears seem long, 
its haunches thin and its tail protrudes markedly. 



176 







FIGURE 93, Desert hare, Lepus tibetanus buchariensis Ogn. , Lower Vakhsh, Tadzhikistan, 
1932 Drawing by K. K. Flerov 



153 



178 



SYSTEMATIC NOTES. The opinions of authors on the systematic 
relationships between the various forms of desert hares differ sharply. 
Authors like Blyth, Hodgson, Blanford, etc. considered some desert 
hares as separate species. They (e.g., Blanford) pointed out, however, 
the close relationships between them, and their great resemblance. 

W.G. Heptner* has recently proposed a new classification. He suggests 
considering the small hares of the Mediterranean subregion, L . 
granatensis, mediterraneus, parnassius and c r e t i c u s , 
as subspecies of Le pu s europaeus, but also subspecies of true 
desert hares such asL. judeae, tibetanus, lehmanni and t o 1 a i . 
Heptner considers all these forms as one species group (Rassenkreis), 
but he excludes from this L, oiostolus, L.hypsibius, L. 
pallipes and L. kozlovi. This seems correct, as all these forms are 
clearly distinguishable from the, common hare. However, there is much 
evidence against considering the common hare, the Tolai and desert 
hares as one species, as proposed by this author. 

We should like to stress the well marked cranial differences between the 
common and the desert hares (see above). The structure of the palate 
of the Central Asian hares closely resembles that ofLepus americanus. 
The same applies to the ratio of the length of the row of lower molars to 
the lower diastema. It is significant that the Central Asian hares resemble 
the American hares more closely than the common hare in these important 
cranial characters. 

The Central Asian hares differ from the common hare also in the 
following characters: l) fur very soft, silky; 2) guard hair relatively 
thin, silky and not curly as in many common hares. 

The range of the common hare does not overlap that of the desert hare. 
The common hare is restricted to the European continent. It apparently 
has a southern, Mediterranean focus of distribution. 

In the course of evolution the southern forms of the common hare became 
sharply different from the typical race and developed into separate 
species. -■'•'■'• These forms have good color and cranial characters. Some of 
these recently developed species occur on islands: Lepus granatensis 
(Balearic Islands), L. mediterraneus (Sardinia), and L . c r e t i c u s 
(Crete). Other species occur in the southern parts of Europe, in the 
Mediterranean subregion (Spain, southern Greece). Miller (Catalogue 
of Mammals of Western Europe, 1912) shows that these species differ 
sharply from the true common hare in their measurements, color and 
skull characters. 

As we have little material, we cannot at present reject Miller's 
analysis of the relationship between the Mediterranean hares and the true 
common hares. 

The common hare apparently spread from its southern European focus, 
but its exact location cannot be determined at present. 

This emigration took place towards the north, east and the southeast. 
It increased during the last century owing to the felling of forests and 
cultivation. The ranges of the common hare and the desert hare (L . t . 
lehmanni) became adjacent in Kazakhstan (in the former Syr -Darya 



♦ G. Heptner, Systematische und tiergeographische Notizen uber einige russische Sauger. Folia 

Zoologica et Hydrobiologica, Bd. VI, No. 1, 1934, S. 21-23. 
** There are no transitions between these forms and the true common hares. 



154 



177 







FIGURES 94-98. Postures of the desert haxe, Lepus tibetanus buchariensis Ogn. 

Drawing by K. K. Flerov 



155 



179 



180 



Region) and these two sharply different species occur together in this 
region. The true common hare also spreads eastwards. The general 
appearance of common hares occupying new territories changes in some 
respects. It is possible that in the future the range of the common hare 
will reach the range of the true Tolai hare, but not that of the desert hare. 

It seems probable that the center of dispersion of the desert hare was 
Central Asia. It is very widely distributed at present in this area and 
forms many subspecies. As pointed out, the Central Asian desert hares 
have much in common with some American hares, e. g., L. americanus, 
in some cranial characters, especially the structure of palate and incisive 
foramina. 

The skull ofL. americanus also shows the cranial characters of 
the blue hare, like the structure of the mandible, the upright position of 
the first lower molar, etc. The skull of the hares in the desert zone of 
Arizona (L . alien i Mearns) has characters of the common hare, but 
also some specific characters, such as a long braincase. This species 
has the following cranial characters: 1 ) coronoid and condyloid processes 
of mandible strongly inclined backwards; 2) lower diastema elongate; 
3) lower molar inclined backwards; 4) incisive foramina wide; 5) sides 
of posterior third of incisive foramina with rounded protuberances. On 
the other hand, the skull of L. alleni has the following cranial 
characters which it has in common with the desert hare: l) length of bony 
palate; 2) narrow mesopterygoid fossa; 3) markedly inflated and rounded 
bullae osseae. 

The pale color ofL. alleni, and the black color of the dorsal side 
of the tail are characters it has in common with the desert and common 
hares. 

Without going into detail, it is interesting to note that the skull of the 
Central Asian hares, and also the general appearance in some species, have 
much in common with those of North American species. 

A general survey of the specific and subspecific groupings of the Central 
Asian hares is given below. This survey is provisional as we have 
insufficient material on many species which we shall have to describe from 
the literature. 

Lepus jarkandensis Giinther (described from the valley of the 
Yarkand) is undoubtedly a good species. This small species has the 
following characters: very pale straw-yellow color (intermediate between 
pinkish buff and avellaneous), pale inner sides of ears (pinkish buff), straw - 
yellow flanks (pinkish buff), very pale chest, with a slight straw-yellow 
tinge on white ground. Color of tail typical; dorsal side of tail with a 
brownish straw-gray dorsal stripe which is yellow laterally (pinkish buff). 

Color of summer fur similar to winter fur, but grayer. Straw-yellow 
tones on chest more marked. 

The skull of this species is typical: bullae osseae very large and 
inflated; roof of braincase flattened; sides of parietals parallel; bony 
palate very long. 

There are transitional forms between L. jarkandensis Gilnther 
and the desert hare. 

A large group of subspecies belongs to Lepus tibetanus Waterhouse 
(1841) which was described from a specimen from Skardu in northern 
Kashmir, and which occurs in Ngari. Lepus tibetanus tibetanus 
is a highland desert form, distinguished by a grayish color. The skull 



156 



181 



(Marcus Ward Lyon, 1904) resembles that of the Pamir desert hare. The 
ranges of these two forms are adjacent. The following subspecies may be 
considered as belonging to L . tibetanus: Lepus tibetanus 
lehmanni Sev. , L.t, pamirensis Blanf. , L. t.buchariensis 
Ogn. , L.t. stoliczkanus Blanf. ,L. tibetanus subsp.? (from 
Kopet-Dagh), L.t.craspedotis Blanf., L.t. zaisanicus Satun. , 
L.t. desertorum Ogn., L.t. centrasiaticus Satun. , L.t. 
quercerus Hollister, L.t. przewalskii Satun., L.t. gobicus 
Satun., L.t. kozlovi Satun. =■= 

The Himalayan-Tibet hares which have completely white tails without 
black or black-brown color on the dorsal side are possibly a separate 
species, L. oiostolus Hodgson. To this color type belong the following 
species: l) Lepus oiostolus Hodgson (1840) from Nepal and Sikkim 
which also occurs in Ladakh according to Hodgson; 2) Lepus pallipes 
Hodgson from Lhasa, considered by W.T. Blanford ("Fauna of British 
India", 1888, p. 452) as a synonym of L . oiostolus Hodgson; 3) 
Lepus hypsibius Blanf. This species is closely related to L . p a 1 1 i p e s 
but differs from it in some external characters, such as color, measurements 
and skull structure (W.T. Blanford, Scientific Results of the Second 
Jarkand Mission. Mammalia, 1879, p. 61). 

There is a large series of hares from northern and eastern Tibet, the 
upper reaches of the Hwang Ho, the vicinity of Shanerda, Nan Shan, 
Kuku-Nor, western Tsaidam (Valley of Winds), and Alyk-Nor, in the 
ZMAN collection. The hair of the back in these species is curly in winter 
fur, light rust-yellow, slightly pinkish. The lower part of the back is 
light smoke -gray; its color differs sharply from that of the middle of the 
dorsal side. These hares are determined as Le pu s oiostolus 
Hodgson in the ZMAN collection. However Lepus oiostolus is 
distributed in Nepal and Sikkim; it is separated from the above-mentioned 
species by the large Tibetan Highlands. The identification as Lepus 
oiostolus is therefore doubtful. 

The characteristic feature ofL. oiostolus, the completely white 
tail, is also shown by some specimens of the series in the KZMAN. Thus 
the tail of a hare from the upper reaches of the Kara-Sai river (eastern 
Tibet, collected by Pevtsov) is completely white. The tail of a specimen 
from the Oi-Tulan-Khutum river (collected by Pevtsov) shows a darkening 
in the proximal quarter of the dorsal side. The same 'character is also 
shown by a specimen from western Tsaidam (No. 12375), from the Valley 
of the Winds (collected by N.M. Przheval'skii). In hares from Gui-Dui, 
the Hwang Ho river, the Humboldt Mountains and other regions of 
eastern Tibet, the dorsal side of the tail is dark brown or black. 

Presence or absence of black color on the dorsal side of the tail is a 
strictly constant character. The above data make it doubtful whether 
this character is constantly present in Central Asian hares, but this 
requires careful study and examination of a large amount of material. 

Specimens of Le pu s centrasiaticus (a subspecies of t i b e t a nu s ) 
are closely related to east Tibetan hares. The color of the back in winter 
fur is very light, clay-yellow in these hares. The lower part of the back 
is whitish, the hair not curly. L, centrasiaticus Satun. is 



* Owing to lack of material, I cannot decide whether Le pus filchneri Matschie, L. aurigineus 
Hollist, andL. swinhoei Thos. belong to this group. Gl. Allen (1927) places L. tolai into this 
group as a subspecies. 

157 



182 






1S8 



181 



apparently distributed throughout Sinkiang, Hami [Qomul] and Kansu, and 
apparently also in Dzungaria. It does not occur in Ala Shan. There are 
specimens from Sachow, Ala Shan, Raskem, Etsin-Gol and Hami in the 
ZMAN collection. 

L. t. centrasiaticus presumably occurs in the far north. It forms 
a more clay-colored, but still very light form, L.t. gobicus Satun. * 
in the Gobi Sands. It would be interesting to determine the relationship 
between this hare and the true Tolai hare, with which L.t. gobicus 
Satun. has some common traits. 

Different hares occur in Tsaidam (Gas, southern Tsaidam). Specimens 
ofLepus t. przewalskii Satun. are closely related to L . t . 
centrasiaticus Satun., but their winter fur is even paler, more 
pinkish, with the lower part of the back white. 

Hares from Khan are also typical (Nam-Chyu river. Basin of the Yangtze 
river, Gui-Dui, Re-Chyu, Bar-Chyu, collected by P. K. Kozlov). K.A. 
Satunin described them as a separate species, Lepus kozlovi Satun. 
They are characterized as follows: l\) nasals narrowed anteriorly; 2) 
anterior lateral part of skull low (in region of maxillaries and 
premaxillaries); 3) frontal area with elliptical depression in the 
interorbital region; 4) lower incisors markedly protruding anteriorly, 
pointing directly forward without upward curvature; 5) postorbital 
processes elevated in profile (Figure 100). This is one of the most typical 
characters of this form. 

This form should be named correctly L.t. kozlovi. It has long 
ears, 130-140 mm along the concavity. Color of dorsum dark with a black 
tinge. Lower part of back different from the upper part by its dark ash- 
gray color. 

Winter fur of L . t . kozlovi resembles that of some specimens of 
L. tolai tolai Pall. However, L. t. kozlovi differs from them in its 
long ears and markedly different skull structure. 
183 The skull of L . t . ko z lo vi resembles that of hares from northern and 
eastern Tibet. However, the color of north Tibetan hares is much lighter; 
the lower part of the back, though grayer, is still lighter. 

Thus, moving to the south in the Kahm mountains, one finds hares with 
a more intense color of the upper part of the back and with a grayer lower 
part of back. 

Hares from Kansu, which is a transitional zone, were considered by 
K.A. Satunin as a separate species, Lepus gansuicus Satun. They 
are closely related to L . t . centrasiaticus Satun., and are apparently 
identical with it. Type No. 2854 (female, 3 June 1890, Kansu, Grum- 
Grzhimailo) is in summer fur and cannot therefore be compared with winter 
specimens of c e nt r a s i at i c u s . However, winter specimens of 
gansuicus (No. 2840, 8283, 2839, etc. ) do not differ from 
centrasiaticus. 

Finally, it should be mentioned that L e p u s kaschgaricus described 
by K.A. Satunin is not monotypical. The specimen of ka s c h gar i c u s 
from the Yarkand Oasis first mentioned in K.A. Satunin's work certainly 
belongs to i ar k an de n s i s . Specimens of L . kaschgaricus from 
Tien-Shan and Yuldus are very closely related to pamirensis. 



* L. tibetanus centrasiaticus Satun. shows gradual transitions to the Turkestan desert hare L. t. 
lehmanni Severtz. 



159 



A detailed systematic analysis of Central Asian hares is beyond the 
scope of this book. I have dealt briefly with this question to clarify the 
relationship between these hares and those in the territory of the U. S. S. R. 

GEOGRAPHICAL DISTRIBUTION of the desert hare and its seven 
subspecies. 

The northern boundary of its range begins from the Buzachi Peninsula on 
the northeastern coast of the Caspian Sea. From there it crosses Ust-Urt 
and continues to the northwestern shore of the Aral Sea. We have no 
exact information as to whether it reaches the lower Emba river, though this 
is very probable. 

A.M. Kolosov informed me that among a large number of Caspian 
common hares, 100km from the estuary of the Emba river, he had seen a 
very small, long-legged, light coloredhare in the summer of 1934, which 
differed clearly from the common hares and was possibly a desert hare. 

The range of the desert hare then apparently crosses the Kara-Kum, 
Sary-Su deserts, the Golodnaya Step' (Bet-Pak-Dala), and extends to 
Lake Balkhash. It is not known where the northern boundary of the range 
passes north of Lake Balkhash. The boundary then continues to Lake 
Zaisan and to the Zaisan Depression, and reaches the Chuya Steppe on the 
Lesser Altai. It certainly occurs in Dzungaria. However, the northern 
and eastern boundaries of the range in Dzungaria cannot be exactly 
determined. It occurs on the Tien-Shan but it is not known how far to the 
east. 

The eastern boundary of the range of the true desert hare also includes 
Kashgar. In eastern Turkestan (along the Yarkand-Darya) a different 
species occurs, Lepus j a r k an de n s i s . The boundary then turns 
southward and includes the whole Pamir, Kashmir, the upper reaches of 
the Indus (Ngari), Afghanistan, Baluchistan, Iran (south to Lazistan, 
Khuzistan and Farsistan). It is not known how far west the range of the 
desert hare extends in this region, but it certainly occurs in northeast 
Iran (Khorasan, Iraq-i-Ajam)''=. L.e.cyrensis Satun. occurs in 
Azerbaidzhan. 

The desert hare was observed in the Buzachi Peninsula by A. Ostrovskii 
(1889); it was recorded from Ust-Urt by I. Brandt (1852) andby Eversmann 
(1850). Eversmann writes that the desert hare also occurs in the steppes 
near the Aral Sea. It is very common in the former Syr-Darya region 
(N.A. Zarudnyi, 1915). Large numbers occur near Kara-Kemken (N.M. 
Dukel'skaya), and near Kzyl-Orda (S.P. Naumov, 1927). N.A. Severtsov 
(1873) points out that it is common on the Kara-Tau, the Hi, along the 
Syr-Darya, as far as the Aral Sea and the eastern coast of the Caspian Sea. 

According to M.N. Bogdanov (1882) it is especially numerous in 
Haloxylon shrubs along the Yany-Darya. The data of A.N. Nikol'skii 
(1887) show that it is very common on sandy ground along the southern 
shore of Lake Balkhash and along the river Lepsa. It is very numerous 
along the Hi river near the village Hi. 

It occurs in the Zaisan Depression from where K. A. Satunin described 
a new species, L.zaisanicus, from near Lake Zaisan. Specimens 
collected there by G.I. Polyakov's expedition are in my collection. It 
occurs in the Chuya Steppe (Chegan-Burgazy, Kosh-Agach). According 



R. E. Cheesman (Report on a Collection of Mammals made by J. Hotson in Shiraz, Persia, Journal of the 
Bombay Natur. Hist. Soc. , 1921) considers hares from Shiraz as Le pus lehmanni Sev. 



160 



185 



to V.N. Shnitnikov (1932) it is distributed throughout Dzhetysu [Semireche] 
at an altitude of 300-3000 m. The distribution of the desert hare in 
Dzhetysu seems sporadic. In some regions, e.g., the Trans -Ili Ala-Tau 
Mountains (near Alma-Ata), it disappeared after 1933. In other localities, 
e.g., along the Chilik river, it is common. N. A. Severtsov (1873) remarks 
that the desert hare is distributed on the Tien-Shan to an altitude of 10,000 
feet and is numerous in the valley of the Chilik river and its tributaries, 
north of Issyk-Kul, on the Soktyube Mountain Pass, the Nyr-Dzhal river, 
on the Alexander Range, and near Merke. It was found on the Alexander 
Range in 1935, by the expedition of A. A. Mashkovtsev. D.N. Kashkarov 
(1932) writes that the desert hare occurs in large numbers on the sands 
of the Terskei-Ala-Tau, in the valley of the Sarychat river. 

According to S. N. Alferaki (1891) it is very numerous near Kuldja and 
common on the Pamir. M. P. Rozanov (1935) writes that the Tolai hare 
(i.e., the desert hare) occurred in large numbers when the expedition 
crossed the Trans-Alai Range (through the Kyzyl-Art Mountain Pass and 
the Markan-Su landmark area). 

The desert hare is widely distributed in Tadzhikistan in Turkmenia 
where it was found by P. Varentsov (1894) in the Tedzhen District, and in 
the Areshsk District from the railway station at Dushak to the village of 
Mean. According to Radde, Walter, Blasius (1890) it is found from 
Krasnovodsk, Mikhailodo, Chikishlyar to the Amu-Darya (where it occurs 
on a large island near Chardzhou). These authors also observed it near 
the Afghan boundary, on the Atrek and Chandyr. According to S. I. 
Bil'ketvich (1918) it is found near Repetek, Kizil-Bai, Tedzhen, and 
especially numerous in the Kara-Kum sands. I saw it on the Kopet-Dagh, 
on the Bolshie Balkhany Range and in the adjacent deserts. 

It is apparently widely distributed in Iran (Thomas, 1905), south to 
Baluchistan (W. Blanford, 1875, 1876), and also throughout Afghanistan, 
though we have no exact data on its distribution in this country. It occurs 
in Kashgar (Dzhigda, Altum-Artush District, vide W. T. Blanford, 1879). 
It is found from northern Kashmir to the upper reaches of the Indus, in 
Ngari (W.T. Blanford, 1879, 1888). 

MODE OF LIFE. The typical biotopes of the desert hare are desert 
and steppe, especially those adjacent to river valleys with tugais*. 
It is also found in deserts among sand dunes overgrown with Halozylon, 
Peganum harmalaL., and Aristida pennata, and in semideserts 
with tamarisks. It avoids open clay desert overgrown with Artemisia 
and Sal so la. 

The desert hare is very common in river valleys and in depressions of 
lakes with thickets of shrubs. Thus, it occurs in large numbers in the 
lower reaches of the Talas river valley, along tugais of the Syr-Darya 
where Peganum grows occasionally, in thickets of Lasiagrostis and 
Hippophae rhaninoides, along the shore of Lake Issyk-Kul or in 
thickets of Nit r aria, on saline soil along the outflow from the Buam 
Ravine into the lake, in tamarisk shrubs in the lower reaches of the Sary- 
Su, and in tamarisk thickets in the vicinity of Lake Baili-Kul (D.N. 
Kashkarov, 1932). 

Foothills, high mountains and the subalpine belt (to about 3500-4000 m) 
are also included in the large range of the desert hare. 



[Tugai — floodland forest in river valleys. ] 



161 



186 



According to M. P. Rozanov (1935) the desert hare lives on the eastern 
Pamir in summer, on moraines and on slopes among screes, and in 
thickets of Eurotia and Artemisia. 

These plants are the main food of the hare in this region. The Pamir 
hare avoids humid meadows and rocks. Towards winter, according to 
local inhabitants, the hares descend in large numbers into the river 
valleys. This shows that the mountain hares make true seasonal vertical 
migrations. M. P. Rozanov (1935) recorded the hare from the pebbly bed 
of the Pyandzh river in the western Pamir. It lies here among sand dunes 
in thickets of Hippophae and tamarisk, and also on the dry, stony slopes. 

W.G. Heptner (1933) writes: "Many hares are found near villages, 
where the banks of the irrigation canals are covered with grass and 
shrubs. These canals cross the cultivated areas, dividing them 
into small squares of corn, cotton, and melon fields. Rows of poplars 
grow along the roads and between the plots. The houses are surrounded 
by fruit orchards. The cemieteries, which are usually near the village, are 
overgrown with weeds and trees. When there is plenty of water, small 
forests are often planted. Hares are occasionally very numerous and 
cause considerable damage to agriculture. They gnaw young wheat sprouts 
and leaves of cotton. They are especially harmful to melons and watermelons. " 

The desert hare is a rather timid animal in many places; in Turkmenia 
they ran away to the distance of a long shot when frightened. In places 
where hares are not used to the presence of man, they are confident and 
careless. They run rapidly and lightly. Their legs are well built 
and long. Their ears are rather long (Figures 93-98). 

They are very active at dusk, but in some places also during the day. 
S.N. Alferaki (1891) writes that the desert hares seen by him near Kuldja 
are completely diurnal. 

We have no exact data on the reproduction or food of the desert hare. 

In Central Asia, the desert hare is scarcely hunted by man as its hide 
is of little value and it has not much meat. 

The desert hare is preyed upon by: Vulpes cor sac, desert foxes, 
wolves, polecats, and various cats, and also by the following birds of 
prey: goldeneagle, steppe eagle. Young hares are probably caught by buzzards. 

Subspecies of the desert hare 

334. Lepus (Eulagus) tibetanus lehmanni Severtzov (1873). 
Syr -Darya desert hare 

1852. Lepus tolai Brandt J. F. , Zoologischer Anhang zu A. Lehmanns Reise nach Buchara und 

Samarkand, S. 308-309. 
1861. Lepus arale nsis Severtsov N. A. , Zveri Priural'skogo kraya. Akklimatizatsiya (Animals of the 

Ural Territory. Acclimatization) II (2):49:70 (Nomen nudum.'). 
1873. Lepus lehmanni Severtsov N. A. , Vertikal'noe i gorizontal'noe raspredelenie Turkestanskikh 

zhivotnykh (Vertical and Horizontal Distribution of Turkestan Animals), Izvestiya Obshchestva lyubitelei 

estestvoznaniya, VIII (2):62, 83-84. 
1882. Lepus butlerovi Bogdanov M. N. , Ocherki prirody Khivinskogo oazisa (A Survey of the Nature of 

the Khiva Oasis), p. 68 (Nomen nudum.') (Tugais along the Amu-Darya). 
1882. Lepus kessleri Bogdanov M. N. , op. cit. , p. 68 (Nomen nudum.') (Tugais along the Amu-Darya) 
1922. Lepus tolai lehmanni Ognev S. I. , Materialy po sistematike russkikh zaitsev (Material on the 

Systematics of Russian Hares), Ezhegodnik zoologicheskogo muzeya Akademii Nauk, Vol. XXIII, p. 482. 
1927. Lepus tolai subsp. Kashkarov D. N. , Rezul'taty ekspeditsii v raione ozera Sary-Chilek (Results 

of an Expedition to the Lake Sary-Chilek Region), p. 101. 



162 



187 



188 



1929. L[epus] tibetanus lehmanni Ognev S, I. Zur Systematik der russischen Hasen, Zoologischer 

Anzeiger, Bd , 84, H. 1/4. S. 78. 

TYPE LCX;AIJTY and deposition. Lower reaches of Syr-Darya river. It is not known whether the 
type is preserved. 

MATERIAL EXAMINED: 24 pelts with skulls. 

DIAGNOSIS. Back pale, straw-gray (intermediate between avellaneous 
and drab). Ground color with blackish mottling weak in summer fur but 
well marked in winter fur due to long black hair and also to hair with 
gray-black bases, broad whitish subapical bands and black tips. Lower 
part of back in both summer and winter fur more ash-gray than rest of 
back, without a straw tinge. 

Top of head whitish gray or whitish straw with marked black -brown 
mottling due to hairs with black-brown tips, broad black-brown bases 
and subapical nmedium whitish or straw- whitish bands. 

Color of neck between pale yellowish straw (intermediate between 
pinkish buff and avellaneous), and reddish brown-gray (avellaneous 
with a reddish rust tinge). Color of neck in winter fur usually richer than 
that in summer fur. 

Chest grayish rust-straw, intermediate between light vinaceous-buff 
and light pinkish cinnamon. 

Color of anterior inner side of ears pale grav with a straw tinge, 
mixed with light blackish brown mottling in summer fur, darker in winter 
specimens. A broad white stripe along the outer fringe of the inner margin 
of the ears. Outer side of ears whitish, occasionally whitish gray with a 
slight straw tinge. Brown-black tips of ears narrow, poorly developed but 
better developed on the outside. 

Anterior side of legs very pale, rust-straw-colored. 

Length of skull 83-88.2 mm (M. 85.6 mm); zygomatic width 38.5- 
42.8 mm (M, 41 mm); length of nasals 34.2-38 mm (M. 36.6 mm); width 
of nasals 14-16 mm (M. 15.7 mm). 

Length of body with head 387-485 mm (M. 455.9 mm); hind feet 108- 
126mm; tail 75-107 mm (M. 97.8 mm); ears (measured along concavity) 
87-lOOmm, ear (measured from occiput) 100-126 mm (M. 119.8 mm). 

ADDITIONAL DESCRIPTION. The above description shows that the 
color of the winter fur does not differ much from that of the summer fur. 
Color of back of summer fur with light blackish brown mottling. The 
color of the back of summer specimens is therefore lighter and sandier 
than in winter fur. Color of chest also somewhat paler, though this is 
not constant. Inner margins of ears of winter specimens have more 
blackish hair than in summer specimens. 

Winter fur easily distinguished from summer fur by its markedly longer, 
softer and silkier hair. 

COLORATION OF YOUNG. Generally paler, grayer and more whitish 
than in adults. 

Color of back straw -gray-white with a light blackish brown mottling. 
Top of head whitish straw with brownish or blackish mottling. Neck very 
pale, straw-pink (vinaceous-buff). Flanks and lower part of back whitish 
gray. Chest very dull yellowish straw color. Color of anterior sides 
of legs richer and yellower. Inner sides of ears pale straw-yellowish 
gray. Outer fringe of inner margin of ears whitish. Inner side of ears 
very pale, whitish straw. Anterior side of ears straw-whitish gray; 
outer side of tip of ear with a distinct narrow black-brown terminal fringe. 



163 



189 



SYSTEMATIC NOTES. The relationships between Turkestan desert 
hares are not clear. 

N.A. Severtsov ("Zveri Priural'skogo kraya" (Animals of the Ural 
Territory), 1861) states that a peculiar hare, Lepus aralensis 
(description not given), occurs in the lower reaches of the Emba and 
further south towards the Aral Sea, Severtsov (1873) described in detail 
the Turkestan desert hare under the name Lepus lehmanni. The 
holotype was a specimen obtained by Lehman in the lower reaches of the 
Syr-Darya. I do not know whether this holotype is preserved. M.N. 
Bogdanov (1882) recorded from the "Tugais along the Amu-Darya" two 
"species" of hares, Lepus kessleri and L. butlerowi, and gave 
their Russian common names, "long-eared" and "roan" hares. 
Bogdanov does not give any description, but on p. 68 gives a poor drawing 
of Lepus kessleri. I had the opportunity of examining the stuffed 
types of these hares in the collection of the Zoological Museum of the 
Leningrad University. The animals are badly preserved, but they hardly 
diffe r from L.t. lehmanni. 

W.G. Heptner" (1934) described a subspecies of the common hare, 
Lepus europaeus turcomanus. Heptner took as holotype a pelt 
and a skull of a hare obtained by S. Naumov and N. Lavrov near the well 
ofAk-Kuyu, 60 km north of Dzhebel (Transcaspia). Hares from the Kopet- 
Dagh which were determined as L e p u s tolai craspedotis Blanf. '■'* 
belong to the same subspecies, in Heptner's opinion (S.I. Ognev and 
W. G. Heptner, 1929). 

Although neither I nor Heptner had a specimen of the true Baluchistan 
Lepus craspedotis, Heptner was probably correct in thinking that 
the Kopet-Dagh hare is different from L. craspedotis. 

I doubt whether turcomanus can be considered as a subspecies 
of europaeus (see above). The Turkestan hares apparently belong to 
L. tibetanus, and not to L. europaeus. 

The systematic position of L. t . turcomanus is not clear. W.G. 
Heptner only gives the following diagnosis "... Steht Lepus europaeus 
lehmanni Sev. nahe, unterscheidet sich aber, wie im Winter-, so auch 
im Sommerfell durch dunklere Farbung mit einem dunkel schwarzbraunen 
Anhauch, durch etwas kiirzeres und grobhariges Fell und dunklere 
Farbung der vorderen inneren Seite der Ohren. . . ". W.G. Heptner described 
Lepus europaeus turcomanus from two specimens of winter fur 
obtained in early March 1932. I have compared these hares with Aral winter 
specimens of L.t. lehmanni and could not find any differences. The 
inner margin of the ears is grayish straw with weak brownish black 
mottling in both specimens from Ak-Kuyu. The inner margin of the ear of 
the holotype (No. 6342) is especially light. Heptner's diagnosis therefore 
does not agree with this holotype. 

In Heptner's opinion, the hares from Ak-Kuyu are identical with hares 
from the Kopet-Dagh, and also with hares from the Bol'shie Balkhany Range. 
We doubt whether this is correct. The inner side of the ears shows marked 
darkening in the summer fur of Kopet-Dagh hares. On the other hand, the 



' W. G. Heptner, Systematische und tiergeographische Notizen iiber einige russische Sauger, Folia Zoologica 

et Hydrobiologica, Bd. VI, 1S34, S. 21-23. 
"• S.I. Ognev and W.G. Heptner, "Mlekopitayushchie srednego Kopet-Daga i prilezhashchei ravniny" 
(Mammals of Centra] Kopet-Dagh and the Adjacent Plain). —Trudy Nauchno-issledovatel'skogo instituta 
zoologii. III (1): 71-73. 1929. 



164 



summer fur of the desert hares (e.g., L. t . 1 e h m a n n i) is lighter than 
their winter fur. The inner side of the ears particularly is of lighter color. 
The winter Kopet-Dagh specimens are probably even darker and differ 
more sharply from both L. t. turcomanus and L.t.lehmanni. 

The taxonomic position of L.t. turcomanus is doubtful. Its color 
differs little from that ofL. t. lehmanni. Considering L.t. 
turcomanus as a synonym of L.t. lehmanni is a simple solution 
to this question. There is, however, a difficulty — the skull of L . t . 
turcomanus is small, and differs clearly from the more northern, 
L.t.lehmanni from the Aral Sea in its small measurements'^, and 
narrower rostrum. 

It is left open for the present whether L. t. turcomanus should be 
considered as a subspecies closely related to L.t. lehmanni, or as 
a natio of it, or whether it should be considered as a synonym of L . t . 
lehmanni. 

GEOGRAPHICAL DISTRIBUTION. The Syr-Darya desert hare is 
widely distributed throughout Turkestan. In the west it enters the Ust-Urt 
Plateau and apparently occurs as far as the lower reaches of the Emba 
river. It also occurs along the northeastern shore of the Aral Sea along 
the Syr-Darya, in the Golodnaya Step', in the Kyzyl-Kum Desert, on the 
Kara-Tau, Alexander Range and on the Talas Ala-Tau. In the east, it 
is sporadically distributed throughout the Dzhetysu [Semireche] plain""- to 
the mountains. A specimen from the vicinity of the village Rybach'ii on 
Lake Issyk-Kul is in my collection. Hares caught in the Zaisan Depression 
and near Dzharkent [Panfilov] belong to the same subspecies (natio 
zaisanicus). The eastern range of L . t . le hm ann i apparently 
includes the northwestern part of Dzungaria. In Turkestan it occurs 
near Tashkent, Fergana, along the upper reaches of the Amu-Darya, in 
Khiva Oasis, and in the northern Kara-Kum Desert. Hares from north 
of Dzhebel are closely related to L. t. lehmanni but their skulls show 
some differences and they apparently belong to a separate natio. 

This survey shows that the southern boundary of the range of L . t . 
lehmanni can not be exactly determined for lack of material and records. 

190 334'. Lepus (Eulagus) tibetanus lehmanni natio zai s an i c us 
Satun. (1907). Zaisan desert hare 

1907. Lepus zaisanicus Satunin K. , Ueber die Hasen Centralasiens, Ezhegodnik Zoologicheskogo 

muzeya Akademii Nauk, Vol. XI (1906), 1907, pp. 7-8. 
1922. Lepus tolai z a isanicus Ognev S. I. , Materialy po sistetnatike russkikh zaitsev (Materials on 

Taxonomy of Russian Hares), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, Vol. XXIII, pp. 483-484, 

TYPE LOCAUTY AND DEPOSITION. Vicinity of Zaisan (male Jan. 1891, collected by Pevtsov). Type 
in the ZMAN collection. 

MATERIAL EXAMINED: 14 specimens. 

DIAGNOSIS. Closely related to L.t. lehmanni Sev. , but differing 
from it by its large skull. The following skull characters are observed in 
series: l) very long incisive foramina; 2) bullae osseae more widely 
separate ventrally; 3) nasal bones longer. 



* See table of measurements. 
'■* I have seen a specimen from Alma-Ata. 



165 



Color of summer fur grayer and duller than in L. t. lehmanni. 
However, there are specimens of both forms which are indistinguishable. 
Color of winter fur ofL.t. lehmanni natio zaisanicus more whitish 
than that of L.t. lehmanni. 

Length of skull 83.8-90 mm; zygomatic width 40-43.7 mm; length of 
nasals 34.1-41.7 mm; width of nasals 15-20.3 mm. 

Length of body and head 443-490 mm; hind feet 120-128 mm; ear 
(measured along concavity) 83- 113 mm; tail 88-116 mm. 

SYSTEMATIC NOTES. As this hare is closely related to L . t . 
lehmanni, I intended to make zaisanicus a synonym of le hm ann i. 
However, a careful examination of the skull showed that there are some 
differences between the Aral and Zaisan desert hares. The color 
characters are of so little importance that they can be ignored. Therefore, 
until more material becomes available I suggest considering zaisanicus 
as a natio of le h m an ni . 

I have noticed that specimens of this subspecies become larger towards 
the north, and smaller towards the south. For instance, the cranial 
differences between zaisanicus and hares from Ak-Kuyu (north of 
Dzhebel) described by W.G. Heptner as Lepus europaeus 
turcomanicus (see above) are much more marked than those between 
specimens from the northeastern shore of the Aral Sea and the 
Zaisan specimens. 

If new material would confirm that the skull of L. t. turcomanicus 
is smaller than that of lehmanni, this form could be considered as a 
separate natio. 

All these questions, can be solved only after examination of more 
material. 

GEOGRAPHICAL DISTRIBUTION. This form is widely distributed in 
the Zaisan Depression. K.A. Satunin (1907) described what he thought to be a 
191 new "species" from Zaisan (collected by N. V. Pevtsov's expedition), and 
also from Kendyrlik='- (collected by Kolomeitsev). G.I. Polyakov (1909) 
obtained a small series of Zaisan hares on the banks of the Kara-Irtysh 
in the Zaisan Depression. A good series of Zaisan hares was collected 
by B. A. Kuznetsov near the village Kara-Bulak (former Semipalatinsk 
Region of the Zaisan District). 

Specimens caught by P. Rohdendorf (1925) correspond in their skull 
structure to this natio. This apparently also occurs in Dzungaria, at 
least in its northwestern part. 

335. Lepus (Eulagus) tibetanus quercerus Hollister (1912). 
Chuya hare 

1912. Lepus quercerus Hollister N. , Proceedings of the Biolog. Society of Washington, Vol. XXV. 

p. 181-184. 

TYPE LOCAUTY AND DEPOSITION. Chuya Steppe, No. 175446, male adult, 28 July 1912, N. 
Hollister. Type in the U. S. National Museum. 

MATERIAL EXAMINED: 6 pelts and 3 skulls. 



* Kendyrlik lies east of Zaisan, near the frontier of the U. S. S. R. 



166 



DIAGNOSIS. General color of back straw colored (intermediate between 
pinkish-buff and avellaneous, plate XXIX and XL), strongly darkened by 
black or black-brown subapical bands of the hair; tips of hair pale straw colored. 

Top of head straw colored (pinkish buff) with strong brown mottling 
which gives a dark appearance to the head. Color of neck pale straw with 
a dark pinkish tinge (light pinkish cinnamon). A straw-yellow (pinkish 
buff) stripe across the chest. Anterior-lateral parts of muzzle gray, 
cheeks either straw-yellow or ash-gray. Eye surrounded by a broad, 
whitish ring. 

Color of anterior inner margin of ears varying. In one specimen it is 
gray with slight blackish brown mottling. In another specimen, the inner 
margin is pale pinkish straw (intermediate between pinkish buff and light 
pinkish cinnamon) and chestnut -brown outwards. The external fringe of 
the inner margin with a well marked whitish stripe. Posterior sides of 
ears whitish with a light straw tinge. Black-brown tips of ears narrow, 
poorly developed and better marked on the outside. 

Flanks gray with a well marked straw tinge (pinkish buff). Anterior 
side of legs pale straw colored. 

Length of skull 84.9-87. 2 mm; zygomatic width 40.2-44 mm; length of 
nasals 36. 7-38. 1mm; interorbital width 13. 2-14. 9mm; length of upper 
molars 13-13. 2mm. N. Hollister gives the following measurements for 
the type: Length of skull 82mm; zygomatic width 41.5mm; length of 
nasals 34.2mm; interorbital breadth 14.8mm; length of upper row of 
molars 14.3 mm. 

Measurements of specimens examined by us for comparison with those of 
Hollister: 



192 



- 


No. 825, Feb, 1936, 
Chegan-Burgazy 


No. 57 o", 30 June 1936, 
Near Kosh-Agach 


cad, 2 July 1912 
According to Hollister 




Length (in mm. ) 


Body 

Tail 

Hind foot 

Ear 

Weight (in grams) 


460 
86 

110 

90 

2520 


460 

86 

110 

90 

1980 


470 

86 

115 



ADDITIONAL DESCRIPTION AND SYSTEMATIC NOTES. A young 
hare obtained by A.M. Kolosov (11 July 1936) in Chegan-Burgazy, Chuya 
Alps (No. 150 male) has the following measurements: body length 95mm 
[sic]; ear 84 mm. It has a pale gray- straw -yellow color on the back with 
a marked blackish brown tinge, caused by black-brown guard hair and 
black-brown tips of some guard hairs which have pale straw-white subapical 
bands. Color of back mottled. 

Color of top of head similar to that of back. Occiput with a distinct 
white spot. Sides of muzzle whitish gray. Cheeks straw-gray. Eyes 
surrounded by a well marked white ring. Color of occiput pale, 
yellowish straw (pinkish buff, Ridgway, 1912). Flanks pale gray with a 
straw tinge. A dull straw-yellow (pinkish buff) stripe across the chest. 
Anterior inner margin of ears dull gray-straw-yellow. A straw-white 
fringe on the outer side of the inner margin of the ears. Posterior sides 
of ears straw -white which is better marked on the proximal parts of the 
outside of the ears. Brown-black tips well marked on the posterior side. 



167 



Color of young (body length 250 mm, weight 150 grams) as follows: back 
pale straw -gray- ye How with weak blackish brown mottling. Color of 
top of head like that of back. Vertex with a well marked oblong white spot. 
Anterior-lateral parts of muzzle pale gray. Cheeks more ash-gray. Eye 
surrounded by a broad white ring. Flanks pale gray with a light straw 
tinge. A dull straw-gray stripe across the chest. Anterior inner parts 
of ears straw colored with strong blackish brown mottling. Tip of inner 
side of ears with well developed straw tones (pinkish buff) giving the tips a 
very marked color. Posterior side of ears whitish. Tip of ears on the 
posterior side with a narrow blackish fringe. 

N. Hollister (1912) gives the following description of the type of 
Lepus quercerus: "General color of dorsum straw-yellow (pinkish 
buff) with a black and well marked vinaceous -straw tinge (vinaceous-buff). 
Sides of muzzle and bases of ears gray. Anterior inner part of ears pale 
isabella color. Tip of anterior inner part of ear straw colored. Posterior 
side of ears almost white. Inner parts white. A black-brown stripe along 
two-thirds of the inner side of the ears. Fringe of inner side straw 
colored (buff). Tips of ears blackish, color descending 15mm along the 
posterior side of the ear, forming also a narrow margin along the anterior 
inner side. Occiput brownish ocher (clay color). Hindquarters grayer 
than back with well developed straw and vinaceous-straw tone. Tail 
white with greater part of dorsal side black. External side of legs pale, 
rust-straw colored (pale cinnamon). Stripe across chest brownish ocher. 
Remaining underparts including inner sides of legs white. " 

Hollister's description agrees well with the Chuya specimens obtained 
in 1936 by A.M. Kolosov. 

There are some differences between the Chuya hares and the Zaisan 
and Semipalatinsk specimens (natio z a i s an i c u s ). Back of Chuya hares 
blacker. Straw tone on flanks more marked. Ash-gray tones marked 
on the anterior-lateral part of the muzzle. Whitish ring around eyes more 
marked. According to Hollister, the ears of L e p u s quercerus are 
longer than those of zaisanicus . However, he does not mention the 
length of the ears of his new species. We did not find any differences 
between our specimens and the Zaisan-Semipalatinsk specimens in this 
respect or in skull structure. General appearance and structure of 
skull is subject to individual variations in Central Asian hares. 

The material at our disposal is insufficient to determine the systematic 
position of the Chuya hare exactly. Furthermore, the adults obtained 
by A.M. Kolosov from the Chuya Steppe had not finished their molt in 
spite of the late date (June-July). Their fur does not, therefore, consist 
completely of fresh summer hair. It is thus difficult to use this material 
for systematic purposes. 

L. quercerus (a subspecies of t i be t anu s ) is thus apparently 
distinguishable from zaisanicus in some color characters but their 
constancy and distinctness should be verified on a large series. 

336. Lepus (Eulagus) tibetanus subsp. Kopet-Dagh desert hau-e 

1928. Lepus tolai craspedotis Ognev S. I. and Heptner W. G. , Zoologischer Anzeiger, LXXV, S. 
261; Ognev S.I. and Heptner W. G. , Mlekopitayushchie srednego Kopet-Daga i prilezhashchei ravniny 
(Mammals of the Central Kopet-Dagh and the Adjacent Plains), Trudy Nauchno-issledovatel'skogo 
jnstituta zoologji, III (1): 71-73. 1929. 



168 



1934. Lepus europaeus turcomanus (partim.') W. G. Heptner, System, und tiergeogr. Notizen 
Uber Saugetiere. Folia Zoologica et Hydrobiologica, Riga, Vol. VI, No. 1, S. 21-23. 
TYPE LOCALITY. No. M. 3064 male, 9 June 1925, near the village Verkhneskobelevskii, Kopet-Dagh, 

Transcaspia. S.I. Ognev's collection. No. M. 3065 male, 17 August, 1925, near the village Sulyukli, 

Kopet-Dagh, S.I. Ognev's collection. 

MATERIAL EXAMINED: 2 further pelts from the Kopet-Dagh and Bol'shie Balkhany Range. Total 4. 

specimens. 

194 DIAGNOSIS. General color of back of summer fur straw-gray with a 
well marked dark m.ottling which gives a gray tinge to the general color 
of the back. Lower part of back grayer. Flanks with marked dull straw- 
brown tones intermediate between avellaneous and wood-brown. 

Top of head straw-gray, sometimes with well marked blackish mottling. 
Neck dull, straw-brown (intermediate between avellaneous and wood-brown). 

Chest dull straw-gray with a light reddish tinge (close to avellaneous). 

Color of anterior inner margin of ears straw-gray with well marked 
blackish mottling which gives the general impression of blackness. A 
white marginal stripe along the outer fringe of the inner side. Posterior 
side of ears whitish with an admixture of blackish hair. Black-brown tip 
of posterior apical part of ears well developed. Anterior side of legs 
straw-gray. 

Length of skull 84-86 mm; zygomatic width 38-42 mm; length of nasals 
36-37 mm; width of nasals 15-16 mm. 

Length of body and head 435-460mm; hind feet 121mm; ear (measured 
along concavity) 108-119mm; ears (measured from occiput) 139-I49mm; 
tail 75-87 mm. 

SYSTEMATIC NOTES. The Kopet-Dagh hares resemble L . t . 
lehmanni closely in their color. They differ from it as follows: l) 
grayer tone on the back; 2) duller color of chest; 3) black anterior inner 
margin of the ears. 

The skull of the Kopet-Dagh hares is smaller than that of L . t . 
lehmanni in the small material examined. Rows of molars especially 
small and weakly developed. Length of upper row of molars 14-14.9 mm 
in our specimens, in L.t. lehmanni 15-16.9 mm. 

W.G. Heptner and myself (1929) provisionally identified the Kopet-Dagh 
hares as L. t. craspedotis which was described by Blanford ("Annals 
and Magazine of Natural History", 1875, ser.4, p. 313 and in "Eastern 
Persia", II, Zoology, 1876, pp. 80-82, p. VIII) from Baluchistan (Pishin). 

The Kopet-Dagh hares are certainly closely related to Baluchistan hares 
in some characters, e.g., the general gray color, black color on the ears, 
length of ears, etc. 

However, the Baluchistan hares are even darker than the Kopet-Dagh 
hares. Blanford unfortunately does not mention the date of collection of 
his holotype. He possibly used a specimen in winter fur, which is darker 
and blacker . However, the far removed range of L.t. craspedotis 
makes it doubtful whether this hare is identical with the Kopet-Dagh hare. 

The relation of the Kopet-Dagh hare to L. europaeus turcomanicus 
Heptner has been discussed above. 

GEOGRAPHICAL DISTRIBUTION. The Kopet-Dagh hare is apparently 

Q restricted to the foothills and mountains, and does not occur in the plains. 

During the Transcaspian expedition (1925) we caught the first specimen in 

tamarisk thickets near the village Verkhneskobelevskii. However, hares 

were rather rare in this region. Near Germab they are more common. 



169 



In the mountains near the village Sulyukli (altitude 5000-6000 feet) they 
are very abundant in some years. They keep mainly to screes and juniper 
thickets in this region. S.I. Bil'kevich obtained them in the Bol'shie 
Balkhany Range in juniper forest, i.e. , rather high in the mountains. 

337. Lepus (Eulagus) tibetanus desertorum Ogn. et Heptner 
(1928). Desert sand hare 

1886. Lepus lehmanni Niko'skii A. M. , Materialy k poznaniyu fauny pozvonochnykh severe- vostochnoi 

Persii i Zakaspiiskoi oblasti (Data on the Vertebrate Fauna of Northeastern Persia and Transcaspia), 

Trudy Sankt-Peterburgskogo obshchestva estestvoispytatelei, p. 7. 
1890. Lepus lehmanni Radde, G. , Walter, A., Blasius, W. , Die Saugethiere Transcasp. , Tiflis, 

pp. 62-64. 
1894. Lepus lehmanni Varentsov P. , Fauna Zakaspiiskoi oblasti (Fauna of Transc.ispia), p. 16, 17. 
1918. Lepus lehm anni Bil'kevich S. I. , Kollektsiya Zakaspiiskogo muzeya (The Collection of the 

Transcaspian Museum), hvestiya Zakaspiiskogo muzeya, I, p. 9. 
1928. Lepus tolai de se rt orum Ognev S. I. und Heptner W. G. , Einige Mitteilungen iJber die 

saugethiere des mittl. Kopet-Dagh etc. , Zoologisch. Anzeiger, LXXV, S. 262. Ognev S. I. and Heptner 

W. G. , Mlekopitayoishchie srednego Kopet-Daga i prilezhashchei ravniny (Mammals of the Central 

Kopet-Dagh and the Adjacent Plain), Trudy Nauchno-Issledovatel'skogo institute zoologii, IIl(l):73-75. 
1934. Lepus europaeus des e rt or u m Heptner W. G. , Systemat. und tiergeogr. Notizen iiber Saugethiere 

Folia Zoologica et Hydrobiol. , Vol. VI, No. 1, S. 22-23. 

TYPE LOCALITY AND DEPOSITION. M. 3066, female, 2 May 1925, near railway station Annau, 
Transcaspia. S. I. Ognev's collection. 

MATERIAL EXAMINED: 9 pelts and skulls. 

DIAGNOSIS. Differing from L. t. lehmanni in its very pale straw- 
sand color of the summer fur with a few blackish hairs. Winter fur on 
back with distinct, irregular blackish mottling on a light straw-yellow 
background. Summer specimens markedly lighter and not as gray as 
specimens caught in winter. 

Length of skull 76.6-84 mm; zygomatic width 37-40.7 mm; length of 
nasals 31. 8-37. 5mm; width of nasals 15. 6-18. 5mm. 

Length of body and head 399-460 mm (M. 431.3 mm); hind paws 102- 
120 mm (M. 118 mm);' ear (measured along concavity) 98.2-105 mm; ear 
(measured from occiput) about 120mm or more; tail 79-100 mm. 

These measurements show that the skull ofL.t. desertorum is 
smaller than that of L.t. lehmanni. 

ADDITIONAL DESCRIPTION. General color of summer fur on back 
very pale straw-sand colored with a light reddish vinaceous tinge 
(intermediate between avellaneous and vinaceous -buff). 

Blackish mottling on back little marked as there is little long black 
guard hair and the blackish hair tips are also little developed. Roots of 
guard hair grayish, followed by broad straw-sand colored bands and short 
blackish brown tips. 
196 Top of head very pale, straw-sand colored. Muzzle with a whitish tinge. 
Eyes surrounded by a whitish ring. Color of neck pale straw-pink 
(vinaceous -buff ) . 

Color of chest very pale, straw-yellow (intermediate between pale 
pinkish buff and pinkish buff). 

Color of anterior inner margin of ears pale, straw-yellow (intermediate 
between vinaceous-buff and pinkish buff), without any marked black or gray 
color, because of the almost complete absence of blackish hair. Outer 
side of inner margin of ears with a broad white fringe. Inner side of ears 



170 



whitish, with relatively few hairs and the skin clearly visible. Posterior 
sides of ears light whitish straw-gray. 






FIGURE 102. Skull of Le pus 
tibetanus desertorum Ogn. 
M. 3066, from Annau (near 
Ashkhabad), holotype, S. I. Ognev's 
collection 



HGURE 103. SkullofLepus 
tibetanus buchariensis 
Ogn. M. 3660, 22 May 1914, 
Khatyn-Rabad, N. A. Pobrinskii, 
S. I. Ognev's collection 



FIGURE 104. Skull of Lepus 
tibetanus subsp. , No. M. 
3065, Sulyukli, Kopet-Dagh, 
S. I. Ognev's collection 



■^grj Anterior side of legs very pale, straw-yellow. 

Belly and groin white. Tail white, with black dorsal side. 

The winter fur differs rather sharply from the summer fur. Its hair is 
much longer and fluffier than in summer fur. Blackish parts of hairs 
broader and more distinctly marked. This gives a rich color and a 
characteristic variegation. Rust tones on occiput and chest well marked. 
Color of occiput intermediate between avellaneous and pinkish cinnamon. 
Color of chest duller. 

General color of winter fur variable. There are specimens with more 
yellowish fur, or with fur of a more pale gray color. 

SYSTEMATIC NOTES. The desert sand hare differs clearly in series 
from L.t.lehmanni in its generally pale color. These differences are 
very distinct in series of summer and winter specimens. 

GEOGRAPHICAL DISTRIBUTION. I shot the first specimen of this 
subspecies in a semidesert with stunted vegetation near the railway station 
at Annau (Transcaspia). I also examined pelts and skulls from near 
Geok-Tepe, near the railway station at Sary-Yazy, near Chatlov (on the 
Atrek), near Uch-Adzhi, near Chelayun 35 km east of Gasan-Kuli, and near 
Karabat. 

The distribution ofL.t. desertorum thus includes the southern 
desert and the sand dunes of Transcaspia (from the southeastern corner 
of the Caspian Sea to the Amu- Darya, to about Chardzhou). The range thus 
includes the southern Kara-Kum desert. 



171 




FIGURE 105. Skull of Lepus tibetanus desertorum Ogn. No. M. 3066, 
Annau, near Ashkhabad 




FIGURE 106. Skull of Lepus tibetanus buch ari ensis Ogn. M. 3060, holotype 

297 338. Lepus (Eulagus) tibetanus buchariensis Ogn. (1922). 
Buchara desert hare 

1823. Lepus tolai Lichtenstein H. , Naturg. Anhang zu Eversmann's Reise nach Orenburg u. Buchara, 

S. 118. 
1922. Lepus tolai buchariensis, Ognev S. I. , Materialy po sistematike russkikh zaitsev (Data on the 

Systematics of Russian Hares), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, Vol. XXIII, p. 475. 
1922. Lepus tolai buchariensis, Ognev S. I., Nekotorye dannye po sistematike nisskikh zaitsev 

(Data on the Taxonomy of Russian Hares), Biologicheskie izvestiya, Vol.1, pp. 102-104. 

TYPE LOCALITY AND DEPOSITION. Khatyn-Rabad, east of Termez. No. M. 3060 male, 22 May, 1914, 
N. A. Bobrinskii, type in S. I. Ognev's collection. 

MATERIAL EXAMINED: 1 1 specimens. 

DIAGNOSIS. Differs from L . t . de se rtorum in the distinct rust -red 
tinge of the pale sand-straw colored back. The longer and softer fur of 
winter specimens has a distinct brownish (a blackish brown) mottling. 
Reddish rust shades are also well marked on the ears of many specimens. 

Length of skull 79-88.3 mm (M.84mm); zygomatic width 38.6-42.2 mm 
(M. 40.5mm); length of nasals 32.2-41 .7 mm (M. 36.6 mm); width of nasals 
13.2-16mm (M. 14.5mm). 
198 Length of body and head 460-530 mm (M. 485 mm); hind feet 102-119 mm 
(M. 112.7 mm); ears (measured from occiput 95-117 mm (M. 123 mm) [sic]; 
tail 78-120mm (M. 109.1mm). 

ADDITIONAL DESCRIPTION. Summer fur. General color of back 
pale, sand-straw colored with a distinct rust-red tinge (intermediate 



172 



between avellaneous and light pinkish cinnamon), with a small admixture 
of long, black guard hair which causes a grayish mottling. Color of 
ordinary guard hair: roots grayish, followed by pale rust-straw 
colored bands and brownish black tips. Color of underfur whitish with a 
rust-straw tinge. Color of top of head similar to that of back but its 
general color is sometimes paler and yellower. Color of anterior part of 
muzzle and cheeks pale straw with a light rust tinge. Eyes surrounded by 
a whitish ring. 

Neck reddish rust-straw colored (intermediate between pinkish 
cinnamon and light pinkish cinnamon). 

Chest of the same reddish rust -straw color, its color sharply different 
from that of the white chin, belly and middle part of groin. A pale rust- 
straw colored stripe on the sides of groin, extending to the inside of the 
hind legs. This stripe stands out clearly on the white background. 

Similar rust-straw colored fur becomes straw-reddish rust colored 
(intermediate between light pinkish cinnamon and pinkish buff) on the 
flanks . 

Color of anterior inner margin of ears pale straw-yellow with a light 
rust and grayish tinge. Outer fringe of inner margin of ears white, 
sometimes a little yellowish. Inner side of ears whitish. A distinct straw- 
rust tinge along the outer margin of the ears. Posterior side of ears 
whitish gray. Posterior apical margin of ears with a blackish terminal 
fringe. 

Anterior side of legs straw-reddish rust colored. 

Tail white. Color of the dorsal side blackish brown. 

The winter fur has longer and softer hair than the summer fur, and the 
brownish and gray brownish mottling on the back is better developed. 
There are no marked differences between the color of summer and winter 
fur in this subspecies. 

SYSTEMATIC NOTES. The coloration ofL.t.buchariensis is 
rather constant, but there are variations both towards stronger development 
of rust-red shades, and towards more reddish and yellowish shades. 

It differs from L. t. lehmanni in the following characters: l) paler 
sand- straw -rust color of the back which is more grayish in L . t . 
lehmanni; this is especially marked in the winter fur; 2) color of neck 
in L . t . lehmanni duller and darker; 3) color of legs rich rust-straw 
colored; in L. t. lehmanni, the legs are pale gray; 4) color of ears in 
L. t. buchariensis paler yellow than inL.t. lehmanni, with rust 
tones which are absent in L. t. lehmanni. 

The range of the Bukharian subspecies is adjacent to that of L . t . 
desertorum. 

GEOGRAPHICAL DISTRIBUTION. N.A. Bobrinskii's expedition 
obtained large numbers of this subspecies near the village Sardab-Buzachi, 
and near Khatyn -Rab ad (east of Termez). L.t. buchariensis lives 
on sandy and on sand-clay-ground in tamarisk thickets. It is apparently 
widely distributed in Tadzhikistan, from where it enters northeastern 
Afghanistan. 



173 



339, Lepus (Eulagus) tibetanus pamirensis Blanford (1875). 
Pamir desert hare 

1875. Lepus pamirensis Blanford W. T. , Journal Asiatic Society of Bengal, XLIV, p. 110. 

1879. Lepus pamirensis Blanford W. T. , Scientific Results of the Second Jarkand Mission, Mammalia, 

pp. 67-68. 
1879. Le pus sp. Severtsov N. A. , Zametki o faune pozvonochnykh Pamira (Notes on the Vertebrate Fauna 

of the Pamir) Zapiski Turkestanskogo otdeleniya geograficheskogo obshchestva, I, p. 64. 
1886. Lepus lehmanni Grum-Grzhimailo G. E. , Ocherk Pripamirskikh stran (A Survey of the Countries 

Adjacent to the Pamir), Izdatel'stvo Geograficheskogo obshchestva, XXII, p. 92. 
1935. Lepus tolai pamirensis Rozanov M. P. , Materialy po mlekopitayushchim Pamira (Data on 

Mammals of the Pamir), Tadzhikskaya kompleksnaya ekspeditsiya 1932 goda, Issue XXXII, pp. 53-54. 

TYPE LOCALITY. Lake Syry-Kul, Pamir. Deposition of type unknown. 

MATERIAL EXAMINED: 6 pelts. 

DIAGNOSIS. Differs clearly from the other subspecies of the desert hare in 
that its summer fur is very soft and delicate and its winter fur is even more 
so. General color of summer fur on back, pale straw-gray with a light 
vinaceous tinge (intermediate between reddish gray and vinaceous buff). 
Flanks yellowish white. Chest very pale straw-yellow. Inner margin of 
ears with a marked chestnut-brown tone which gives this part of the ear a 
dark appearance. 

Winter fur with conspicuous grayish brown, sometimes blackish, 
mottling. Specimen No. S. 5004, in the collection of the Moscow 
Zoological Museum, shows a marked black-brown color in the middle of 
the back, on head and ears. 

Length of skull 83-89 mm; zygomatic width 40-41.4 mm; length of 
nasals 34. 3-39. 8mm; width of nasals 14.2-16.3 mm. 

ADDITIONAL DESCRIPTION. General color of summer fur on back pale 
straw-gray with a slight vinaceous tinge (intermediate between drab-gray 
and vinaceous -buff). Black hairs few in number, giving a grayish tinge. 

Color of top of head resembling that of back, but grayer. 

Anterior part of muzzle and cheeks with a whitish shade. Eyes 
surrounded by straw -white rings. 

Occiput very dull, pale straw-rust colored, intermediate between 
light pinkish cinnamon and vinaceous -buff. 

Chest very pale, straw-yellow (intermediate between pale pinkish buff 
and pinkish buff). 
200 Flanks more whitish and lighter than back. Belly and groin white. 

Color of anterior inner margin of ears chestnut-gray-brown (between 
hair-brown and benzo-brown). Fringe of inner margin whitish. Inner 
side of ears whitish gray. Posterior side of ears dirty- straw -white. 
Apical part with a brownish-black margin. 

Anterior side of legs pale straw yellow. 

Tail white, its dorsal side black. 

Winter fur more luxuriant and dense. Brownish gray mottling distinct 
on the general straw-yellowish gray background. 

There is an interesting specimen in the Moscow Zoological Museum 
caught on the Pamir by Voskoboinikov (1898) (No. S. 5004). The middle 
part of the back is deep black-brown, it has much black hair on top of the 
head, and only a straw -gray mottling on a general black-brown background. 
Flanks much lighter than back, straw-yellowish gray. Chest and neck 
pale, straw-rust colored. Belly and groin white. 



174 



Anterior inner margin of ears straw- gray with a strong admixture of 
black-brown hair. Outer fringe of anterior inner margins white, contrasting 
with the dark background. 

Anterior side of feet pale straw-yellow. 

Blanford (1879) describes L.pamirensis as follows: "General tone 
very pale, sandy-brown, almost Isabella colored on back and flanks. 
Hindquarters grayish white. Dorsal side of tail black. Anterior side of 
ears of the same color as the back. Outer margin of distal parts of ears 
black. Chest light reddish brown (rufous). Venter usually white. Fur 
fine, dense and soft. . . " 

It appears from this description that Blanford had specimens with light 
anterior inner margins of the ears. The dark color of these parts in our 
specimens cannot therefore be considered as a reliable and constant 
character. 

SYSTEMATIC NOTES. This high-mountain form is characterized by its 
luxuriant and soft fur even in summer. 

It differs clearly from other desert hares by the texture of its fur and 
some color characters. 

As mentioned above, specimens of Le pu s kaschgaricus Satun.from 
the Bol'shoi Yuldus and Tien-Shan (obtained by M.N. Przheval'skii) are 
closely related to the Pamir hare. 

GEOGRAPfflCAL DISTRIBUTION. According to N. A. Severtsov, this 
form is common on the Paraiir. In G. E. Grum-Grzhimailo's opinion (1886) 
it is very common in this region. M, P. Rozanov (1935) writes that the 
Pamir expedition found large numbers ofL.t.pamirensis in the 
Markansu area, beyond the Kyzyl- Art Pass in the Trans -Alai. 



Subgenus Lepus s. str. Linnaeus (1758) 

''"I 1904. Lepus (subgenus!) Lyon M. , The Hares and their Allies, Smithson, Miscell. Collections, 
Vol. I, parts 3-4, p. 394 and 334 (partim.'). 
1902. Lepus (group) Barrett-Hamilton H. , A History of British Mammals, p. 292. 

Head relatively large. Ears short, ll/lO — 1 1/8 as long as head 
(measured from occiput). 

Tail with terminal hair much shorter than hind foot. 

Tail whitish gray in summer fur, markedly gray on the dorsal side. 
Tail completely white '■= in winter fur. Teats 8. 

Winter fur sharply different from brownish gray summer fur in most 
forms'''*. 

Species of this subgenus completely white in winter. Only the tips of 
the ears remain black. Nasals measured from anterior corner to posterior 
outer corner as long as, shorter than, or rarely longer than frontal suture. 

Width of incisive foramina ll/8toll/4as long as the bony palate. 
Anterior lower premolar slightly inclined backwards. 

Geographical distribution. Boreal and arctic regions of Eurasia and 
North America t. 



In arctic forms, tail completely white both in sumnaer and winter. 

With the exception of Le pus hibernicus Pall. 

Small American hares of the group of Lepus americanus and its subspecies are often placed into the 

subgenus Lepus (e. g.,H. E. Anthony, 1928). This is incorrect in our opinion. These hares differ 

clearly from true Lepus in cranial and morphological characters. American hares of the group 

americanus belong to a separate subgenus Poe c i 1 ol ag us Lyon. 

175 



160. Lepus timidus Linnaeus (1758). Blue hare 

1758. Lepus timidus Linnaeus C. , Systema Naturae, editio 10, p. 57. 

NAMES: The mountain or blue hare (English); Nordischer Schneehase, Veranderlicher Hase (German); 
Le lifevre blanc ou le changeant (French). 

LOCAL NAMES: ushkan (Russian in Siberia); zaets' bilyi or bilyak [white hare] (Ukrainian); numala 
(Mordvinian); Koetsch, or lotets (Votyak, Permyak); meran (Mari, Cheremis); molgach (Chuvash); 
Koetsch (Komi-Zyryan); nummel (Lapp, on the Imandra); njammel (eastern Lapland); shaur, or shogur 
(Khanty-Ostyak, on the Ob); toegur (in Surgut); kuyan (Bashkir, Meshchera); shichve, or sheching (Mansi 
Vogul); nevu or n'y^va (Nenets along River Ob); neuko (along Pechora river); n'eva (Yurak); taivusi (Nenets 
along Taz river); nyaba (Nenets in the Turukhansk Territory); nyuma (Selkut, Ostyak in the Turukhansk 
Territory); teas' (Yurak); n'oa or nioa (on the Narym); n'omu (Nganasan, Tawgi); mundukan (Evenki, 
Tungus) ; kuobakh, kobak, dveleken, tavyskhan (Yakut) ; tuksaki (Evenki, in the Olekma-Vitim Territory); 
Chagechantu (Orochen); cholgoro (Odul, Yukagir); oiegon (tundra inhabitants, Odul-Yukagir); chyi, chyik 
(Nivkhi-Gilyakof the western Sakhalin Coast); ossik, or kanak (white) (Nivkhi-Cilyak of central and eastern 
Sakhalin); ossuki (Ainu on Sakhalin); toksa, or tochsa (Nanai, Goldi-Samagir); tochsa and golmochomg 
(Nanai, Goldi between Geong and Ussuri); gurmakho (in the region between Ussuri and Sungari); toksake 
(Birar, Manyagr, Orochen); milut (Nymyllan, Koryak), khalluk (Aleut); bilchuch (Kamchadal) ; melotal'gin 
(the first 1 pronounced softly; Chukchi). 

PRINCIPAL FIGURES: 1) Pennant Thomas, Zoologia Britannica, IV, p. 40, pi. 47, 1770; 2) Schreber 1. , 
Die Saugethiere, Erlangen, Bd. IV, 1792, S. 885-891, Taf. CCXXXV A (summer fur); Taf. CCXXXV B 
202 (^^intsr fur) ; 3) Bingley W. , Memoirs of British Quadrupeds, 1809, p. 308 (color pi ate)^; 4) Nilsson Sv. , 

Illuminerade Figurer till Skandinav. Fauna, 1831-1832, Vol.1, H. 6, pi. 22 (Lepus borealis varietas - 
entire animal in color), H. 6-7, pi. 16, 19 (Lepus borealis - entire animal in winter and summer fur 
in color); 5) Simashko Yu. , Russkaya fauna. II, Mlekopitayushchie (Russian Fauna. II. Mammals), 1851, 
pp. 785-787; pi. 60A,figs. 2-3 (summer fur in color); pi. 61, fig. 2 (winter fur in color); pi. 61, fig. 3 (skull); 
pi. 62, fig. 3 (winter fur in color). Lepus variabilis in color: pi. 62, fig. 1 (Lepus canescens in color): 
6) Middendorf A. Th. , Uber die als Bastarde Angesproch. Mittelformen zw. L. europaeus und. L. 
variabilis, Bullet, phys. -mathem. de I'Academ. Imp. de St. -Petersb. , IX, No. 14, 1851, pp. 25-29 
(drawing of skull, p. 29); 7) Blasius J. , Naturgeschichte d. Saugethiere Deutschl. , 1857. S. 420-425, 
Fig. 229 (structure of teeth and bony palate); 8) Meves W. , Holmgren A. E. , Atlas ofver Scandinaviens 
Daggdjur, Stockholm, 1873, pi. XV, Fig. 1-7 (Le pu s timidus L. - c anesc e ns et bor e al is in color); 
9) Brehm A. , Zhizn' zhivotnykh. II (Animal Life, Vol.11), 1874, p. 252; 10) Nordquist O. , Anteckninger 
och studier till Sibirska Ishafskustens Daggdjurs-Fauna, 1833, pp. 84-90, figs. 8-10 (skull ventral, dorsal 
and lateral); 11) Trouessart E. L,, Histoire Naturelle de la France, .Mammifferes, Paris 1885. p. 187, 
fig. 78 (entire animal); 12) Fokht K. , Mlekopitayushchee v opisaniyakh (Descriptions of Mammals), 
Moscow, 1885, p. 403 (half-tone plate of hare in winter fur); 13)- Flower W. H. , Lydekker R. , An Introduct. 
to the Study of Mammals, 1891, p. 493, fig. 218 (entire animal): 14) Brehm A. E. , Zhizn' zhivotnykh. 
II. (Animal Life, Vol.11), p. 664 (half-tone plate of entire animal); 15) Lydekker R. , A Hand-Book of the 
Brit. Mammalia, London 1895, pi. XXVUI (entire animal in color) ; 16) Kashchenko N. F. , Rezul'taty 
Altaiskoi ekspeditsii (Results of the Altai Expedition), Tomsk, 1899. pp. 57-59, pi. 11, fig. 4 (drawing of 
ears); 17) Forsyth Major, On Fossil and recent Lagomorpha, Transactions of the Linnean Society of London, 
Vol. VII, Zoology, 1896-1900, p. 468, fig. I-II (drawing of incisors, ventral); 18) Lyon M. W. , Classification 
of Hares and their Allies, Smiths. Miscellaneous Collections, Vol.1, pp. 3-4, 1903(1904). pi. LXXX and 
LXXXI , fig. 3 L . tschuktschoru m (skull dorsal and ventral) ; fig. 4L. variabilis -(skull); 19) 
Gake, V. , Zhivotnyi mir (The Animal World), Vol. I, 1901, pi. 29 (color plate); 20) Millais J. C. , The 
Mammals of Great Britain, Vol. Ill, 1906, pi. 50 ^color), pi. 51 (color), pi. p. 32 (biological scene), pi. 
p. 34; 21) Hilzheimer M. , Die Hasenarten Europas, Jahreshefte d. Vereins fur vaterd. Naturk. im Wiirtt. , 
1908, S. 385-387, Taf. V., Fig. 8a-c (skull, mandible, teeth); 22) Meerwarth H. und Soffel K. , 
Lebensbilder aus der Tierwelt, III, Leipzig, 1912, S. 64-65, 69, 71 (photograph of animals); 23) Miller 
G. S. , Catalogue of the Mammals of Western Europe, London 1912, p. 525, fig. 103 L. t . sc o t ic u s (skull 
lateral, ventral and dorsal); p. 526, fig. 104 (teeth); 24) Barrett-Hamilton G. , A History of British 
Mammals, 1912, part. XI, pi. XV (entire animal); p. 230, fig. 2 (skull lateral); p. 232, fig. 3 (palate); 
part XII, pi. XIX fig. 8 (hind leg); pi. XX, fig. 8 (hind leg); 25) Ognev S. I. , Mlekopitayushchie Moskovskoi 
gubernii (Mammals of the Moscow Province), 1913, pp. 298-307, pi. Ill, pp. 45-50 (seasonal variations cf 
fur); fig. 53 (skull of young blue hare); pi. IV, figs. 57, 60, 62 (photograph of skull); 26) Brehm's Tierleben, 
Saugetiere, II, 1914, S. 66 (half tone drawing of entire animal); 27) Filipchenko Yu. A. , Izmenchivost' i 
nasledstvennost' cherepa u mlekopitayushchikh (Variability and Heredity of Mammalian Skull), Russkii 
arkhlv anatomii, gistologii i embriologii, I (2), 1916, pi. XVI, fig. 13 (skull dorsal, ventral); 28) Thorburn 



176 



A. , British Mammals, Vol. II, 1921, pi. 34 (in autumn fur); pi. 35 (in winter fur); 29) Klshida K. , An 
Explanatory Illustration of Japanese Mammals, ed. II, 1926, pi. II (in Japanese); 30) Wollebaek A. , Norges 
Pattedyr, Oslo 1927, p. 22, fig. 17 (skull lateral); p. 23, fig. 18 (entire animal); 31) Kaverznev B. N. , O zaitse 
i ego dobyvanii (The Hare and its Capture), Moscow 1928, p. 6 (track of a blue hare); p. 7 (tail of blue hare, 
its color compared with that of the common hare, a litter of the blue hare), p. 27 (running blue hare); 
32) Bobrinskii N, A. , Opredelitel' okhotnich'ikh i promyslovykh zverei (A Key to Game and Commercial 
Animals), 1928, p. 41, fig. 25 (drawing of tail); 33) Abe J. , A Synopsis of the Leporine Mammals of Japan, 
Journ. of Science of the Hiroshima University, ser. B, Div. I (Zoology), Vol.1, 1931, p. 48, fig. 1 
(Lepus ainu); p. 50, fig. 2 (Le pus timidus saghaliensis), pi. 1 (Skulls of Le pus ainu); pi. II 
(skull ofL.t, saghaliensis); 34) Stakhrovskii V. G. , Zayats-belyak (The Blue Hare), in the book: 
Verkhnevychegodskaya ekspeditsiya pod nachal'stvom S. V. Lobacheva (The Upper Vychegda Expedition under 
the Direction of S. V. Lobachev), Issue I, Moscow 1932, pp. 176-195, fig. 34 (an ensnared hare); 35) Heptner 
W. G. , Zaitsy (Hares), Izdatel'stvo Soyuzpushniny, Moscow 1933, p. 4, fig. 1 (blue hare in winter fur - an 
excellent drawing); 36) Vinogradov B. S. , Mlekopitayushchie SSSR (Mammals of the U. S. S. R. ), Leningrad, 
20c 1933, p. 85, fig. 76 (A), fig. 77 (A) (bony palate and skull dorsal); 37) Manteifel' P. A. , Zaitsy SSSR (Hares 
of the U. S. S. R. ), in the book: "Biologiya zaitsev i belok" (Biology of Hares and Squirrels), Moscow 1935, 
p. 18, fig. 2 (posture of female during copulation); p. 27, fig. 4 (one-colored blue hare); 38) Bobrinskii N. A., 
Opredelitel' zverei SSSR (Key to Animals of the U. S. S. R. ), Moscow 1935, p. 50, fig. 40 (blue hare in winter 
fur); 39) Aspisov D. I. , Zayatsbelyak (The Blue Hare), Volzhsko-Kamskaya zonal 'naya stantsiya. Issue IV, 
1936, p. 45, fig. 2 (embryos); p. 118, fig. 15 (diagrams of molt); p. 122, fig. 16 (photograph of molting 
specimens in spring); p. 125, fig. 17 (diagrams of molt); 40) Kolyushev 1. 1. , Mlekopitayushchie krainego 
severa zapadnoi i srednei Sibiri (Mammals of the Extreme North of Western and Central Siberia), Trudy 
Biologicheskogo nauchno-issledovatel'skogo instituta, Tomsk, 1936, pi. I-II (photograph of skulls of L. t. 
begitschevi and L. t. sib iricorum ); 41) Gureev A. A. , Kraniologicheskie priznaki zaitsa belyaka i 
rusaka (Cranial Characteristics of the Blue and Common Hare), Trudy Zoologicheskogo instituta Akademii 
Nauk, III, 1936, pp. 356, 357, 362, figs. 1, 3, 5, 7 (skulls and details of their structure). 
MATERIAL EXAMINED: 205 pelts and 246 skulls. 

DIAGNOSIS. Head relatively large. Ears short, 1 l/lO to 1 1/8 as long 
as head (measured from occiput). Tail with terminal hair much shorter 
than hind foot (without claws). 

Tail whitish gray in summer fur, markedly gray on dorsal side. 
Tail completely white in winter. 

Winter fur of most formis sharply different from summer fur (except for 
Lepus hibernicus Pall,). Hares of the subgenus Le pu s completely 
white in winter, only the tips of the ears remain black. 

Summer fur coarse, guard hair dense, straight, not curling on back, 
Underfur curling. Hair of winter fur softer, longer and more delicate than 
that of summer fur, Underfur in winter much more luxuriant. 

Teats: p. 1-1; a. 3-3, totals. 

Nasals measured along oblique line connecting anterior inner corner to 
outer posterior corner as long as, shorter than, or rarely longer than 
frontal suture. 

Incisive foramina in their posterior third with rounded protuberances 
which are sometimes little marked. 

Ratio of length of bony palate (measured lateral to the anterior process) 
to width of incisive foramina in their posterior third 70,24%, 

Ratio of length of bony palate to maximal width of mesopterygoid fossa 
72.7%. 

Ratio of length of row of lower molars to lower diastema 86.9%. 

Lower anterior premolar slightly inclined backwards. 

Mandibular condyle large, slightly inclined backwards. 

Posterior-ventral plate of mandible (from angular process to notch at the 
beginning of the horizontal process) broad, longer than the distance between 
anterior alveolus of first premolar to tip of lower incisor. 
204 Bullae osseae small and moderately inflated, slightly protruding 
anteriorly in profile. 

177 




FIGURES 107-109. Skull of blue hare, Lepus timidus kozhevnikovi Ognev. M. 3167, 17 November 
1922, Shipov Experimental Forestry near Pavlovsk, S. I. Ognev's collection 

Drawing by Yu. A Kostylev 



178 



204 Length of ear (measured from occiput) 110- 118 mm; length of ear (along 
concavity 77- 115 mm; hind feet 132-I90mm. 

ADDITIONAL DESCRIPTION. Skull not as elongate and slender as in 
the common hare. It appears shorter because of the massive and wide 
zygomatic arches. 

The characteristic differences between the skulls of the blue and common 
hare have been given above. The orbit is delimited by massive, zygomatic 
arches which rise steeply backwards. The zygomatic arch therefore forms 
a sharper angle at the anterior edge of the orbit than does that of the common 
hare. In series, or even in individuals, the species of a hare may be 
determined from the form of the orbit. 

Nasals generally shorter than in the common hare. The ratio of length 
of the bony palate to the width of the incisive foramina and the 
mesopterygoid fossa gives reliable characters for identification. 

The mandible is very massive. 

Another reliable difference in the skull of the blue hare is the more 
vertical position of the lower premolar which is markedly inclined 
backwards in the common hare. 

For biometrical differences between the skulls of common and blue hares, 
see description of the common hare. 

COLOR. The color of summer fur of blue hares varies greatly in 
different localities. The color of most northern blue hares is darker and 
grayer than that of specimens from southern localities. Northern 
specimens also show marked blackish tones on the ears. We shall 
describe the color of the typical Moscow blue hare (L. timidus 
kozhevnikovi) as an example. We shall then describe the differences 
between this and a specimen of the nominate race (L. timidus) from 
northeastern Europe. 

The color of an old Central Russian blue hare in summer fur is as 
follows. Chin and lips white. Anterior part of muzzle, cheeks and region 
around the eyes rich rust -brown (usually between cinnamon and sayal- 
brown, Ridgway, 1912, pi. XXIX). An admixture of black hair gives a 
duller tinge to the general tone in these parts. Black hairs become very 
numerous on the forehead, vertex and between the ears. Hair on forehead 
and vertex of several colors. In this region are guard hairs with gray-black 
roots, rust-colored bands in the middle and blackish tips. There are also 
completely black guard hairs in this region. Hair of underfur with gray 
roots and dull rust-colored tips. General tone of back rust -brown with 
inconspicuous grayish black mottling. Fur of back much longer than that 
of head but of the same color. There are three types of guard hairs: l)most 
numerous and typical are guard hairs with blackish gray roots, broad 
cinnamon-buff middle bands and blackish tips; 2) thinner guard hairs with 
paler and not so well developed middle bands; 3) the longest and coarsest 
guard hairs are completely black. Underfur on back curling, with 
grayish roots and very pale rust-colored tips. 

Color of lower part of back near tail, flanks and a broad belt across 
the neck and upper part of the chest more gray than the parts mentioned 
above. The fur of these parts consists of guard hairs with gray roots, pale 
straw-gray middle bands and blackish tips. Longer guard hairs are also 
present here which are either completely black, or have narrow pale straw- 
gray subapical bands. Underfur dense, very curly, straw-smoke-gray. 



179 



206 




FIGURE 110. Mandible of Le pus timidus kozh e vniko vi Ognev. M. 3167, 
17 November 1932. Shipov Experimental Forestry near Pavlovsk, S. I. Ognev's 
collection 

Drawing by Yu. A. Kostylev 

Anterior side of fore- and hind legs with rust-brown hair with a strong 
admixture of black hair. Soles of hind feet with long fluffy dirty-gray fur. 
Tail uniformly gray. 

Color of the anterior inner margin of the ears generally rust-brown. 
Inner sides of ears whitish gray. Outer margin with a dull rust-gray 
oblong patch. External margin of the ears with a white fringe. Posterior 
side of ears dirty brownish gray. Tips of ears blackish. 

Vibrissae on cheeks and muzzle black and white. Color of longest 
vibrissae either black or white. There are occasionally long black 
vibrissae with white tips. 
207 In the northern blue hare, the color of cheeks and anterior part of muzzle 
is dull straw-rust-gray. Color around the eyes the same. Forehead and 
occiput with a strong admixture of brownish gray hair. Hairs on these parts 
have broad blackish brown roots, narrow, pale rust-gray subapical bands 
and black-brown tips. There also are completely black guard hairs. 
General color of back dark, grayish brown with blackish gray mottling. 
Color of flanks and transverse stripe on chest grayer and duller than back, 
dull gray-brown (hair-brown, pl,XLVI, Ridgway, 1912). 

Anterior side of legs dull gray-brown-rust colored. Ears with well 
developed black tone, especially on the inner side along the margin. 

In contrast to the variable summer color, the winter color is constant, 
pure white. The following variations occur, however: l) the color of the 
underfur may range between snow-white (e, g, , in most Siberian blue hares, 
and in those from northeastern Europe), and delicate straw-white (e,g. , 
in the center of the European part of the U.S.S.R. ); 2) variation in length 
silkiness and luxuriance of hair. Blue hares from the northern and 
northeastern areas have longer, more luxuriant and silky fur. Specimens 
rarely occur in which the winter fur is not completely white, e.g., in the 
southwestern part of the range of the blue hare. Some gray hair tips 



180 



persist in such specimens, giving the animals a peculiar smoky, grayish 
color*. 




FIGURE 111. Blue hare in winter fur. 

Drawing by A. N. Komarov 

208 The weight of the blue hare varies with the locality: in the Verkhoyansk 
District, it reaches 1.600 kg [sic]; the heaviest blue hares from the Moscow 
Region weighed 4.400 kg; hares from the central Volga Region weighed up 
to 4.800 kg; large Chukchi blue hares weighed up to 5.800 kg. D.I. 
Aspisov (1936) gives the following weights for hares of the Tatar Republic. 
Females: average weight of 158 specimens 3.758 kg (minimum 2.600, 
maximum 5.100 kg); males: average weight of 217 specimens 3.336 kg 
(minimum 2,200, maximum 4.400 kg). There are some doubtful records of 
specimens weighing more than 5.800 kg. 

SYSTEMATIC NOTES. Not enough attention has been given to the 
subspecific classification of the blue hare. The following subspecies are 
known in western Europe: 1) Lepus timidus timidus L. 
(Scandinavian Peninsula); 2) L.t.varronis Miller (European Alps, 
to an altitude of 1300 m and more); 3) L.t.scoticus Hilzheimer 
(high mountains of Scotland, northern England and Ireland); 4) L . t . 
hibernicus Bell. (Ireland). This is considered by many authors (e. g. , 
G. S. Miller, 1912; G. Barrett -Hamilton, 1912) as a separate species. 
There are no important cranial differences between the Irish hare and the 
blue hare. The general appearance ofL. hibernicus resembles that 
of the blue hare. The only differences are as follows: l) fur does not 
become white in winter; 2) measurements larger; 3) ears longer; 4) 
interorbital space flatter; 5) postorbital process less elevated; 6) 
incisors larger and thicker. The Irish hare is possibly a well 
differentiated insular form of the blue hare. 



* A blue hare shot in the Spassk County, RyazanProvince (according to V. N. Shchepot'ev "Nature and 
Hunting" (Priroda i okhota), 1886, p. 83) apparently has such a winter color. The fur of this specimen 
is light ash- blue like that of the blue polar fox (A lop ex lagopus beringensis). 



181 



209 



Nine forms of Le pus timidus are known in the U. S. S. R. at present. 
This number may still be increased. 

The taxonomic relationship between the blue hare and the North American 
hares in Alaska, Canada and Greenland can be determined only after 
examination of large series of Annerican and European hares. American 
zoologists distinguish the following species and subspecies, which become 
white in winter or retain their white color in summer, from northwestern 
Alaska to Newfoundland: l) Lepus arcticus (with subspecies) from 
Baffin Island to Newfoundland, Labrador, Banks Island, Prince Albert 
Peninsula to the tundra zone of Canada, from the Bear Lake, the northern 
coasts of the Great Slave Lake to Keewatin; 2) Lepus othus Merriam 
(tundra in north and northwest Alaska); 3) Lepus poadromus Merriam 
(Alaska, near Bristol Bay; 4) Lepus (Boreolepus) 

groenlandicus Rhoads (northwestern coast of Greenland, to the southern 
coast of Disko Bay). 

Some of these species do not exist in fact; they form part of larger 
specific groups. For example, L. groenlandicus Rhoads belongs to 
L. arcticus Ross. 

As was recently shown by Magnus Degerb^l and F. Wimpffen Braestrup 
("Vidensk. Medd. fra Dansk. naturh. Foren", Bd. 98, 1934, pp. 197-205), 
the incisors ofL. groenlandicus which protrude much anteriorly are 
not a constant character; they are found only in the northernmost 
specimens of this race. 

The arrangement of teeth is normal in hares in southern Greenland 
(L. a. porsildi Nelson). These authors came to the conclusion that the 
Greenland hare should be considered as a subspecies of arcticus, and 
that the creation of a separate subgenus Boreolepus Barrett -Hamilt. 
was not justified. 

After comparison of the skull of Lepus arcticus with that of our 
blue hares, I came to the conclusion that they are very closely related. * 
These hares resemble each other in general and in morphological 
characters. It is highly probable that L . arcticus is only a 
subspecies of L . timidus. 

According to A. Howell (1936), there are seven subspecies of Lepus 
arcticus: l) L.a. monstrabilis Nelson (King Oscar Land); 
2) L.a. arcticus Ross, (northern part of Baffin Island, Cockburn Land, 
Melville Peninsula); 3) L.a.andersoni Nelson (Prince Albert 
Peninsula, northern Mackenzie); 4) L.a.labradorius Miller 
(southern part of Baffin Island, northern part of Labrador); 5) L.a. 
bangs i Rhoads (Newfoundland, northeastern coast of Labrador to Cape 
Harrison); 6) L.a. groenlandicus Rhoads (northern part of 
Greenland); 7) L.a. porsildi Nelson (southern Greenland). If 
L. arcticus is considered as a subspecies of timidus, all the above 
forms should be considered as subspecies of the blue hare. 

The position of Lepus othus Merriam can be determined only 
after examination of good material. In D. G. Elliot's opinion (1901), 
Lepus tschuktschorum Nordquist occurs in northern Alaska. 

According to A. Howell (1936), Lepus othus Merriam is closely 
related to tschuktschorum . Their skulls are very similar both in 
general and in measurements. The only difference between them is the 



* Skulls of Greenland hares also belong to the "blue hare type". 



1254 



182 



color of the summer fur. Lepus paodromus Merriam is smaller, 
and its sum,mer fur has some specific characters. 

If this view is accepted, North America from Alaska to Greenland is 
inhabited by various subspecies of the blue hare (Lepus timidus). 
This can be decided only after examination of series of American and 
European blue hares. 

GEOGRAPHICAL DISTRIBUTION. The northern boundary of the range 
of the blue hare begins in eastern Europe, in Lapland and the Kola 
Peninsula. According to F. D. Pleske (1884) Le pu s timidus occurs 
throughout the Kola Peninsula wherever it finds birch forests and willows. 
According to A.N. Formozov (1929) it is not found on Kil'din Island, 
but is common on the adjacent parts of the Murmansk Coast. In K. F. 
Kessler's opinion (1868) the blue hare was very common in the former 
Olonets Province. At present, it is widely distributed and occasionally 
very common in the Olonets Territory, where it reaches the coast of the 
White Sea and Onega Bay. Farther east, the boundary of the range of the 
blue hare includes the greater part of the Kanin tundra, excluding its 
extreme north and the Timan and Bol'shezemel'skaya tundras. In these 
regions, the blue hare is distributed as far as the Arctic Sea. It does not 
include the Kolguev and Vaigach islands, and passes to the tundra north 
of the Urals. It is not exactly known where the northern boundary passes 
in this region, and especially where it crosses the Pai-Khoi Range to reach 
the Baydaratskaya Gulf. 

The northern boundary of the range of the blue hare passes the Yamal 
Peninsula north of 70°N (near Lake Neite). B. M. Zhitkov (1913) observed 
the blue hare even farther south, along the river Khadyta. The boundary 
then follows the coasts of the Ob and Taz bays. The boundary of the range 
of the blue hare on the Gyda Peninsula coincides with the northern 
boundary of the tundra, i. e. , 7l°N. The species is very rare north of this 
line. According to W. G. Heptner (in litt. , 1928), it occurs on the western 
Taimyr Peninsula, and even on Dikson Island. In summer it is common 
near the Pavlovskii Base (30 miles from Dikson). It is rare between 
Dikson Island and Cape Polyn'ya. It occurs in large numbers near Lake 
Pyasino. Near Dudinka on the Yenisei, where there is shrubbery, it is 
numerous and resident. Farther east, it occurs on the Taimyr Peninsula. 
In A. Middendorf's opinion (l867), the northern boundary of its distribution 
on the Taimyr is 75° N, where a litter was found. It occurs along the lower 
reaches of the rivers Khatanga (according to A. F. Chirkova), Anabara, 
Olenek and Lena, reaching their estuaries. 

I received detailed observations on the distribution of the blue hare on 
the western Taimyr from A. A. Kirpichnikov (in litt., 1936): "The blue 
hare occurs in small nunnbers throughout the year along the southwestern 
Taimyr Coast to the Pyasina river. It was also found sometimes in winter 
on Dikson Island. In the second half of November 1933, a blue hare was 
caught on Rastorguev Island south of 74°N. The blue hare becomes very 
rare near the Pyasina Bay. It is not found on the Minin Cliffs nor on the 
coast of Khariton Laptev. Latitude 74°, which passes along the right 
coast of the Pyasina Bay, is possibly the northern boundary of the range of 
the blue hare on the western side of the Taimyr. " 

E.O. Shereshevskii contributed important observations (in litt., 1936) 
on the distribution of the blue hare in the northern parts of East Siberia. 
Tracks and fecal pellets were found frequently during Shereshevskii's 



183 



1935 expedition along the coast from Cape Chelyuskin to Andrey Island, 
and also on the Samuel, Fram, and Faddey islands. In Pronchishchev 
Bay, a blue hare was caught in a trap (April 1935). Shereshevskii has no 
doubts that the blue hare reaches the sea in the Taimyr in years of 
intensive multiplication and migration, and passes to the littoral islands 
across the ice. The islands mentioned above are 2-12 km from the coast. 

According to Shereshevskii, the blue hare occurs in the valley of the 
Yana (below Ust'-Yansk), in the Khroma Basin, and reaches the sea. It 
also occurs in the estuary of the Kolyma (according to S. A. Buturlin). 

The blue hare does not occur on the New Siberian Islands. According to 
fur traders, it was caught once or twice on the Bol'shoi Lyakhovskii Island. 
212 The boundary thus passes along the eastern tundra of Siberia as far as 
the northern coast of the Chukchi Peninsula. 

We shall now examine the eastern and southern boundaries of the range, 
from eastern Siberia to the west of the European part of the U. S. S. R. The 
blue hare occurs in the Chukchi and Koryak territories as far as the 
Olyutorski Gulf, the Penzhinaand Gizhigabays. It is also found in Kamchatka. 
According to S. Barmantov (1933), it is not numerous in the Kronotski 
Reserve. The occurrence of the blue hare in Kamchatka is also mentioned 
by S. Krasheninnikov in the chapter "Terrestrial Animals" (O. Zveryakh 
zemnykh) of his book "A Description of Kamchatka" (Opisanie zemli 
Kamchatki, second edition. Vol. I, 1786, p. 210). 

The eastern boundary includes the whole Maritime Territory, the entire 
coast of the Sea of Okhotsk, and the Tatar Gulf (to about 45°N). The blue 
hare also inhabits Sakhalin and Hokkaido. 

In the Far East, the southern boundary crosses the region of Lake Khanka 
(but does not reach Vladivostok), and continues westward, excluding 
Manchuria. In A. de Sowerby's monograph (1923) this species is 
reported neither for Manchuria nor for Barga [Hulunbuir District] of the 
He ilungkiang Province* (A.S. Lukashkin, 1933). The southern boundary 
probably runs northwest from Khanka crosses the river Sungari, and 
possibly continues as far as the Amur. From Aigun, the southern boundary 
turns south and the species occurs in the northern and central parts of the 
Great Khingan (G. Radde, 1862; A.S. Lukashkin, in litt. , 1936). Farther 
west it is encountered in the valley of the Onon river together with the Tolai 
hare (Raddle, 1862). 

The blue hare apparently occurs in the Hentey, although it has not been 
recorded from there. Farther west it is found in Urianghai [Tuva 
Autonomous Region] and near Lake Hobsogol in northwestern Mongolia 
(Radde, 1862). The exact course of the southern boundary west of this 
point is not known. The blue hare certainly occurs in the Sayan Mountains 
and on the Altai. It is absent in the Tarbagatai, as the environment is not 
suitable. A.M. Nikol'skii (1887) recorded it from Sergiopol [Ayaguzj. 
According to V.N. Shnitnikov (1925), it occurs in northern Semirech'e 
(Dzhety-Su), where it is sometimes numerous in the Lepsinsk District and 
in the former Kopal County. However, I doubt whether it occurs farther 
west than Abakumovskoe. It occurs in the Ala-Kul and Sasyk-Kul districts. 
West of Sergiopol, the southern boundary turns north towards Karkaralinsk 
and Akmolinsk [Tselinograd], The exact boundaries in this region are not 
known . 



[Now Inner Mongolian Autonomous Region. ] 



184 



From Akmolinsk, the boundary probably continues to Atbasar, passing 
between Kustanai and Turgai. It then turns southwest passing south of 
Aktyubinsk along the northern foothills of the Mugodzhar Mountains, and 
touches the middle reaches of the Ilek river (N.A. Zarudnyi, 1897). It 
then reaches the Ural river, and includes Obshchii Syrt south of Uralsk 
(approximately 51°N). The exact course of the boundary fromi Obshchii 
Syrt to the Volga is not known. It turns sharply north to the Samara Bend, 
excluding the Trans -Volga Steppe. In the lower Trans -Volga, M.N. 
Bogdanov (1871) observed this hare in the northern part of the former 
Kamyshin County, where it was very rare. It was not recorded from this 
district by V.E. Martino (1913). We assume that the southern boundary of 
the range of the blue hare in the Volga Region passes between Volsk and 
Saratov. It then turns north to Atkarsk and Serdobsk, continues to the 
former Penza Province and passes between Morshansk and Tambov, The 
blue hare is sporadically distributed in the Tellerman Wood on the Khoper, 
near the town Borisoglebsk (A. Silant'ev 1894), farther south in the Grafskoe 
Forestry, north of Voronezh and in the Shipov Forest in the Pavlovsk 
District. 

From the Morshansk and Tambov region, the southern boundary runs 
towards Ranenburg [Chaplygin], where the blue hare is very rare. The 
boundary then passes north of Tula, turns sharply south to Odoevo, 
Bolkhov and passes near Khotynets (Riga-Orel Railroad). Farther south, 
the boundary includes the basin of the Desna with its tributaries Navlya 
and Nerusa (former Trubchevsk County, Orel Province). The main 
southern boundary then turns west towards Novgorod-Severskii (52°N) and 
continues to south of Novozybkov (Dm. Vilinskii, 1885). Farther west, the 
blue hare occurs in the forests of the Mozyr Polesye (A. G. Fedyushin, 1927) 
and farther south, in the forests of the Ovruch District (M. Baklashov, 
1877). The boundary then turns sharply north, includes the area of the 
Pinsk Polesye and continues in Poland towards the area north of Bialystok 
and Novogrudok (E. R. Lubicz-Niezabetowski, 1934). 

In addition to this "main" southern boundary in the southern Ukraine, 
there are scattered, small, isolated areas where the species has perhaps sur- 
vived until the present. Forinstance, a blue hare was found near Sumami. In 
the collection of the Moscow Zoological Museum is a pelt and skull 
obtained by A. V. Apollonov (1913) from the northern part of the former 
Sumy County of the Khar'kov Province. According to N.I. Gavrilenko 
(1928), blue hares rarely occur in the former Zenkov, Poltava and 
Konstantinograd [Krasnograd] counties of the Poltava Province. In his opinion, 
they are the offspring of specimens introduced by the former landowner 
Kharitonenko in the former Bogodukhov County of the Khar'kov Province. 

Professor K. F. Kessler (1850) recorded blue hares from the Volhynian 
and Kiev Provinces. In 1844 a blue hare was obtained in the Radomysl 
County. In 1848 such a hare was observed in the vicinity of Kiev, where 
it was still very rare. According to E. V. Sharleman (1915), it occurs very 
rarely in the former Kiev Province. It was observed in a drive near 
Korsun', former Kanev County of the Kiev Province. 

M. Shcherbina (1924) reported that he observed a blue hare on 
4 February 1917, near Ivintsy, former Zhitomir District. V. Khranevich 
(1925) states that it is very rare in Podolia. One was killed near the 
village Cherepovaya of the Proskurov County. Another was killed near 
Zin'kovtsy of the former Kamenets Podolski County. 



185 



Except for these fragmentary records, we have no data on rare and 

' , , accidental captures in the southern and central Ukraine. We repeat that 
214 

these isolated records lie outside the constant range. 

The population of the blue hare is subject to great fluctuations due to 
ecological conditions which vary from year to year. 

In conclusion, the discovery of fossil remains of the blue hare in the 
Crimea may be mentioned (S.K. Dal', 1928). During the construction of 
the Ayanskii Basin on the Petrovskie Rocks near Simferopol, numerous 
remains (mainly nnandibles) of blue hares were found in Karst caverns 
which were partly filled with sand. In S.K. Dai's opinion, the presence 
of the blue hare in the Crimea was short lived. It apparently entered the Crimea 
during the Wiirm glacial, when the ice reached 48-50°N, and the land 
bridge connecting the Crimea with the Continent was 200 km wide. At 
the end of this epoch, the blue hare disappeared and the common hare 
settled in the Crimea. 

Although some details of S.K. Dai's theories are questionable, there 
is no doubt that the blue hare was formerly distributed farther south, and 
that it was driven back north as a result of cultivation. * This results in 
restriction of the range of the blue hare, contrary to the phenomenon 
observed in the common hare. As shown above, the range of the common 
hare gradually expands, especially in northern, eastern and southeastern 
directions. 

MODE OF LIFE. Habitats. In the central belt of the European part of the 
U.S.S.R. , the blue hare is very frequently found in dense undergrowth with 
willow and alder shrubs, and in undergrowth consisting of birch and 
spruce. It occurs among fallen trees in clearings covered with branches 
and logs, or in broken terrain in which forest alternates with ditches and 
forested gullies. It is also attracted to spruce forests mixed with birch 
and aspen (opened by felling), where the ground is mossy, especially 
where fallen trees are numerous. 

The blue hare occurs in definite habitats in southern European forests, 
on the southern border of its range. For example, in the Shipov Forest, 
former Pavlovsk County of the Voronezh Province, and in the Tellerman 
Forest on the Khoper, the blue hare prefers aspen groves. According to 
P. A. Polozhentsev (1935), it occurs in all types of plantations in the 
Buzuliik Forest. However, it prefers shrub thickets and pine nurseries. 
It occasionally causes them great damage by gnawing the bark and 
„ (- eating buds. 

In the north of the European U. S. S. R. , e.g., the Kola Peninsula, the 
range of the blue hare is connected with birch forests and osiers. It also 
enters the tundra. 

In western Siberia, the blue hare is widely distributed in the forests, 
mainly in floodlands with willows. For instance, L. P. Naumov observed 
it in such habitats in the Surgut District. It occurs in the same habitats m 
the Yenisei forests, e. g. , along the Tunguska. 

"The blue hares are most numerous in valleys of large rivers with 
dense willow thickets, or on the shores of lakes and at the edge of small 
forests in the tundra. It is often found on the banks of small streams 
overgrown with forests of larch, spruce and cedars with dense alder and 



* See M.N. Bogdanov's map of distribution in the article "Game and Trade Birds and Animals of the 
European Part of Russia" (Okhotnich'i i promyslovye zveri i ptitsy Evropeiskoi Rossii, "Okhota i 
konnozavodstvo", No. 3, 1873, p. 63 (Map 3)). 

186 



216 



217 



willow underbrush. It is rare on mountains tops and in the bare Taiga 
almost devoid of underbrush. It is very fond of dense undergrowth on 
mountains, where it makes many tracks. It was not seen on bare, stone- 
covered high mesas. " 

According to G. G. Doppelmeyer (1926), the blue hare is common 
during the summer near Lake Baikal, in willow thickets of estuaries of 
rivers, in birch groves, and in pine forests. According to D.K. 
Solov'ev (1920), it is found on the Sayan Mountains up to the "belogor'ye"*. 
According to A. M. Kolosov (1933) it is found on the Altai in the high- 
mountain tundra (e.g., in the Teletskoe Lake Region) in dense thickets of 
polar birch (Be tu la r o t un di fo li a ) and willows (S ali x herbacea, 
S. reticulata). According to G. N. Gassovskii (1927), in eastern 
Siberia (in the Gilyui-Ol'doi Upland) blue hares occur in large numbers 
in regions overgrown with P inu s pumila, on the crests of ranges 
between the Great and Lesser Yurmakit Mountains, along the Gilyui and 
Getkan, and especially among willow and black-crowberry shrubs. In 
Transbaikalia, blue hares occur not only in the forest zone, but also in the 
Daurian Steppe, e.g., along the Onon river. 

In the West Siberian forest-steppe, at the southern limit of their range, 
blue hares often occur in reed thickets along the shores of lakes, and in 
willow thickets. They are also found in birch and oak groves and in 
vegetation along rivers. I found numerous blue hares in these habitats 
in the valley of the Belaya river in Bashkiria. 

In the forest steppe and steppe, the blue hare typically lives among so- 
called "black alder pollards" overgrown with nettles, Umbelliferae, shrubs 
of dewberry, wild rose and raspberry. Such habitats are also typical of 
the Bashkirian blue hare. In north Siberia it lives in the stunted forest 
tundra. It is also widely distributed in the open tundra. 

S.P. Naumov writes: "The blue hare concentrates in river valleys 
overgrown with trees and shrubs, in the forest tundra, but its tracks are 
often seen on watersheds. . . this changes completely when we enter 
the tundra, especially the part in which the last high shrubs (Alnus 
fruticosa) disappear. The absence of large trees and shrubs compels 
the hare to seek other shelters. It finds them in the high, precipitous, eroded 
cliffs common in some parts of the tundra. Here the sporadic distribution of 
the blue hare is even more striking. One may pass tens of versts without 
finding a single animal or its tracks. However, in the habitats mentioned 
above the blue hare is very numerous; one may see sometimes ten or more 
without leaving a place. Only frightened hares run far into the tundra. 
However, they do not go beyond the limits of the broken, hilly area. " 

GENERAL DESCRIPTION AND HABITS. The movements of the blue 
hare are rapid. It is an agile animal. According to a popular concept 
the hare is very timid. The blue hare was considered especially so, 
and was therefore named "timidus" by Karl Linne. After observing its 
behavior in nature it is hardly justifiable to consider timidity as one of its 
main characteristics. 

V.N. Kaverznev (1928) writes: "It is not justifiable to consider timidity 
peculiar to the hare. The fact that it takes flight when it senses peril does 
not prove that it is timid. Had it failed to do so, it would have been 



[Flat alpine massifs. ] 



187 




FIGURE 112. Blue hare in summer fur 

Photograph by S. I. Ognev 

exterminated. It is badly equipped for resistance, but has enemies 
at every turn. All animals try to escape danger. Even the most 
redoubtable predators usually avoid meeting man. They fight only 
exceptionally, prompted by hunger or despair. 

"It has been stated that a hare trembles when it is caught, which is true 
especially of the nose, lips and whiskers. If this nervous phenomenon 
really exists, it is not peculiar to the hare, and should not be ascribed to 
fear alone. 

"Confusion is one of the main symptoms of fear. The hare does not 
show panic when hunted by greyhounds, and in a hopeless situation. Its 
dodges, hooks, and jumps in face of the dogs are proof of purpose which 
would not be present in a state of panic. " 

Its sense of hearing is best developed. Its sight is poor, but its sense 
of smell is well developed. The blue hare is a taciturn animal. According 
to P. A. Manteifel' (1927) males and females make sounds resembling a 
moan during estrus. 

A hunter of the Military Hunting Company, M. Potapov, informed me 
that the spring cry of the male resembles the tawny owl's cry "uik-uik", 
but is shriller. It is intermediate between the cry of the tawny owl and that 
of a blue hare when it is caught by hounds. The female makes a peculiar 
smacking sound produced by drawing air through pursed lips. According 
to Potapov, males are attracted by this sound. Hunters from Shar'ya told 
A.N. Formozov that the sound of the female described by Potapov is not the 
female's voice, but the voice of young hares. They claimed that male and 



218 



female run to help the young in response to this sound. There is even a 
method of hunting using this sound to attract the hares. 

Other hunters stated that the cry of the male blue hare sounds like that 
of the forest owl (A s io otus), like "o-kho-kho-kho". N.R. Obrecht 
maintains that he saw male blue hares raising themselves on their hind 
paws, emitting this cry and afterwards running with lowered head, sniffing 
the track of a female v^hich had passed. Other hunters gave the same 
description of the spring cry of the hare. 

Even on this simple question, opinion is divided and the problem should 
be further investigated. 

A wounded hare captured by hounds or men usually emits a shrill 
squeal like that of screaming piglet or child. 

Footprints of the hare in the snow clearly reflect its various gaits. 
When it is foraging, it moves slowly, leaving in the snow parallel 
elongate footprints of its hind feet followed by consecutive rounded footprints 
of its forefeet. When it runs, the footprints are more elongate. The 
traces of the hind feet are parallel, but one of them slightly in front 
of the other. The traces of the forefeet are further behind those of the 
hind feet. Sets of traces are separated by a large space, the length of 
which depends on the speed of the hare. A hare in a great hurry leaves 
tracks with even wider intervals and greater spacing of the footprints. 

Before going to sleep, the blue hare "covers" its track by jumping to 
the side, away from its main direction. It may also go along paths or ruts 
in the snow, where its track is usually not perceptible. Its sideways jumps 
are called "hooks". Many naturalists and hunters think that the hooks of 
the hare are proof of its cunning. The hare thus covers its tracks and leads 
its persecutors astray. The hare often passes from recent snow to paths 
or ruts and makes hooks for a considerable distance in order not to leave 
a plain track. This may be instinctive cunning . Hooks may be regarded 
as a behavior pattern acquired during evolution and are probably a reflex 
activity not involving any conscious cunning. 
219 V.N. Kaverznev (1928) writes: "I would consider the inventiveness of 
the hare in covering its track to the lair as cunning, and not as inherited 
from preceding generations. The method follows principles, according to 
which the animal performs stereotypic double and triple hooks. 1 would 
also consider as cunning the rare cases in which the hare lies in a 
depression, in an empty fox burrow, or between boughs of a fallen tree, 
if it were possible to prove that these places had not been chosen 
accidentally. These activities reflect an instinct to conceal the lair, and 
not the track leading to it. In other words, they are manifestations of the 
animal's efforts to hide. 

"For instance, I do not consider double and triple steps='= a method of 
confusing the track. I consider the double step an everyday occurrence. 
After covering some distance in search of a lair or food, the hare returns 
to make sure it has not missed a place or an object, or in order to rest. 
In some cases, I observed a hare making a double step, nibbling a plant, 
and finally covering its track. 

"After finding a suitable lair, the animal makes a hook towards it. If 
the lair is unsuitable, it continues its course. This is the reason for the 
absence of hooks from triple steps. The hare's habit of performing hooks 



* Double and triple steps mean a double or triple repetition of tracks by returning along the line of the track. 



189 



before resting is probably the result of its old habit of jumping aside and 
lying down when chased. It probably generalized this habit to everyday 
cases of lying down to rest. " 



/■ M .^ -. '^ ^ 







nCURE 113. 



Tracks of slow walk of a foraging blue hare 

Photograph by S. I. Ognev 



220 It often stops by the way, and changes the direction of its course. All 

this confuses its track. Using an anthropomorphic argument, all these 
phenomena can be explained from the point of view of human logic, by 
ascribing cunning to the hare. However, such anthropomorphic 
interpretations are probably not valid. 

The hare is a typical land animal and its rapid progression is therefore 
on the ground. It does not run rapidly only on even ground. It can move 
safely and quickly on mounds and steep slopes. Difficulties appear when the 
hare runs quickly down a steep mountain. As it has long hind legs, it 
sometimes turns end over end in a somersault, as if shot. The hare 
climbs precipitous mountains rapidly and safely owing to the build of its legs. 



190 



Even when the animal could escape by running over an even space, 
choose to climb a mountain, rapidly ascending the steep slope. 



it may 




221 



FIGURE 1 14. Track of a huirying blue hare 

Photograph by S. I. Ognev 

The blue hare is a strong and good swimmer. During floods, herds of 
hares may collect on islands. If necessary, they swim across the water 
to escape floods. When chased by hounds, the hare often jumps into the 
water and easily crosses large rivers. It holds its ears backwards 
during swimming. 

Floods in spring are often fatal for the hares. The tired, soaked 
animals have to escape from one island to another. Finally they die of 
exhaustion. During this difficult period, the hares are also hunted by 
hawks and eagles and by poachers. 

The hares may escape to ice floes, small spits and islands, and gather 
on stumps and heaps of brushwood. The hunters do not shoot them, but 
kill them with clubs. They may kill 20-30 specimens daily, including many 
gravid females. 



191 



DAILY LIFE. The blue hare is a crepuscular animal, its activity 
beginning in the evening, when it comes out to feed. During the day and in 
the late morning the hare usually lies in a shelter or moves slowly about 
a small area in sheltered, safe places. The activity of the blue hare ceases 
at night and begins again early in the morning. 




FIGURE 115. Track of slowly moving blue hare 

Photograph by S. I. Ognev 

V.G. Stakhrovskii (1932) observed blue hares in a large open-air cage. 
He describes their daily life as follows. The hares emerged from the 
lairs at sunset (which was very early in winter and much later in spring 
and summer). They rose one after another, and 15-20 minutes after the 
first hare rose, all were feeding. In the morning they fed and basked in the 
sun for a long time. In summer they were often seen walking at 6.00a.m. 
On several occasions, Stakhrovskii saw hares in the early morning hours 
playing with each other, rolling in the dust of a sown oat field. The animals 
rolled on their backs and from side to side, their long hind legs dangling 
ridiculously in the air. 



19: 




FIGURE 1 16. Blue hare in winter 



Photograph by S. I. Ognev 



222 



V. G. Stakhrovskii could not find out whether the hares are active at 
night. Some observations indicate that they are not. This view was 
confirmed by S. V. Lobachev, who watched at night in sheds on oat fields, 
and noticed hares only in the evening or in the morning (not between the 
hours 11.00 p.m. and 3.00 a. m. ). During quiet, moonlit, winter nights, 
they feed and leave footprints even after midnight. 

NESTS AND BURROWS. In the European part of the U. S. S. R, the 
blue hares do not build lairs or burrows to raise their young. The female 
gives birth in a concealed place under shrubs or fallen trees. According 
to P. A. Manteifel, the young are active on their first day. In autumn, 
especially when there is no snow, the hare, which has become white, 
conceals itself very carefully. It hides under overhanging roots, under 
a fallen tree, shrubs, etc. In winter, when there is deep snow, the hare 
lies down under a snow-covered bush after making a hook. It sometimes 
breaks the snow and builds a well-concealed shelter. When it is discovered 
it jumps energetically, spraying snow in all directions. After a moment, 
the hare becomes visible on the white background of the landscape. 

In eastern Siberia, blue hares occasionally make large burrows in the 
snow. L.O. Belopol'skii (inlitt., 1936) writes that in winter, along the 
Anadyr river, the hare usually remains in the region of shrubs and dwarf 
forest. It rarely enters the tundra or the mountains. Among shrubs, 
Belopol'skii occasionally found burrows in the snow 80-100 cm deep, for 
instance, near the estuary of the Anadyr river (23 November 1930). 

Blue hares in northeastern Siberia occasionally Duild permanent 
communal burrows on hills and steep river banks. Such a colony was 
described by A. F. Girikova (in litt. , 1935). 

"l had the opportunity of observing hares during our journey to the lower 
reaches of the Khatanga (from village Khatanginskoe up to the Bol'shaya 
Korcha locality). We saw very few hares in the forest belt. They appeared 



193 



223 



only in the tundra which begins south of the Krestovskii Peninsula, The 
first striking fact was the "symbiosis" between two predaceous birds, 
buzzards and merlins, and hares. We observed the same phenomenon in 
four places along the river. The presence of these birds proved to be a 
reliable indication of the presence of hare burrows. Small groups of hares, 
possibly including young, could always be found near nests of the birds, as 
if they were under their supervision. The nests of the birds were always 
on top of the escarpments. Below them were the colonies of hares. I 
carefully observed such a colony near the village Novoe Rybnoe. This group 
of hares lived near a merlin nest, and apparently consisted of one litter 
(five hares killed proved to be young of the same year='')- The hares 
occupied seven conical hills along the Khatanga river (for about 1/2 km). 
The cliffs here reached a height of 20-30 m. The surface of the escarpment 
was sandy with occasional clay outcrops from the underlying bedrock. 
Here and there it was covered with fine shingle. The sandy parts of the 
conical mounds were covered with dwarf tundra vegetation consisting of 
weeds, tundra birch and osier. When people appeared, the hares began 
running anxiously in circles about the conical hills, always returning to 
certain points. Near these points we found their burrows. 

"These burrows were clearly inhabited. Around them we found numerous 
tracks, pellets and remains of food. I saw hares emerging from the 
burrows. At first we thought that these were abandoned burrows of polar 
foxes, which had been temporarily occupied by the hares. Closer examination 
showed that they had been dug by the hares themselves. They were 
usually located at half the height of the conical hill and had one or two 
exits. The burrow went first to a depth of 1-2 m into the ground. This 
section was followed by a turn. We could not excavate the burrows, because 
they led into the depth of the hills. One inhabited burrow was on a small 
hill in a ravine descending to the river. It had one exit with three small 
air vents located 2m above the exit and on either side. We concluded 
that the burrow branched in the ground on a small platform above the 
exit, which was 16 cm high and 39 cm wide. 

"The air vents of another burrow were located along a crack in the ground. 
The largest burrow (25X35 cm) was on the greatest conical hill. It had one 
exit on either side of the hill and apparently passed through it. The 
distance between the two exits was 7 m. There was a track on the surface 
of the hill connecting the two exits. Both the exits and the track were 
located on a shingle -covered slope, so that even a small animal could 
not approach the burrows without making a noise. Several burrows with 
one exit were empty. We also found some recent, unfinished burrows. 

"On top of one mound we found a whole network of collapsed corridors 
and small round pits which had been inhabited by young hares. Their 
small pellets were still there. The burrows of the young hares were very 
close to the merlin nest (about 10m). " 

A. F. Chirkova's description of these colonial burrows of the Khatanga 
blue hares is interesting in many respects. No other subspecies is known 
to have such habits. The Khatanga hares are a well-differentiated 
subspecies. 

NUTRITION. The blue hare lives on a diet of grain, grasses, berries 
(e.g. , bilberry and raspberry) and twigs. The hare avidly feeds on aspen 



It seems doubtful that one litter could occupy such a widely spread system of burrows. 



194 



224 



and willow twigs. V. G. Stakhrovskii (1932) reports that during the 
Verkhnevychegod expedition he repeatedly saw blue hares in pine forests, 
bilberry and pine thickets with spruce, especially near marshes. It seems 
that the blue hares fed mainly on bilberry, especially its roots. V. G. 
Stakhrovskii (1932) studied the food of the blue hare in the open-air cages 
of the Losinoostrovskaya [Babushkin] Biological Station. The animals fed 
on the following plants which grew in the cages: Poa trivialis, P. 
amnia, Carex pilosa and raspberry. They also ate willow, birch 
and aspen shoots. In September the hares began to feed on the bark of 
willows, aspens, nut trees, young oaks, and even bark and branches of 
spruce. They visited especially sown fields of oats, sunflowers and 
clover mixed with millet. Their favorite food was unripe oats. They did 
not touch potatoes, parsley or carrots. Experiments showed that the blue 
hares accepted branches and bark of the following trees: aspen, poplar, 
willow, birch, nut-tree and also oak. They completely ignored honeysuckle, 
spindletree, elder, buckthorn, linden and European bird cherry. 

Detailed observations on the food of the blue hare are given by D.I. 
Aspisov (1936). In summer, blue hares in the Tatar Republic are 
especially fond of M e li c a nutans, C a 1 am agr o s t i s , Greek valerian, 
cornflowers, and Picris hieracioides. They paid little attention to 
Aegopodium and ground ivy . 

In dry mixed forests hares generally live on birch, aspen, nut trees and 
willow in winter. In floodlands they live on osier, willow, oak, and aspen. 
In oakwoods they live on maple, oak, nut trees and rarely on birch and 
aspen. 

When there is little snow, they feed on the following plants: sprouts 
of winter rye, Brassica campestris L,. , Agropyrum, 
Calamagrostis, angelica. Chicory, Chenopodium, Artemisia, 
Arctium and nettle. The first two are especially favored by the hares, who 
oftenhave to digthe plants out of the snow. They usually bite off the parts 
protruding from the snow. 

REPRODUCTION. In the central part of the European U. S. S. R. , the 
estrus usually begins in the middle of February and lasts until the end of 
July. The females have 2-4 litters duringthe summer . D.I. Aspisov (1936) 
thinks that blue hares normally have 3 litters each summer . In western 
Siberia, for instance in the Tara District, estrus begins in March 
(I. Shukhov). In northeastern Siberia it begins much later, in the Kolyma 
Territory only in late April (S. A. Buturlin). The genital organs of all 
hares caught in early May were clearly swollen. According to Aspisov, 
the first litter usually consists of 3 young, the second of 5 and the third of 
3. Most femiales have two litters a year, producing 8 young. The natural 
death rate among the young reaches 35-40 per cent. The progeny of a 
female in late summer and early autumn is thus reduced to about 5 young. 
The young generation is about four times as numerous as the adult 
population. 
225 In the north of the European U.S.S.R. the first litters sometimes appear 
as early as in March. The hunters call the young of these early litters 
"March hares" or "hares of the frozen-snow period". The last litters appear 
in September or later, and are called "autumn hares" by the hunters. Inthe 
hunting literature we find many cases of late pregnancy in blue hares. In the 
the "Hunter's Gazette" (Okhotnich'ya gazeta. No. 17, 1889) we read that a 
gravid female was caught in late December in the former Petersburg 



195 



226 



Province. According to Aspisov (1936) litters normally appear in Tataria 
in early May, the last days of June and the first days of August. According 
to Aspisov, earlier or later litters should be considered as accidental, 
which do not give a true picture of the normal biology of the hare. 

P. A. Manteifel' ("Hares of the U.S.S.R." (Zaitsy SSSR) in "Biology of 
Hares and Squirrels" (Biologiya zaitsev i belok), Moscow 1935) reported 
the following observations on the reproduction of the blue hare in the 
Moscow Zoological Garden. 

He writes: "The males started chasing a female blue hare on 5 April 
1927, and another on 7 April 1927. The estrus was rapid and violent. Four 
males chased one female. The female ran away from them and made 
sudden sideways jumps. The males drove each other away from the female. 
They collided while running, leaped and struck each other with their hind 
legs so that fur flew. When they halted, they bit each other and quarreled, 
scratching each other with their forelegs. They huddled behind the female 
which suddenly stopped. The most agile male leaped on the female. The 
female raised herself high on her feet and curved her back (Figure 76). 
The other males began scratching and biting the copulating animals, jumped 
over them and kicked the male with their hind legs. 

"Only a very persistent male could have fertilized the female under these 
conditions. After copulation the female lay down in a depression, breathing 
heavily. The males did not touch her, but occasionally they attacked each 
other. The males sometimes mounted each other. When they had lost 
sight of the female, they tried to find her, running about and sniffing the 
ground. During this they often sniffed each other quickly and then 
continued the search. Their sense of smell is apparently not sharp enough, 
because they often passed a female lying on the ground, without noticing 
her. 

"The estrus of the female, during which she copulates frequently, does 
not last more than 24 hours, generally only 10 hours. Copulation is brief 
as in rabbits. The penis looks like a long hook with a sharply curved end. 
This forces the male to push his hindquarters under the legs of the female 
while he holds her with his forelegs. It also compels the female to rise 
high on her legs and to curve her back. 

"The estrus of the male is equally active during day and night. 
Observations in the Moscow Zoological Garden showed that the female had 
intercourse about every half hour. It was mounted most of the time by the 
same male. " 

According to P. A. Manteifel' gestation lasts 48-51 days. The young 
are generally born on the 50th day. They are born with open eyes and are 
covered with curly wool. Their ears stand up as in adults, and the upper 
and lower incisors are visible outside the gums. The stomach of one -day- 
old hares contains curds. Their weight at this age is 84-140 grams. The 
young suck about 40 grams of milk on their first day. The milk of the female 
is very fat. Milk in the stomach of a newborn blue hare contained 23.6% 
fat. Immediately after birth the newborn young suck their fill, scatter 
and hide in the grass. Their development is influenced by the following 
peculiar maternal instinct. When the puerperal estrus (which lasts about 
10 hours) is over, and she comes across the track of a hiding young she 
immediately looks for it, runs to it and suckles it. When the female 
approaches, the young hare runs towards her. The young hare usually 
begins to move towards its mother when she is 2-3 m away. P. A. Manteifel' 



196 



noticed that females suckle not only their own young but also other animals 
who leave tracks. Only after 2-4 days do the young hares leave the lair. 
The duration of the rest period depends on the quantity of milk obtained. 
When young hares are taken by men, they cry loudly. The female does not 
actively protect her offspring. However, she runs past and hits the ground 
with her hind legs, as do rabbits. On the eighth or ninth day the young hares 
already begin feeding on grass. Later on, the female suckles her young 
not more than two or three times. 

The female eats the placenta imnnediately after birth. P. A. Manteifel' 
writes: "By eating the placenta the mother sets the young free. This act 
is useful for the mother and for her offspring. Flesh is an unusual food 
for the hare, and causes diarrhea. This is necessary because in the last 
days of pregnancy the large fetuses compress the intestine which becomes 
blocked with feces. If the placenta had not been eaten, it would have 
decomposed and the odor would have attracted predators, which might have 
found the young and killed them. " 

The female can be fertilized again immediately after giving birth. The 
short northern summer is exploited so that not a single hour is wasted for 
reproduction. 

P. A. Manteifel' describes birth and postnatal events as follows: "We 
observed a female in the Zoological Garden which gave birth after 50 V 2 
days of pregnancy. We saw a wet young hare, energetically crawling out from 
227 under her belly. The newborn raised its head trying to get hold of the 

teat. It sought it by smell and with its lips. At the same time the female 
turned over and with her teeth pulled another young one from her vagina. 
The newborn was in a thick layer of membranes. The female tore the 
membranes and swallowed them in large pieces. After freeing the young one 
she licked it. The newborn disappeared under her belly and made 
smacking sounds with its lips. Two more were born, and disappeared 
under the mother's belly. A large male was sitting near the female. This 
male tried to copulate with the female already during birth, and once she 
even assumed her copulating posture. When the female raised herself on 
her legs and curved her back the three young did not cease sucking and 
remained almost suspended in the air. Copulation did not take place 
because the fourth young covered with bloody membranes was still protruding 
from the vagina. The male jumped off and sat nearby while the female 
extracted the bloody mass with her teeth. Holding it with her forelegs, she 
tore the membranes and swallowed them. Her mouth still bloody with the 
membranes, she rose on her legs and curved her back. The male jumped 
onto her and copulated. There was only little bleeding during birth. The 
whole process lasted about five minutes, including the copulation attempt. " 

D.I. Aspisov (1936) provides interesting data on the growth of the young. 
This is particularly rapid in later litters, so that they become strong 
enough to resist the beginning of the cold. This is clearly illustrated by 
a comparison of weight increase of the young of different litters. From 11 
July to 31 August the weight of a leveret of the second litter increased by 
80.3 % while that of a specimen of the first litter increased by only 52.0% 
during the same period. From 11 July to 30 October the weight of a leveret 
of the second litter increased by 88.3 % as compared to 65.8 % in a specimen of 
the first litter. 



197 



228 



MOLT. All authors who wrote on the molt of the blue hare agree about 
the spring molt. However, data about the autumn molt are contradictory. 

Loewis* suggested that in autumn the hair of the blue hare grows and 
becomes depigmented instead of having a true molt. I came to the same 
conclusion after examination of large series of central Russian blue hares 
(S.I. Ognev. "Mammals of the Moscow Province" (Mlekopitayushchie 
Moskovskoi gubernii), 1913, pp. 306-307). I. V. Zharkov recently came 
to the same conclusion ("Fur Structure and Autumn Molt of the Blue Hare" 
(stroeniyi mekha i osennyaya lin'ka zaitsa belyaka), Raboty Volzhsko- 
Kamskoi biologicheskoi stantsii, Vol.1, 1931, pp. 153-167). He thinks that 
the autumn molt consists of the following stages: 1) appearance of new 
hair bulbs and renewed activity of old bulbs; 2) growth; 3) depigmentation. 

P. A. Manteifel' (1935) made experiments in the Zoological Garden to 
determine whether the fur of the hare becomes depigmented in autumn or 
whether the fur is replaced by new hair during the molt. He wrote (1935): 
"We observed that the hares became white, and asked ourselves whether 
the gray summer fur is replaced by new white fur, or whether the summer 
fur persists wholly or in part, losing its dark pigment. In order to 
determine this the following method was used. In July, two blue hares in 
summer fur were dyed with henna and hydrogen peroxide. As a result the 
fur became rust colored. In the middle of November the hares became 
completely white. We then examined the dyed animals. Much dyed hair 
persisted under the very long white fur which completely concealed the 
short summer fur. It thus appears that most of the summer fur (mainly 
the undercoat) is retained towards winter and its length remains almost 
unchanged. " 

This seems a strange conclusion, which in our opinion does not follow 
from the results of the experiment. Some of the summer fur (undercoat) 
was certainly retained by these hares when they were examined. However, 
after the end of the molt it would probably have been completely replaced 
by winter fur. 

In the recent edition of Brehm's Tierleben revised by L. Heck** we find 
that the fur of hares in the Berlin Zoological Garden was dyed with aniline 
dyes. These experiments showed clearly that blue hares have a normal 
autumn molt without depigmentation. J. Allen (1894) and Professor 
E. Lbnnberg (who wrote to L. Heck) came to the same conclusion. 

Molt in the blue hare was recently studied by B. A. Kuznetsov and E. A. 
Pavlova. 

They write: "It was thought until recently that the sharp color and 
structural changes of the fur in autumn were caused by its depigmentation, 
by its growth and by the appearance of a new undercoat. These changes 
were not thought to be due to replacement of summer fur by winter fur. 

"Our work consisted of the following steps: 1) observation of light 
specimens in the Moscow Zoological Garden; 2) morphological and 
histological studies of the pelts of animals from various regions of the 
U. S. S. R. ; 3) dyeing of summer fur of light specimens. We proved that 
the former notions on autumn changes in structure and color of the fur 
were erroneous. Our conclusion that the hair is completely replaced in 



O. von Loewis, Bemerkungen iiber den Schneehasen Livlands. Zoologisch. Garten, 1877, S. 16-20. 
Brehm's Tierleben, Saugetiere, Bd. II, 1914, S. 68. 



229 









ij 


/K^-*^^ 


«s 




N 


<cY 


<M 


r^ 











^^"s^-Y 




-V' -1 




199 



230 




200 



autumn was based on the following: 

1. Some parts of the fur were dyed with permanent colors in midsummer. 
During the autumn molt the colors became gradually paler and had 
disappeared completely at the end of November. 

2. The length of dark hair was measured every five days between 1 
October and 15 November. It was found that the length of the hair is 
constant and that the hair does not grow. 

3. The hair of summer fur of animals caught in July- August always had 
a keratinized, bulbous root with an occluded cordiform canal. This proves 
that growth in autumn could not have taken place. 

4. We never found partly depigmented hair in autumn fur. The latter 
always included summer hair which had retained its dark color. 

5. The number of white hairs per surface unit growing in autumn usually 
corresponded exactly to the number of hairs on the same area in the 
completed winter fur. 

6. When pelts of animals shot in autumn are combed, the dark colored 
hair falls out easily. 

7. The structure of summer and winter fur is different in the 
blue hare. We never found summer hair in the winter fur. 

230 "All this proves that the fur of the blue hare is completely replaced 
during the autumn molt, as in other animals, " 

Molt takes place in a definite sequence. The white winter fur appears 
in the following sequence (Figure 117): on the tail, the hind legs, posterior 
part of back (behind the sacrum), the upper parts of the ears, forefeet, 
chest, neck, flanks, and head, behind the eyes. The following parts molt 
later: back, forehead, sides of the head (especially tip of the nose), 
cheeks and the area below the eyes. 

The spring molt begins slowly, but then becomes very rapid (i.e., the 
fur falls out in whole tufts). It proceeds from the head towards the 
hindquarters. The summer fur appears first on the anterior part of the head, 
then on the ears, upper part of back, and flanks. It appears latest on the 
lower part of the back and the feet. Molting takes place diffusely and 

231 uniformly. According to D.I. Aspisov (1936), the spring molt (including 
the very first slow stage) lasts about three months. According to the 
locality, the spring molt begins in March-April, and even in May. In the 
extreme north of Siberia, e.g., in the lower reaches of the Yenisei, Taz 
and along the Kolyma, it begins in the middle of summer. There occur blue 
hares in mid- June which have not yet completed their spring molt and do 
not have complete summer fur. 

I examined a series of Kolyma blue hares obtained by S. A. Buturlin. In 
No. 7 (female, 7 June 1905), I found tufts of white fur on the cheeks, 
occiput and upper part of the neck. Tail, feet and throat (except for a gray 
transverse stripe on the throat) were white. In No. 4 (male, 4 June 1905), 
caught near the village Pokhodskoe, remnants of the winter fur were even 
more numerous, covering most of the head and forming large white patches 
on the back, flanks and legs. The fur of these specimens appeared as if 
at the height of the spring molt. Together with such specimens occur 
hares (No. 2, female, 4 June 1905) which have an almost complete dark 
summer fur. The molt occurs last on the feet. In specimen No, 2 
(female) the feet are completely white. 

In hares of the extreme north, some parts of the body have hardly 
enough time to complete their molt into summer fur. For example, in blue 



201 



hares obtained by L.A. Portenko near Markovo (Anadyr Territory) on 
5-7 July 1932, the hind legs and tail are still completely white. 

Autumn molt in the Moscow Region begins in the middle of September 
and ends in December. The time of completion of the molt varies 
considerably in different specimens, depending on age (old specimens molt 
earlier) and on the individual. The hares begin molting normally even in 
snowless late autumn and early winter. They are then in a very 
disagreeable situation. Their white fur is very conspicuous on the dark 
grass, when the frightened animal moves among the leafless shrubs. 

Farther north the autumn molt begins much earlier. In specimen No. 3 
(old, 3 September 1935) of the series of Kolyma blue hares collected by 
S. A. Buturlin, there is white hair on the ears and on the lower part of the 
back. In No. 8 (3 September 1905) the ears are almost completely white 
and there is much white hair on the flanks, the anterior part of the head and 
the hind part of the back. A hare caught along the Kureika river by F. F. 
Shillinger on 25 July 1925 (S.I. Ognev's collection No. M. 3176), had clearly 
begun its autumn molt. White hair had appeared on top of the head, ears, 
flanks and in some places on the back; feet completely white. Many 
Khatanga specimens (collected by A. F. Chirkova in August) had begun 
molting. Summer fur is thus worn by blue hares of the extreme north only 
for a short period, little more than two months. 

POPULATION DYNAMICS AND MIGRATIONS. The blue hare population 
of a locality is not constant but is subject to considerable fluctuations. 
232 Both in Russian and foreign literature there are many reports on the large 
scale of these fluctuations, and the factors regulating them are discussed. 

These data have been summarized recently by A.N. Formozov. 
("Population Dynamics of Commercial Animals" (Kolebaniya chislennosti 
promyslovykh zhivotnikh), Moscow, 1935, pp. 12-32). After analyzing data 
from the literature and discussing observations by correspondents, A.N. 
Formozov comes to the following conclusions. The increase and decrease 
in populations of hares recur periodically and regularly. The rhythm of 
fluctuations remains the same for large regions with similar physico- 
geographical conditions. In the south and southwest of Russia the fluctuation 
cycles are short, recurring every five years. These periods become 
longer farther north. In the Leningrad Region the cycle lasts 5-6 years; 
in Karelia 9; in Yakutia 9 years or even more. " 

American naturalists also observed a periodicity in the fluctuation 
in numbers of Le pus americanus. Ch. Elton (1924) compared the 
cycle of the American hare (every 10 years) with the sunspot cycle. In 
his opinion, the peaks of populations coincide with years with a minimum 
of sunspots. This, however, seems inexact because there are exceptions 
to this rule. Elton's conclusion therefore seems hardly justified. 

To illustrate the scale of fluctuations of the populations we give a few 
examples of mass reproduction. Some regions become overpopulated and 
the hares begin migrating. M. Sulima (1911) describes such a migration 
in a "fertility year": "In the autumn of 1908 a large migration of hares 
from the east to the west was observed in the former Vologda and 
Arkhangelsk provinces. The hares proceeded in masses, feeding on their 
way on haystacks, aspen thickets, etc. In the Vychegda District they 
destroyed so much hay that they caused considerable damage. Hares were 
caught by all methods along the Pechora river in that year. The hunters 
did not have time to lay traps. In M. Sulima's village 150 sled dogs lived 



202 



233 



only on hares during the whole season. In the winter of 1909-1910, the 
number of hares fell off markedly. None of the hares shot during this 
season were young. The young apparently had died, and multiplication 
decreased. 

A.N. Formozov's correspondent wrote that hares (apparently mainly 
blue hares) were unusually numerous in the Kamyshlov District in December 
1921. Aspens, birches and willows 1.5 cm thick were eaten completely. 
No guns were used. Hares were caught with nets. Groups of 25 people 
covered distances of 200 m with nets and caught as many as 80 hares each 
time. 

Mass multiplication and migrations have been described also from 
Siberia (M. Konstantinov, "Fur Trapping and Fur Commerce in the 
Yakutsk Territory" (Pushnoi promysel i pushnaya torgovlya v Yakutskom 
krae), 1921; N.N. Beretti, "In the Extreme Northeast" (Na krainem 
severo-vostoke), 1929, etc. ). 

After studying recent statistics A.N. Formozov came to the conclusion 
that the number of pelts treated in "productive years" may be 50 or more 
times as large as the number of pelts obtained in "poor years" in the 
_„. Ivanovo Region, 25 times as large in Karelia, 20 times as large in the Western 
and Leningrad regions, 35 times as large in the Moscow Region, etc. The 
amplitude of the fluctuations is thus very large. 

There are many causes for the increase of numbers. In A.N. 
Formozov's opinion, one of the main causes is the natural purification and 
disinfection of land in the 2-3 years after epizootics, during which the 
population of hares is very small. After populations of hares have been 
very high, there are large numbers of oocysts of coccidia and eggs of 
parasitic helminths in the soil. This results in epidemic outbreaks which 
cause mortality among the hares. However, the parasites apparently die in 
2-3 years. Hares which survive the epizootic are apparently immune to 
infection. The survival rate of the young is certainly increased in the 
area in which the pathogens have disappeared. 

Another important factor is the increased fertility of females at the 
beginning of a "productive" year, MacFarlan observed for Lepus 
americanus that in productive years the number of young per litter 
increased from 3-4 to 6-8 (or even 10), One of the prerequisites for mass 
multiplication is favorable food and climatic conditions, e.g., sufficient 
food in winter, a low snow cover, etc, A deep snow layer would have 
covered the young shrubs growing after a year with a high population of 
hares. 

Two related factors cause increased population: abundance of food and 
increased fertility of females. 

As stated above, peaks of blue hare populations have a definite 
periodicity, the intervals of which increase towards the north, because as 
a result of the short summer the blue hares in the north have fewer litters 
in summer than do the hares in the south, where the summer is longer 
and hotter and the autumn warm and prolonged. 

Among the main factors which reduce populations are epizootics. The 
denser the population of a species, the quicker epizootics spread. 

A.N. Formozov (1935) writes: "The scale and course of an epizootic 
depend on the following factors: l) the population of hares; 2) climatic 
conditions; 3) presence of disease carriers and vectors of parasites; 
4) landscape features (especially characteristics of the soil). The disease 



203 



235 



236 



appears first in separate, small foci. These expand rapidly, encompassing 
progressively larger areas until they become confluent, leaving only small 
healthy islands in which small numbers of hares survive after the animals 
around them have been killed by the epizootic. Such "islands" have been 
repeatedly observed by hunters during recent large epizootics. They were 
generally located on isolated, high and dry plots in which the population of 
hares was moderately dense. " A.N. Formozov collected material on 
the great "plague" of hares which occurred between 1926 and 1930. This 
epizootic broke out with great violence in the south, mainly among common 
hares. As the plague moved northward, it also began to affect populations 
of blue hares. 

In 1928-1929 reports were received on mortality of hares in the Ukraine, 
the central Chernozem Region, Belorussia, the Urals, the former Saratov 
and Stalingrad territories, Moscow, Ivanovo and Western regions. Isolated 
reports came also from the Northern and Nizhni Novgorod (Gorki) 
territories. In 1929-1930 the plague had ceased in the south and on the 
Urals, where, after reaching its minimum, the population of hares again 
began to increase. The epizootic occurred in the northern area and in 
the Leningrad Region in 1929-1930, continuing in these areas during 1930- 
1931. The decrease of hares was irregular during these years. It was 
especially marked during the second year of the epizootic. 

The victims of the epizootics are very numerous. A.N. Formozov (1935) 
reports that in the spring after the 1929 plague in the Shar'ya District of 
the Nizhni Novgorod Territory, when rubbish was being burned in felling 
areas, a number of blue hares, still covered in winter fur, were found in 
each plot. 

Mortality of hares is also increased by the following factors: l) deep 
snow; 2) snow storms; 3) strong frosts; 4) ice-covered soil; 5) rainy 
summers. 

One of the important parasites of hares during plague years is the liver 
fluke (Fasciola hepatica). The hares also contract strong infections 
of Taenia serrata. Tens of thousands of larvae of this parasite 
(oncospheres) are deposited with feces on the ground and among the grass. 
They are then ingested with the food. 

A dangerous lung parasite of the hares is the helminth S ynt h e t o c au lu s 
commutatus. According to M. P. Liubimov (1935), epizootics of 
Synthetocaulus are very severe in rainy years in low and humid 
pasturelands. Strong infections with this parasite cause symptoms of 
pneumonia with frequent coughing and sneezing. The presence of 50-100 
worms in the lungs of a hare produces a pathological picture of pneumonia. 

Trichostrongylosis caused by Trichostrongylus retortaeformis 
and T. instabilis is very common in hares . Infected hares are 
emaciated, anemic, and have chronic intestinal catarrh. 

The liver of the hares is often parasitized byLinguatula taenioides 
(Pentastomida). 

According to M. P. Lyubimov (1935), about half the hares suffer from 
coccidiosis. Eimeria stiedae infects the liver and E i m e r i a 
perforans and E . magna, the intestine . 

More information about diseases of hares is given in M.P. Lyubimov's 
"Diseases of Squirrels and Hares" (Bolezni belok i zaitsev), in "Biology 
of Hares and Squirrels" (Biologiya zaitsevibelok), pp. 80-140, Moscow, 1935. 

The blue hares cause damage to forest nurseries, undergrowth and 
shrubs. They damage trees by gnawing rings around the trunk. Young 

204 



trees suffer severely during years of mass multiplication of hares. The 
blue hares often damage gardens and melon fields. 

The blue hare is hunted by foxes, lynxes and wolves. It is also attacked 
by smaller predators such as ermines and weasels which kill mainly young 
hares. According to I.I. Kolyushev (1936), the polar fox usually does not 
attack the blue hare. It does not attack a hare which is lying down nor does 
it drive it from its lair. The following predacious birds hunt hares: 
eagles (e.g., golden eagles), goshawks and great horned owls. 

The greatest enemy of the hares is man, who hunts them for their meat 
and fur. 

COMMERCIAL VALUE. Pelts of hares are of great commercial value. 
They are used for fur articles and are also shorn and dyed for the 
manufacture of felt. Pelts of hares are an important article of Russian export. 

Before the revolution about 6 million pelts of hares of all species were 
processed in Russia. 

Modern stocks of pelts of blue hares are subject to great fluctuations 
depending on "productive" years of the hares. According to B. A. 
Kuznetsov (1932), the stocks of pelts may range between 100 and 502% 
during six years. The number of pelts of blue hares constitutes 1.2- 
6.1% of the total fur production of the U.S.S.R. 



232. 




FIGURE 119. Blue hare in beginning of spring molt 



Photograph by S. I. Ognev 



205 



Subspecies of the blue hare 

KEY TO SUBSPECIES OF THE BLUE HARE (LEPUS TIMIDUS L.) IN THE U.S.S. R. 

1 (8). Anterior inner margin of ears black or black-brown. 

2 (5). Skull massive; zygomatic bones wide; length of skull 98-116mm(M. 103mm); zygomatic 
width 51-57 mm (M. 52.8 mm). 

3 (4). Summer fur on back dull grey-brown-straw-colored with a little blackish mottling. Lower part 
of back grayer than middle, ash-gray. Anterior irmer margin of ears with marked blackish tones; its 
color clearly different from that of back. 

L. timidus tschuktschorum Nordquist. 

4 (3). Color of summer fur on back pale gray. Lower part of back gray with an ash-blue tinge. 
Anterior inner margin of ears pale gray with a slight rust tinge. 

L. timidus begitschevi Koljuschev. 

5 (2). Skull less massive and zygomatic bones narrower; length of skull about 100mm, usually less; 
zygomatic width usually less than 50 mm. 

6 (7). Anterior inner margin of ears black-brown with slight straw-rust mottling. Measurements large: 
length of skull 95.2-101. 7mm (M, 97mm); zygomatic width48-50. 8 mm (M. 49mm). Weight about 4 kg or more. 

L. t. timidus L. 

7 (6). Anterior inner margin of ears usually black. Length of skull 88.7-99. 2 mm (M. 96.6 mm); 
zygomatic width 45-50mm (M. 49.3mm). Weight usually less than in L. t. timidus, about 2. 7-3. 1 kg. 

L. t. kolymensis Ognev. 

8 (1). Anterior inner margin of ears not black or black-brown, but of lighter color. 

9 (10), General color of back rich rust-brown. Anterior inner margin of ears rust-brown. 

L. t. kozhevnikovi Ognev. 

10 (9). General color of back not rich rust-brown. Anterior inner margin of ears not rust-brown. 

11 (12). Color of back of light and dull color: straw- earth- gray or dull gray. 

Color of summer fur on back dull, light straw- earth- gray without any rust tone or light straw mottling. 
Color of lower part of back not markedly different from that of other parts of the back. 

Anterior inner margin of ears dull straw-earth-gray, their color not different from color of back. 

L. t. sibiricorum Johansen. 

12 (11). Color of back brownish gray. 

13 (14). Color of lower part of back in front of tail deep gray, clearly different from color of other 
parts of the back. Color of back dark chestnut-brown-gray with a marked black tone. Anterior inner 
margin of ears brownish gray, sometimes with blackish and slight straw tone. 

L. t. gichiganus J. Allen. 

14 (13). Color of lower part of back in front of tail light gray with a brown tone, not sharply different 
from that of other parts of the back. 

Color of back lighter than inL. t. gichiganus, between rust-brown with straw and gray mottling to 
duller, more brownish tones with a slight straw-gray and blackish gray mottling. Color of anterior inner 
margin of ears not sharply different from that of back. 

15(16). Measurements small: length of skull 86-93.2 mm; zygomatic width 45-49 mm. Length of 
body 440-515 mm; hind feet 132- 152 mm. 

L. t. transbaikalicus Ognev. 

16 (15). Measurements large: length of skull 88-96 mm; zygomatic width 47-51 mm. Length of 
body 523-570 mm; hind feet 155- 180 mm. 

L. t. mordeni Goodwin. 



206 



340. Lepus timidus timidus Linnaeus (1758). Northern blue hare 

1758. Lepus timidus Linnaeus C. , Systema Naturae, editio 10, p. 57. 

1777. Lepus timidus alpinus Erxleben J. Ch. , Systema regni animalis per classes, ordines, genera 
etc. , Mammalia, I. , p. 328. 

1778. Lepus variabilis Pallas P.S,, Novae species quadrupedum e Glirium ordine, pp. 1-17. 
1778. Lepus algidus Pallas P. S. , op. cit. , p. 2 (New Name for the Alpine hare, Thomas Pennant, 

British Zoology, IV, 1770, p. 40). 
1778. Lepus borealis Pallas P. S. , op. cit., p. 2. 
238 1795. Lepus septentrionalis Link H. Fr, , Beitrage zur Naturgeschichte, Rostock, B. II, p. 73. 

1816. Lepus albus Leach W. E. , Systematic Catalogue of the Specimens of the Indigenous Mammalia 

and Birds that are Preserved in the British Museum, p. 7 (Nomen nudum.'). 

1819. Lepus albus Leach W. E. , in Sir John Ross's Voyage of Discovery, editio I, app. II, pp. XLV-XLVI; 
editio II, Vol.11, app. IV, pp. 17, 170 (Nomen preoccupatum.'). Bechstein J. M. , Gemeinniitzige 
Naturgesch. Deutschlands etc. , 1801, editio I, Vol.11, S. 1096 (Lepus timidus albus from 
Thiiringia). 

1820. Lepus borealis Nilsson S. , Skandinavisk Fauna, Lund, B.I, p. 211. 

1829-1832. Lepus borealis c ol li nus Nilsson S. , Illuminerade Figurer till Skandinav. Fauna, Lund, 

I, pi, 16, 19. 
1829-1832. Lepus borealis sylvaticus Nilsson S. , op. cit., I, pi. 22. 
1844. Lepus canescens Nilsson S. , Ofversigt af Kongl. Vetensk. Academ. Fdrhandl. , Stockholm, I, 

p. 133. 
1884. Lepus variabilis Pleske Th. , Ubersicht der Saugethiere und Vogel der Kola-Halb. , 1884, S. 

108-111. 
1906. Lepus medius scoticus Hilzheimer M. , Zoologischer Anzeiger, XXX, S. 511. 

TYPE LOCALITY AND DEPOSITION. In the tenth edition of "Systema Natiurae" we find the following: 
"Habitat Europa. Hyeme in frigidis niveus exceptis auribus nigris". The type locality is not given. G. S. 
Miller (1912) proposes to consider the vicinity of Upsala as the type locality of the nominate race. Holotype 
not preserved. 

MATERIAL EXAMINED: 12 pelts and 13 skulls. 

DIAGNOSIS. In summer fur anterior part of muzzle, cheeks and 
forehead dull brownish gray-rust-colored. A considerable admixture of 
blackish hair on the upper part of forehead, vertex and between the ears. 
These parts of the head therefore appear duller and darker. 

Color of back dark, dull brownish gray with a delicate pattern of straw - 
gray and blackish gray mottling. Anterior inner margin of ears black- 
brown with slight straw-rust mottling. A dull grayish brown (hair-brown) 
stripe across the chest. Winter fur completely white, underfur white, hair 
not very luxuriant, soft or silky. 

Length of skull 95.2-101.7 mm (M.97mm); zygomatic width 48-50.8 mm 
{M.49mm). 

Length of body 590-600 mm; hind feet 155-164mm (M. 158.6 mm); 
ears (measured along concavity) 84-94 mm (M. 89 mm). 

SYSTEMATIC NOTES. Typical northern blue hares are characterized 
as follows: l) rather dark brownish gray summer fur; color of 
back mixed with hair-brown and sepia; flanks with grayer tones (hair-brown 
and mouse -gray) 2) black-brown color of anterior inner margin of ears; 
straw-rust-colored mottling on the background of this dark color. 

In spite of variations of the general color the dull gray color of the 
summer fur may be considered as typical for L. t. timidus. 

Winter fur with snow-white undercoat. 

Not only specimens from the north of the European part of the U.S.S.R. 
belong to the nominate race of the blue hare, but also some from the 
„„_ northern part of the Tobolsk Territory, e.g., from near Surgut and Narym. 
Specimens which are transitional to the western Siberian lighter race, 
L.t. sibiricorum Johans . occur in Narym . 

207 



Owing to the courtesy of the late Professor G. E. Johansen I had the 
opportunity of examining the holotype of L . lugubris Kastsch. = obtained 
on 20 June 1928 near Ongudai on the Altai. A second specimen (cotype) 
was caught along Um'gumen' river on the Altai. 

A comparison of Le pu s lugubris with typical blue hares showed 
that they are closely related. It is almost impossible to distinguish 
lugubris from the nominate race of the blue hare. 

Comparison of series showed the following differences between Le pus 
t.timidus and L . lugubris Kastsch: l) general color of fur of the 
Altai hare with a stronger straw tone especially on the head; 2) dark color 
of ears (i.e. , black color on anterior inner margin of ears) weaker than 
in the north European race. 

The Altai blue hare is only a natio of the nominate subspecies and it 
should be named L. t. timidus natio lugubris . 

G.E. Barrett-Hamilton (Proceedings Zoolog. Society of London, 
parti, p. 90, 1900) described the subspecies L . t . alt ai c us from a single 
specimen (pelt and skull) in the collection of the British Museum. It is 
labelled "Altai Mountains". Barrett -Hamilton does not give a description, 
but only records the basal length of skull and blackness of the ears. If 
Lepus t. altaicus really came from the Altai Mountains, the blackness 
of the ears places it with L. t. lugubris, and makes it a synonym of 
this. 

The boundaries of the range of the northern blue hare in western Siberia 
are not known. Blue hares from Narym are transitional between L . t . 
sibiricorum and L. t.timidus in some characters . The Narym 
hares have practically no black anterior inner margins of the ears. 
Specimens from Tomsk, Novosibirsk and Omsk are typical L . t . 
sibiricorum. L. t. sibiricorum thus seems to separate the Altai 
range of L . t . lu gub r i s from that of the nominate subspecies. The 
ranges ofL. t. timidus and L . t . lu gub r i s possibly overlap in a 
narrow strip beyond the Yenisei Mountains and farther north. 

Melanic specimens are very rare among the blue hares. One such 
specimen obtained on 2 October 1936 near Arkhangelsk was sent to me by 
V. Ya. Parovshchikov. The color of the guard hair on the back of this 
specimen is a brilliant black. Undercoat and roots of hair gray. Color of 
underparts, especially of belly, lighter, whitish gray. 

GEOGRAPHICAL DISTRIBUTION. The typical race occurs in 
Scandinavia, Finland, Lapland, Murmansk Coast, Karelia, Onega 
Territory, the Valdai Hills, in the Vologda, Kostroma, Vyatka [Kirov] 
and Perm territories. It is still difficult to establish the southern boundary 
of the range of the race in this region. It apparently passes near 57-58° N. 
In western Siberia the nominate race occurs near Berezovo, Tobolsk, 
Surgut, and Narym. A slightly different form (natio lugubris Kastsch. ) 
occurs on the Altai. 

341. Lepus timidus kozhevnikovi Ogn. (1929). Central Russian 
blue hare 

1895. Lepus ti m i dus Satunin K. A. , Pozvonochnye Moskovskoi gubernii. Mlekopitayushchie (Vertebrates 
of the Moscow Province. Mammals), Izvestiya Obshchestva Lyubitelei Estestvoznaniya, antropologii i 
etnografii, Vol. XXXVl, p. 14. 



* N. F. Kashchenko, Results of the Altai Expedition, Tomsk, 1899, pp. 57-59, tab. II, fig. 4. 

208 



240 1913. Lepus timidus Ognev S. I, , Mlekopitayushchie Moskovskoi gubernii (Mammals of the Moscow 
Province), Vol.1, pp. 298-307. 
1929. Lepus timidus kozhevnikovi Ognev S. I. , Zur Systematik der russischen Hasen. Zoologischer 
Anzeiger, Bd. 84, H. 1/4, S. 79-80. 

TYPE LOCALITY AND DEPOSITION. No. 2368 male, 16 October 1911, near village Chemaya, 
Bogorodsk County, Moscow Province, collection of the Moscow Zoological Museum. 
MATERIAL EXAMINED: 89 pelts and 125 skulls. 

DIAGNOSIS. In summer fur, anterior part of muzzle, cheeks and forehead 
rich rust-brown. Upper part of forehead, vertex and area between ears with 
some blackish hair. The color of these parts is therefore generally more 
intense and darker. 

General color of back rust -brown with very light grayish black mottling. 

Anterior inner margin of ears intense rust-brown, occasionally with a 
black tinge caused by black-brown hair and black-brown hair tips of the 
other guard hair. 

A broad dull straw-gray stripe (between hair-brown and drab) across 
the chest, which has a marked straw-rust-colored tone. 

Winter fur white. Undercoat with light yellowish roots. Yellowish 
tone of the hairs of undercoat with a slight rust tinge. Hair of winter fur 
coarse and less dense and silky than in northern and northeastern forms. 
Length of hair on back 35 mm; hair on lower part of chest and sides of belly 
up to 60 mm. 

Lengthof skull (Figures 120, 121) 97.8-105.6 mm (M. 100mm); zygomatic 
width 50-53 mm (M. 52.3 mm). 

Length of body 450-630 mm; hind feet 155- 180 mm (M. 170.8 mm); ears 
(measured along concavity) 80-100 mm; ears (measured from occiput) 
110-ll8mm (M. 113.8mm). 

SYSTEMATIC NOTES. The Central Russian blue hare is well 
differentiated from the northern subspecies; the color of its summer fur is 
richer. According to N. A. Severtzov (1855), the summer fur of Moscow 
blue hares is more reddish. The hair of these southern specimens is rust- 
colored with black rings, which results in a yellow-brown ground color. 

I had not enough blue hares from the southern parts of the range in 
eastern Europe. I mentioned already that towards the south the color of 
the fur becomes lighter and richer. Moscow blue hares are more reddish 
and of a richer color than those from Murmansk, Arkhangelsk, etc. 

Large series should be examined to determine how far south the tendency 
of southern specimens to have lighter and richer color continues. It seems 
that even if the Voronezh and other southern blue hares are lighter and 
richer colored than the Moscow hares, this feature is still transgressive, 
and not sufficiently well marked. It is still premature to speak of a south 
European race of the blue hare. 

Among series ofL. t. kozhevnikovi one finds together with typical 
specimens, markedly darker specimens from the same locality.. For 
example, on 22 August 1933 in a humid forest in the floodland of the river 
Belaya (near the village Angasyak in the Dyurtyuli District of Bashkiria, 
former Birsk County of the Ufa Province), N.E. Schultz shot a male, the 
color of which was rather dark with the anterior inner margin of the ears 
black. On the same day I shot in the same forest a female with the normal 
rich color characteristic of the subspecies kozhevnikovi . 

A series of typically colored hares obtained in Bashkiria by S. V. Kirikov 
included a female (shot on 20 May 1931 in tugais along the river Belaya) which 
had the anterior inner margin of the ears markedly black. 



209 



242 



241 






FIGURES 120-125. Skull of subspecies of Le pus timidus. 120 — Lepus t. kozhevnikovi Ogn. , 
No. 2343, 23 Oct. 1911 Bogorodsk County Moscow Province, Collection of the Moscow Zoological Museum; 
121 —Lepus t. timidus natio 1 u gubris Kastsch. , type No. 3177, 20 June 1898, Altai, Ongudai, 
N. F. Kashchenko. Photograph by S. I. Ognev; 122 —Lepus t. sibiricorum Johans. , type No. 4858, 
Chulym River; 123 —Lepus t. tiansbaikalicus Ogn., type No. M. 3178; 124— Lepus t. 
begitschevi Koljushev, No. 18786, KZMAN; 125 —Lepus t. tschuktschorum Nordquist, 
No. M. 4346, S. I. Ognev's collection, from the Chukot Peninsula. Photograph by S. I. Ognev. 



210 



GEOGRAPHICAL DISTRIBUTION. The southern and southwestern 
boundaries of the range of this race are the same as the southern boundary 
of the range of the blue hare in eastern Europe. 

The northern boundary of the range is not quite clear. However it 
coincides with the southern boundary of the range of the typical race (see 
above). Richly colored blue hares occur in the Baltic Region, near 
Yamburg, Peterhof and Novaya Luga. That is, hares colored like L . t . 
kozhevnikovi occur in the western parts of the Leningrad Region and 
apparently near Leningrad. All specimens from the region examined 
belong to this subspecies. Hares from the Valdai Hills in the former 
Novgorod Province belong toL.t.timidus. 

In the northwestern part of the U. S. S. R. , the range of L . t . 
kozhevnikovi reaches farther north than in the central parts of the 
European part of the U . S . S . R . 

It is difficult to state where this subspecies disappears in the east. 
Turgai specimens are much more closely related in color to L . t . 
sibiricorum* than to L. t. kozhevnikovi. Hare s from the southe r n 
Urals and Bashkiria belong to the central Russian form. 

342. Lepus timidus sibiricorum Johans. (1923). West Siberian 
blue hare 

1877. Lepus variabilis Peters W. , Ubereicht iiber die wahrend der Sibirisch. Expedition von Dr. 

Finsch gesamm. Saugethiere, Monatsberichte der Konigl. Akadem. der Wissensch. z. Berlin, S. 735 

(Samarovo on the Irtysh). 
243 1886. Lepus variabilis Stepanov P. , Zoogeograficheskii ocherk lesisto-bolotistoi polosy mezhdu rek 

Om'yu, Taroi i Irtyshom (A Zoogeographical Survey of the Forest-Marsh between the rivers Om', Tara 

and Irtysh), Zapiski Zapadno-Sibirskogo otdeleniya Geograficheskogo obschestva, 8 (l):2S-38. 
1923. Lepus timidus natio sibiricorum Johansen G. E. , Po Chulymu (Along the Chulym), 

Izvestiya Tomskogo gosudarstvermogo universiteta, p. p. 58-59. 
1923. Lepus timidus Martino V. and E. , Materialy po sistematicheskomu i geograficheskomu 

rasprostraneniyu mlekopitayuschikh Kirgizskoi stepi (Material on the Systematic and Geographical 

Distribution of Mammals in the Kirghizian Steppe), Ezhegodnik Zoologicheskogo muzeya Akademii Naok, 

Vol. XXIV, p. 29 (Semipalatinsk Region). 
1925. Lepus timidus Zverev M. D. , Obzor gryzunov Kuznetskoi stepi (A Survey of the Rodents of the 

Kuznetsk Steppe), p. 133. 
1928. Lepus timidus Shukhov I. , Promyslovye zveri Tarskogo okruga (Fur Animals of the Tara District), 

"Okhotnik" No. 7, p. 22. 

TYPE LOCALITY AND DEPOSITION. Western Siberia, valley of Chulym. Type No. 4858 female, 7 
January 1914, river Chulym, Artarek Island, near village Novokuskovo, G. E. Johansen, collection of the 
Zoological Museum of the Tomsk University. 
MATERIAL EXAMINED: 17 pelts and skulls. 

DIAGNOSIS. In summer fur, color of anterior part of muzzle, cheeks 
and forehead pale and dull straw-rust-gray; general color of these parts 
much duller and more yellow than in Moscow blue hares, between 
avellaneous and cinnamon. 

Color of upper part of forehead and vertex duller and darker because 
of a slight admixture of blackish tips of some hairs. 

Color of back dull and light straw-earth- gray without any rust tinge. 



* After comparing series of L. t. sibiricorum with hares from Kustanai, I found that their color is 
richer than that of typical sibiricorum. However they do not have the rust tones of L. t. 
kozhevnikovi. Some specimens are more closely related to L . t . timidus. 



21 



244 



Color of anterior inner margin of ears straw-earth-gray, not 
different from general color of back. 

A broad dull straw-gray stripe across the chest. Hair tips in this region 
whitish straw. 

Winter fur white; undercoat completely white. 

Length of skull (Figures 122,128) 92. 3-100. 2mm (M. 98.2mm); zygomatic 
width 46.7-53.2 mm (M. 49.7 mm). 

Length of body 433-550 mm; hind feet 145-155 mm (M. 152. 1 mm); ear 
(measured along concavity) 86-92 mm; ear (measured from occiput) 104- 
117mm (M. 110.3mm). 

Systematic notes. This subspecies is clearly distinguished from 
the central Russian blue hare by the following characters: l) light earth- 
straw-gray color of back without any admixture of reddish rust color; 
2) color of head much lighter straw-rust-gray; 3) color of ears pale 
straw -earth- gray. 

The West Siberian blue hare differs even more sharply from the 
nominate race than from L. t. kozhevnikovi. 

GEOGRAPfflCAL DISTRIBUTION. The range of this subspecies is not 
well known. It occurs mainly in the plains of western Siberia. It is 
widely distributed in the Tomsk Territory. The northern boundary of its 
range passes between the Chulym and Tara rivers. In the south it occurs 
as far as Abakan, Semipalatinsk and farther south to the Samara District 
(south of the Kalba Range). This record is very interesting. I have 
a series of hares identical with true sibiricorum, which were collected 
in the Samara District by V.I. Dasenkov. Farther south this subspecies 
occurs in the Zaisan Depression (specimens in the collection of the 
Zoological Museum) and also in northern Dzhety-Su (the Lepsinsk, Kopal, 
Alma-Ata districts). After examination of specimens given by V.N. 
Shnitnikov to the ZMAN I decided that the Dzhety-Su specimens belonged 
to L.t. sibiricorum. Its distribution towards the west is still unknown. 
It occurs in plains from the Zaisan Region towards Akmolinsk [Tselinograd], 
Atbasar and Kokchetav. 

According to N. M. Mikhel' (1934), 1^ e p u s timidus sibiricorum 
occurs in the Borovoye Forestry south of Kokchetav. In his opinion blue 
hares from the Troitsk District do not differ from sibiricorum . 
Specimens from the Kustanai District (south of Kustanai, the Naurzum 
Reserve) are intermediate between L. t. timidus and L.t. 
sibiricorum, and differ clearly from L. t. kozhevnikovi 

The Kustanai-Turgai belt is apparently inhabited by a transitional 
type, and the ranges of three races meet in this region. As the habitats 
of the Turgai blue hare are rather specialized (see below) we may assume 
that this region is inhabited by a separate natio. The boundary of the range 
of L.t. sibiricorum passes west of the line Atbasar-Petropavlovsk. 

343. Lepus timidus t r an sbaikalicus Ognev (1929). 
Transbaikal blue hare 

1910. Lepus timidus Kashtchenko N. F. , O kollektsii mlekopitayushchikh iz Zabaikal'ya (A Collection 
of Manvmals from Transbaikalia), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, XV, p. 271. 

1926. Lepus t i m i dus Soronov 1. , Fauna gryzunov v raione Ivanovo-Arakhleiskikh khutorov (Rodent 
Fauna ih the Region of the Ivanovo-Arakhleiskie Hamlets), in "Regional Studies in Transbaikalia" 
(K kraevedcheskim raboiam v Zabaikal'e) Izdanie Chitinskogo gosudarstvennogo muzeya, p. 41. 



212 



245 



1926. Lepus timidus Svatosh Z. F. , Pushnoi i okhotnichii promysel (Hunting of Fur Animals and Fur 

Tradey, in G. G. Doppelmeyer "The Sable Trade Along the Baikal Coast" (Sobolinyi promysel na 

poberezh'e Baikala), p. 176. 
1929. Lepus timidus Kuznetsov B. A. , Zverovye promysly vostochnogo Zabaikal'ya (Animal Trade in 

Eastern Transbaikal), Trudy po lesnomu opytnomu delu, Tsentral'naya lesnaya opytnaya stantsiya, 

issue VI, p. 92. 
1929. Lepus timidus transbaik al icus Ognev S, 1. , Zur Systematik der russischen Hasen, 

Zoologischer Anzeiger, Bd. 84. H. 1/4, S. 81. 

TYPE LOCALITY AND DEPOSITION. No. M. 3178, 23 December 1924, Sosnovka, Barguzin Taiga, 
near Lake Baikal, in Transbaikal. S.I. Ognev's collection. 

MATERIAL EXAMINED. 12 pelts and skulls. 

DIAGNOSIS. In summer fur, anterior part of muzzle, cheeks and 
forehead dull brownish gray-rust, colored sometimes with an admixture 
of richer rust color. 

A greater admixture of blackish hair on the vertex and between the 
ears. These parts are duller than the anterior part of forehead and cheeks. 

Color of back dark, dull brown with a slight straw-gray pattern and 
blackish gray mottling. Color of anterior inner margin of ears almost 
the same as that of back, dull brown with straw-gray and blackish mottling. 

A dull grayish brown (hair -brown) stripe across the chest. Color of 
chin and throat of all specimens examined dull gray with a slight brown 
(light drab) tinge. 

Winter fur white, undercoat with a slight yellowish straw tone (at the 
roots of the hairs). Hair of winter fur more luxuriant, silky and soft. 
Hair of middle of back 42 mm long; hair of flanks and belly up to 6 2 mm 
long. 

Length of skull (Figures 123,129) 86-93.2 mm (M. 90.7 mm); zygomatic 
width 45-49 mm (M. 46.7 mm). 

Length of body 440-515 mm; hind feet l32-152mmL; ear (measured 
along concavity 77-92 mm. 

SYSTEMATIC NOTES. The above diagnosis shows that the color of 
summer fur ofL. transbaikalicus differs little from that of the typical 
form. The general color of the Transbaikal blue hare is lighter. Its 
ears do not show an admixture of blackish tones. Measurements of the 
Transbaikal blue hare smaller than those of the typical form. 

Examination of a small series of Transbaikal blue hares showed that 
their color is variable. 

No. M.4506 female, 22 August 1928 from the Chita District (collected 
by B. A, Kuznetsov) closely resembles specimens from Novgorod in the 
color of the back. However, the latter have a markedly black anterior 
inner margin of the ears. The color of the ears of the Transbaikal 
subspecies is identical with that of the back, grayish brown with a slight 
straw mottling. 

The color of No. M.4507 male, 22 August 1928 (from the same locality) 
shows a stronger brownish rust tinge. The color of this hare is 
practically the same as that of a specimen from the Kanin Peninsula (Semzha 
river, 18 June 1913, No. S. 2739, collection of the Moscow Zoological 
Museum). However, the blue hare from Kanin, like all northern blue 
hares, differs from the Transbaikal subspecies in the dark blackish gray 
color of the anterior inner margin of the ears. 

The color of young L.t. transbaikalicus is typical. They differ 
clearly from young L . t . ko zhe vniko vi from the Moscow Region. Their 
color is much grayer than that of the latter. The general color of the back 



213 



246 




.- d 



o ;5 



»■ ? 



« .- 
^ * 



6 I 

'i^ 00 ^ 
CM 00 



d 2 



1254 



214 



247 



245 of young Transbaikal hares is rather dull with a straw-brown tinge and 
slight straw- white mottling (color close to Jiair-brown and buff -brown). 

This mottling is caused by a pale straw-gray subapical band of the hair. 
Anterior part of head, i.e., nose, lips and eyes more rust-colored. Color 
of anterior inner margin of ears like that of back. Flanks grayer than 
back. Transverse stripe on chest gray with a slight brownish tinge. 

GEOGRAPHICAL DISTRIBUTION. Throughout Transbaikalia, in the 
taiga, forest and forest steppe, and steppe. The population of blue hares 
is subject to yearly fluctuations. Hares from the Great Khingan 
certainly belong to the Transbaikal subspecies. A blue hare sent by A.S. 
Lukashkin is in my collection. The specimen was obtained from the upper 
reaches of the Eimanatchi river along the Bol'shaya Yadora river, in the 
Haimin Kungssii Forest Concession inthe upper reaches of the river Imin Gol. 
This specimen is very small, especially its skull. The Khingan hares are 
perhaps even smaller than the Transbaikal hares. 

344. Lepus timidus gichiganus J. Allen (1903). Gizhiga, or 
Okhotsk blue hare 

1903. Lepus gichiganus Allen J. Report on the Mammals collect, in North-east. Siberia, Bull. Amer. 

Mus. Natur. History, XIX, 
1929. Lepus timidus gichiganus (Partim.') Ognev S. I. , Materialy po sistematike russkikh zaitsev 

(Material on the Systematics of Russian Hares), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, 

Vol. XXIII, pp. 491-496. 
1926. Lepus timidus gichi g anus Ognev S. I. , Mlekopitayushchie severo-vostochnoi Sibiri (Mammals 

of Northeastern Siberia), Vladivostok, p. 74 (partim.'). 

TYPE LOCALITY AND DEPOSITION. Gizhiga, shore of Sea of Okhotsk, No. 18286 male ad. , 11 
January 1901, N. G. Buxton. Tye in the collection of the New York Natural History Museum. 

MATERIAL EXAMINED: 20 specimens*. 

DIAGNOSIS. In summer fur, color of anterior part of muzzle, cheeks 
and forehead rust-straw-gray (between clay color and cinnamon, but with 
a strong dull gray tinge). 

Color of upper part of forehead and vertex darkened by black-brown hair. 

Color of back dark chestnut -brown- gray with a marked black tinge 
caused by the black-brown hair tips (with a straw-rust-colored subapical 
part) and some completely black-brown hair. 

Color of anterior inner margin of ears brown- gray with a slight straw 
tinge, like that of back. There are occasionally specimens in which 
anterior inner margin of the ears is markedly black. 

Color of lower part of back in front of tail deep gray (between dark 
mouse -gray and deep neutral gray). A broad dull grayish brown stripe 
across the chest. Winter fur white. 

Length of skull 90.2-95mm (M. 94.7mm) (93mm in the holotype**): 
zygomatic width 47-50 mm (M. 49 mm) (49 mm in holotype). 

Length of body 550-624 mm (582 mm average for 20 males, 577 mm 
average for 17 females); hind feet 146-173 mm (M. 164mm) average 
for 20 males, and 162 mm average for 17 females); ear (measured along 
concavity) 78-80 mm. 



J. A. Allen (1903) described the new species (which is actually a subspecies) from 37 pelts and skulls. 
According to J. Allen's measurements (1903), the length of the skull is 88-89 mm (94mm average for 
males and 91 mm for females). 



215 



SYSTEMATIC NOTES. In former publications I stated that L . t . 
gichiganus is widely distributed between Gizhiga and Penzhina (on the 
coast of the Sea of Okhotsk) and farther south along the coast of the Tatar 
Gulf and the Sea of Japan. I thus assumed that it occurs in the Amur 
Territory, the Ussuri Valley, north of Vladivostok and in forests along the 
Sikhote-Alin'. 

I now have collections from the coast of the Olyutorski Gulf (collected 
by A.S. Samorodov) and from the river Penzhina in the vicinity of Penzhino 
(collected by V. S. Bazhanov). A comparison between these small 
collections and specimens from the Maritime Territory showed that the 
hares from the Ussuri-Maritime Territory differ from the northern 
„ .„ specimens from Okhotsk. 

The following general rule was found: farther north the color of summer 
fur of blue hares becomes darker, with gray and blackish brown shades. 
It should be stressed that blue hares from the mountain region of the 
Maritime Territory Coast (from Sikhote-Alin') differ from those in the 
plains of the Maritime Territory. They closely resemble the Kamchatka 
hares in their darker color and in the black color on their ears. They 
apparently belong to the race gichiganus,. 

The Gizhiga hare is transitional between the very dark, black-eared 
L.t.kolymensis Ogn. and the much lighter, rust-colored L . t . 
mordeni Goodw. from the Maritime Territory. 

The Kolyma blue hare is also larger than the Gizhiga hare. According 
to S. A, Buturlin, the weight of the Kolyma hare is 2.7-3.1 kg. 

According to V. S. Bazhanov the weight of 44 adult hares caught near the 
village Penzhino was 1.6-3.0 kg- (2.560 kg average). 

According to A.I. Cherskii (in litt. ) the weight of hares from Maritime 
Territory is 2.6-3.4 kg, i.e., more than that of the Gizhiga blue hare. 

The Kamchatka blue hares which I saw in the ZMAN collection (e.g.. 
No. 12442 female ad., August 1847, Kamchatka, vicinity of Bol'sheretsk, 
Vosnesenski) are characterized as follows: l) intense brown color of back; 
2) saturated rust-brown color of top of head; 3) straw tinge on the nose; 
4) a pale straw ring around the eye; 5) lower part of back in front of 
tail gray. 

Lengthof skull 94-98 mm; condylobasal length 86.2-88.9 mm; zygomatic 
width 48.2-49.9 mm; length of nasals (along oblique line) 37.3-39.8 mm; 
length of upper row of molars 18.6-19.2 mm. 

In my opinion, the Kamchatka blue hares can be considered as belonging 
to L.t. gichiganus. In the ZMAN collection are some blue hares from 
the upper reaches of the Iman river on the Sikhote-Alin' (collected by K. K. 
Flerov), Color of back of these specimens intense chestnut-brown with 
black mottling caused by black hair tips of guard hair. Top of head with 
marked rust color (between cinnamon and sayal-brown, Ridgway, pi. 
XXIX 1912). Color of lower part of back not gray but duller, more 
blackish, differing from the chestnut -brown color of the back. 

These specimens and the Kamchatka blue hare are closely related to 
gichiganus . 

GEOGRAPHICAL DISTRIBUTION. J. Allen (1903) described L.t. 
gichiganus from a specimen from Gizhiga (on the Sea of Okhotsk Coast). 
Specimens from near the village Penzhino (along the Penzhina) and also from 



* Only two specimens of this series weighed 3 kg. 



216 



249 Apuka (Olyutorski Gulf) are in my collection. During the Chukchi 
expedition of the All-Union Arctic Institute, L. A. Portenko collected 

a good series of skulls from near Markov (near the Anadyr river). These 
skulls are small and generally resemble those ofL. t. gichiganus. 
Series of summer pelts from near Markov should be examined to determine 
their status. Judging from the limited material available L , t . 
gichiganus apparently occurs in this region. Transitional forms to a 
more northern race probably occur here. On 6 July 1932 near the 
Shchuch'ya river L. A. Portenko obtained a specimen with marked black 
anterior margin of the ears. Another specimen shot on 7 July 1932 did not 
show such black margins, and the color of the ears was generally like that 
of the back. The Gizhiga blue hare occurs in Kamchatka, along the northern 
coast of the Sea of Okhotsk and in Central Yakutia. The northern boundary 
of the range ofL.t. mordeni and the southern boundary of the range of 
L. t. gichiganus can only be determined after examination of more 
material. I have only a few specimens from the middle reaches of the Amur. 
More material may show that this region is inhabited by L . t . 
gichiganus . 

Blue hares from the Sikhote-Alin' (collected by K.K. Flerov) are closely 
related to gichiganus in their color (see above). It is possible that in 
the middle reaches of the Amur there occurs a form which is closely 
related to the Sakhalin form. On the other hand the Sakhalin blue hare 
differs very little from L. t. mordeni of the Maritime Territory. More 
material should be examined in order to establish the relationship between 
the Sakhalin subspecies L . t . orii Kuroda and the Maritime Territory 
subspecies L. t. mordeni Goodwin. 

345. Lepus timidus mordeni Goodwin (1933). Maritime territory 
blue hare 

1858. Lepus variabilis Schrenck L. v. , Reisen und Forschungen im Amur-Lande, I, S. 145-147 (partim.') 

1859. Lepus variabilis Maak R. , Puteshestvie na Amur (A Journey to the Amur), p. 107. 

1870. Lepus variabilis Przheval'skii N. M. , Puteshestvie v Ussuriiskom krae (A Journey to the Ussuri 

Territory), p. 257. 
1922. Lepus timidus gichi ganus Ognev S. I. , Materialy po sistematike russkikh zaitsev (Material on 

the Systematics of Russian Hares), Ezhegodnik Zoologcheskogo muzeya Akademii Nauk, Vol. XXllI, 

pp. 491-496. 
1927. Lepus timidus gichiganus Gassovski G. N. , Gilyui-Ol'doiskii okhotnich'e-khozyaistvennyi 

raion (The Gilyui-Ol'doi Commercial Hunting District), p. 527. 
1927. Lepus timidus Emel'yanov A. A. , Promyslovye zveri zemli orochei (Commercial Animals of the 

Land of the Orochs), in "Productive Resources of the Far East" (Proizvodstvennye sily Dal'nego Vostoka), 

IV, p. 262. 
1933. Lepus timidus mordeni Goodwin G. G. , Mammals Collected in the Maritime Province of 

Siberia by the Morden-Graves North- Asiatic Expedition. Amer. Mus. Novit. , 681, pp. 1-18. 

TYPE LOCALITY AND DEPOSITION. Monom river near village Troitskii in the Maritime Region. 
No. 85408 male ad. , 16 January 1930, G. Goodwin. Type in the American Museum of Natural History. 

MATERIAL EXAMINED: 8 pelts and skulls. 

DIAGNOSIS. In summer fur, color of anterior part of muzzle, cheeks 
and forehead light rust- straw-colored (between cinnamon-buff and pinkish 
cinnamon, but with a marked grayish tinge). 

250 Color of upper part of forehead, vertex and part between ears mixed with 
brownish- gray caused by blackish hair and blackish tips of many guard hairs. 



217 



Color of back generally rust-brown with a straw tinge and gray-brown 
mottling. 

Anterior inner margin of ears with mixed rust-colored and blackish 
hair; rust tone marked in this region. Color of anterior inner margin of 
ears almost like that of back, only slightly duller. 

Color of lower part of back in front of tail light gray with a brownish 
tinge (between deep mouse- gray and mouse-gray). 

A broad dull straw-gray stripe across the chest. 

Winter fur white. Roots of undercoat pale rust-straw colored. Hair of 
middle part of back about 45 mm long, that of flanks up to 65 mm long. 

Length of skull 88- 96 mm (M. 93.2 mm); zygomatic width 47-51 mm 
(M. 48.7 mm). 

Length of body 523-570 mm (M. 540.8 mm); hind feet 155- 161 mm- 
(M. 156.8 mm); ears (as measured along concavity) 95-97 mm (M. 95.6 mm). 

SYSTEMATIC NOTES. As I had not enough material, I wrote previously 
that L.timidus gichiganus is widely distributed south of the coast 
of the Sea of Okhotsk and also that it occurs in the Maritime Region and the 
Ussuri Territory. After comparing specimens from Ussuri and the 
southern Maritime Territory with specimens from the Okhotsk Coast I 
came to the conclusion that L. t. gichiganus differs from L . t . 
mordeni, which is distributed throughout the Maritime Territory. 

G.Goodwin (1933) who described L. t. mordeni from one winter 
specimen could not give a precise description. He compared the skull 
and teeth of L. t. mordeni with those of L. t. gichiganus, and found 
that the skull of the Primorye hare is smaller, but that its teeth are larger. 
• According to Goodwin, the width of the lower incisors of L. t. mordeni 
is 3 mm and ofL.t. gichiganus 2.5 mm. We obtained the following 
measurements in our material: width of lower incisor of L. t. mordeni 
3; 3; 3; 3.1; 3.2; 3.2; 3.4mm (M. 3.14mm); of L . t . gichiganus 2.8; 
2.9; 3.3 mm (M. 2.75mm). These differences are very small and not 
significant. According to Goodwin the transverse width of the crown of 
the first upper premolar of L. t. mordeni is 5 mm against 4-4.5 mm in 
L. t. gichiganus. I obtained the following measurements from our 
material: in L. t. mordeni 4.3; 4.5; 4.8; 5; 5; 5.1mm (M. 4.63mm); in 
L. t. gichiganus 4; 4.2; 4.2; 4.5 (M. 4.2 mm). According to Goodwin, 
the width of the largest upper molar of L. t. mordeni is 6.8 mm and 
5-6 mm (M. 5.7 mm) in L.t. gichiganus. I obtained the following 
measurements from our material: in L. t. mordeni 5.7; 6; 6; 6; 6; 6.3; 
6.5 mm (M. 6 mm); in L. t. gichiganus 6; 6.1; 6.2; 6.2 mm (M. 6.1 mm). 

These data show that the differences of the teeth of the Gizhiga and 
251 Maritime Territory blue hares are very small. The conclusions of 

Goodwin, reached after comparison of a single skull of L. t. mordeni 
with a skull of L. t. gichiganus are, to say the least, premature. 

The following real differences exist between specimens from Gizhiga 
and those from the Maritime Territory: 1) color of summer fur of 
Gizhiga subspecies darker and grayer; 2) measurements and weight of 
Gizhiga blue hare smaller. Examination of more material may prove that 
the measurements of the skull of the Gizhiga blue hare are also smaller. 
The differences between L. t. gichiganicus and L. t. mordeni are 



* According to G.Goodwin, the hind feet are 180mm long in the holotype ofL. t. mordeni. 



218 



thus very small. They cannot, however be placed into one subspecies at 
present. More material is necessary in order to make the differentiation 
more specific. 

GEOGRAPHICAL DISTRIBUTION. This subspecies is widely distributed 
in the Maritime Territory, the valley of the Ussuri, Khungari and perhaps 
along the middle and lower reaches of the Amur. In my collection is a 
specimen from Nikolaevsk-on-Amur which is closely related to L . t . 
mord eni . 

It is difficult to say how far north this subspecies is distributed. Its 
range seems to pass into that ofL.t.gichiganus. As stated above, a 
darker form which hardly differs from L. t. gichiganus occurs in the 
Sikhote-Alin Range. 

The southern boundary of the range ofL.t.mordeni is not known. 

A.de Sowerby (1923) did not record this subspecies from Manchuria. 
The late A, I. Cherskii (in a letter of 17 Feb. 1914) wrote as follows on the 
distribution of the blue hare in the Ussuri and Maritime Territories: "The 
blue hare is sold in the Vladivostok market as 'imported game'. According 
to Maksimovich (apud Radde), the southern boundary of its range passes 
north of Lake Hanka. I found hares of this subspecies in 1911, in the upper 
reaches of river Odarka, in the Lake HankaSystem, and 27 km east of the 
Yevseyevka Station of the Ussuri Railway. This moves the southern 
boundary south by a whole degree. In the mountain forest of the upper 
reaches of the Odarka river blue hares are more common than the 
Manchurian hare. " 

346. Lepus timidus orii Kuroda (1928). SaJkhalin blue hare 

1889. Lepus timidus Nikol'skii, A.M., Ostrov Sakhalin i ego fauna pozvonochnykh zhivotnykh (The 
Vertebrate Fauna of Sakhalin Island), pp. 126-127. 

1890. Lepus variabilis Suprunenko, Pozvonochnye Sakhalina (Vertebrates of Sakhalin), p. 39. 
1923. Lepus tim i dus Sowerby A. , The Naturalist in Manchuria, II-III, pp. 173-174(SakhalinIsland). 
1928. Lepus timidus o r i i Kuroda N. , The Mammal Fauna of Sakhalin, — Journal of Mammalogy, 

VoL9, No. 3, pp. 223-224. 
1931. Lepus timidus saghaliensis Abel., A Synopsis of the Leporine Mammals of Japan, Joum. of 

Science of Hiroshima University, ser. B, div. I, pp. 49-50. 

TYPE LOCALITY AND DEPOSITION. Sakhalin Island, Naihoro(westerncoastof central partof Japanese 
Sakhalin), No. 1234, male ad. , 6 May 1926, Nagamichi Kuroda's collection. 

MATERIAL EXAMINED: 4 specimens. 

252 DIAGNOSIS. Owing to lack of material I cannot give a diagnosis of this 
subspecies, the existence of which as a systematic unit should be verified. 

In summer fur, color of upper part of forehead, vertex and region 
between ears dark brownish gray. 

Color of back brownish gray with a straw tinge caused by yellowish 
straw colored subapical bands of hair. Tips of hair blackish brown. 

Color of anterior inner margin of ears gray-brown with a straw tinge, 
little darker than back. 

Color of lower part of back in front of tail brownish gray, almost without 
straw tone, and thus duller and darker than middle of back. 

A dull straw gray stripe across the chest. 

Winter fur and under fur white. 

SYSTEMATIC NOTES. The Sakhalin subspecies was described by 
N. Koruda (1928) as L. t. orii. According to N. Koruda it differs from 



219 



L. t. gichiganus as follows: 1) skull large, tail and hind feet shorter; 

2) ears of L . t . o r i i longer than those of L.t. gichiganus; 3) in 
summer fur, color of forehead and sides of head pale rust-brown and 
reddish, (gray-brown inL. t. gichiganus*); color of back, flanks and 
haunches grayish brown or reddish brown (dark gray in gichiganus). 
Lower part of back of L . t . o rii usually grayer and darker than middle of 
back. 

In 1931 I.Abe again described the L. t. orii from a specimen from 
Otomari [Korsakov] (Japanese Sakhalin), as L. t. saghaliensis. He 
did not mention N. Kuroda's description and only referred to Kishida's 
book "An Explanatory Illustration of Japanese Mammals", 1924, in 
Japanese. In this work the Sakhalin hare is called Lepus gichiganus 
subsp. ? 

According to I, Abe the Sakhalin hare is characterized as follows: 
1) elevated postorbital processes; 2) ears longer than inL. t. gichiganus; 

3) hind feet longer, 180 mm long. This last character differs from the 
measurement given by N. Kuroda. 

On Hokkaido (Yezo) L . t im id u s ainu Barrett -Hamilton occurs, which 
I.Abe considers, without any justification, as a separate species. He 
gives the following characters as typical for Lepus ainu: 1) low 
posterior part of the postorbital processes (as inL. hibernicus); 
253 2) large skull: length of skull of the hare from Hokkaido 98-101 mm. 

All these data indicate that a subspecies which is closely related to 
L . t . orii and possibly identical with it occurs on Yezo. The relationship 
between the Sakhalin hare and hares from Hokkaido will have to be 
determined after examination of more material. 

In conclusion I give measurements of the Sakhalin blue hare and the 
Hokkaido blue hare (according to N. Kuroda and I. Abe). 



Skull measurements of Le pus timidus or i i Kuroda (in mm) 





















<u 


<u 






























J3 








rt 




Ji 


Xi 








T3 


j: 


„ 


H 


42 


di 


'T3 w 

S 









■s 

CO 


5 

o 


T3 


15 


a 




3 


B o 


B 


u 




B 




■B ° 


"o 


S 


% 


-B -S 





B 
z 


a 

2 




B 




O 

*-• 

2 


^1 


a 


43 ^ 


J3 


II 


op 
X 


660 


100 


80.5 


51 


37.1 


- 


46 


21 


17 


80 


53.1 


1229 


94.5 


76 


51.5 


36 


16.5 


40.1 


21.5 


17.9 


74.5 


48.5 


1230 


94.5 


76 


49.9 


32 


15 


39 


20.5 


18 


76.5 


49.5 


1233 


97.2 


77.5 


52 


34.1 


16 


41.5 


20.5 


17.5 


75 


50 


1234 


99.5 


79 


50 


31.1 


13.5 


42.1 


20.1 


19 


77.5 


51.5 


Type 























+ N. Kuroda gives an inexact description of the color of the forehead and sides of the head of L. t. 

gichiganus. In specimens examined, these parts were rust-straw-gray with variable degrees of rust 
tone. The color of the back of gichiganus is not dark gray, but a dark chestnut-brown-gray color. 



220 



Measurements ofL. t. saghaliensis Abe (male No. 28) 









H 










<u 




















1 


■s 


^ 




w 


rt 


r 


■(S 


4) 
> 






u 




:? 








t 
a 








a 


1 


o 

^^ 
O 


§ 


.i9 
o 


.3 
1 




a, 

3 






o 

•2 


•3 






"S 


t? 


^ 


^ 


■?! S 


"S 


<4H (H 

a 


o ;^ 


rt 

p 


° ^ 


s 2 


O o 


o 


'S 









60 




,£ 
■S 




a H 


5 B 

73 rt 


a 


5 E 


11 




^^ 


5 M 


J3 
CO 


■3 

bO 


Z 


N 


^ & 


^ 


;ii 


^ g 


^ >2 


H-l 


^ ^ 




d 




^ * 


:ii i 




^ 


98.5 


79 


49.5 


17 


19 


40 


43 


16 


9 


10.5 


23.5 


25 


16.5 


115 


95 


180 


149 



The measurements of specimen in the ZMAN collection are as follows 
(in mm) : 

Length of skull 94. 3 

Zygomatic width 46.8 

Width of skull behind orbit 15.2 

Width of nasals at the base 21.2 

Length of upper tooth row 19. 2 

Length of mandible 65. 8 

Height of mandible 44. 1 

The data on the Sakhalin blue hare show that there is insufficient 
material to clarify the status of this form. 
254 The descriptions by Kuroda and Abe are not complete, being based on 
too little material and giving accidental characters which are hardly typical. 
The Sakhalin blue hare is certainly closely related to the Primorye hare 
(mordeni). It is however impossible to consider them as identical. 
L. t. mordeni is closely related to L. t. gichiganus in various 
characters. L.t. mordeni is probably only a lighter natio of 
gichiganus, and the Sakhalin subspecies is probably synonymous with 
the Primorye subspecies. The Okhotsk -Maritime Territory and Sakhalin 
are thus probably inhabited by one species with a lighter, southern natio. 

GEOGRAPHICAL DISTRIBUTION. The subspecies occurs only on 
Sakhalin Island. It possibly also occurs on the continent, along the middle 
reaches and northern part of the Amur (near the coast). The small 
collections examined from the lower reaches of the Amur were closely 
related to L. t. mordeni 

347. Lepus timidus kolymensis Ogn. (1922). Kolyma blue hare 

1884. Lepus variabilis Bunge A. , Naturhist. Beobachtung. im Lena- Delta. M6m. Biolog. de 

I'Academ. Imp. St-Petersb. , XII, S. 33 (Lena estuary). 
1887. Lepus variabilis Bunge A. , Berichte iiber die Expedition nach Neusibirischen Ins. S. 39-40 

(river Yana, Yana estuary). 
1914. Lepus tschuktschorum Allen G. I. Proceedings New Engl. Zoological Club, V, p. 66 (Bol'shoi 

Baranovyi). 
1922. Lepus timidus kol ym ensis Ognev S. I. , Nekotorye dannye po sistematike russkikh zaitsev 

(Data on the Systematics of Russian Hares), Biologicheskie Izvestiya, Vol. I, p. 106. 
1926. Lepus timidus kol ym ensis Ognev S. I. , MIekopitayushchie severo-vostochnoi Sibiri (Mammals 

of Northeastern Siberia), pp. 72-77. 

TYPE LOCAUTY AND DEPOSITION. Pokhodskoe Village, 80km below Nizhne-Kolymsk along the 
Kolyma river. Type No. 8028 female 4 June 1905, S. A. Buturlin, ZMAN collection. 

MATERIAL EXAMINED: 15 pelts and skulls. 



221 



DIAGNOSIS. In summer fur, color of anterior part of muzzle, cheeks 
and forehead dull brownish gray with a slight rust tinge. 

Color of upper part of forehead, vertex and region between ears darker 
as a result of presence of black hair and black -brown tips of numerous 
hairs which have straw- gray subapical bands. 

Color of back dark, dull brownish gray with a slight straw-gray tinge. 

Anterior inner margin of ears almost completely black. Hairs in this 
area have dark gray roots, straw-gray subapical bands and broad black 
tips. They are so strongly developed that they give an almost black color; 
this is typical for this subspecies. 

Color of lower part of back in front of tail dark gray. A dull gray stripe 
across the chest. 

Winter fur and undercoat completely white. 

Length of skull 93.2 -99.2 mm (M. 96.6mm); zygomatic width 47.7-51 mm 
255 (M. 49.3 mm). 

SYSTEMATIC NOTES. As stated above, the Kolyma hare is larger 
than the Gizhiga hare, as shown by its weight and the size of its skull. 

The Kolyma hare is characterized as follows: 1) color of back black 
and without any rust tone; 2) blackish tone especially marked in middle 
of back, on ears and occiput; 3) head and cheeks dull brownish gray. 

The Kolyma blue hare is closely related to the Gizhiga hare. It is a 
darker race confirming the rule that farther north the summer fur becomes 
darker and blacker, 

GEOGRAPHICAL DISTRIBUTION. The Kolyma blue hare occurs along 
the Kolyma river and apparently also in the Indigirka Valley. In the east, 
beyond the Stanovoi Range to the Anadyr river in the north, L . t . 
gichiganus occurs. Another subspecies possibly occurs in the valley 
of the Yana river. According to S. A. Buturlin (1913), the hares in this 
region are very small and weigh only 1.4-1. 6kg. This could be decided if 
we could obtain material from the region between the Tas-Khayakh-Takh 
and Verkhoyansk ranges. 

348. Lepus timidus begitschevi Koljuschev (1936). Taimyr blue 
hare 

1933. Lepus spec? Kolyushev 1. 1. , Materialy po faune i promyslu zapadnogo ugla Taimyrskogo 

poluostrova (Material on the Fauna and Animal Trade of the Western Comer of the Taimyr Peninsula), 

Materialy po izucheniyu Sibiri, Vol. IV, pp. 15-16. 
1936. Lepus timidus be gi tsch e vi Kolyushev 1. 1. , Mlekopitayushchie krainego severa, zapadnoi i 

srednei Sibiri (Mammals of Extreme Northern, Western and Central Siberia), Trudy Biologicheskogo 

nauchno-issledovatel'skogo instituta, Vol. II, Tomsk, pp. 304-306. 

TYPE LOCALITY AND DEPOSITION. Western coast of the Pyasina Gulf. Type obtained on 7 March 1930, 
in 1. 1. Kolyushev's private collection. 

MATERIAL EXAMINED: 22 pelts (most of them with skulls). 

DIAGNOSIS. Muzzle grayish brown with a marked admixture of rust 
colored hair, especially marked on cheeks. 

Back pale gray. Neck and sacral region with a marked ash-blue tinge. 
Scattered black hairs on the back, especially in the lower parts. 

Ears pale gray with a weaJc rust tinge and a few white hairs. 

Length of skull 98 -108 mm (M. 103,4 mm); condylobasal length 88-95.2 mm 
(M. 92.6 mm); zygomatic width 50.6-54 mm (M. 52,7mm); length of upper 
row of molars 18. 6-20. 5mm (M. 19.5mm). (See Figures 130, 131.) 



222 



Length of body with head 575-740 mm; hind feet 143-171 mm; ear 
80-94 mm. 

SYSTEMATIC NOTES. An excellent series of Taimyr blue hares 
collected by E. O. Yakovlev is in the ZMAN collection. Some of these hares 
originate from the eastern Taimyr and some from the Khatanga Gulf. They 
correspond in color and measurements to 1. 1. Kolyushev's description. 

In the Moscow Zoological Museum is a series of 8 pelts (unfortunately 
256 without skulls) which were obtained by A. F. Chirkova in the lower reaches 
of the Khatanga in the tundra near the village Khatanginskoe, 

These hares differ from east Taimyr blue hares in their color and 
smaller size. 

Description. In summer fur, color of middle part of anterior half of 
muzzle and cheeks very pale gray- straw-yellow (close to pinkish buff with 
a cinnamon-buff tinge). This color is much duller because of gray and 
blackish hair. Sides of muzzle and lower part of cheeks dull gray -straw - 
white. Color of forehead duller straw-gray with a slight rust tinge. Eye 
surrounded by a completely white ring. 

Color of back dull gray (mouse-gray) with light straw mottling. This 
mottling creates a straw -gray color on the back, very similar to drab. 

Flanks, lower part of back in front of tail and stripe across chest gray 
(light mouse -gray) with a slight blackish tinge caused by black hair tips or 
gray -black subapical bands. 

Color of anterior inner margin of ears straw -gray -brown, similar to 
color of back. * 

Length ofbody 470- 550 mm; hind feet 132-157 mm (M. 141.6 mm); ear 
(measured along concavity) 82-93 mm (M. 87.7 mm). 

The skulls of the series have been lost. We therefore cannot say 
whether these specimens are identical with those collected by E. O. 
Yakovlev. It seems that blue hares from the lower reaches of the Khatanga 
are different from the Taimyr specimens. 

The Khatanga blue hares are characterized as follows: 1) dull straw- 
gray -yellow color of head; 2) gray color of back with slight straw mottling; 
3) light anterior inner margin of ears. This form differs from L . t . 
timidus in the lighter gray fur without the brownish tones which are well 
marked on the back of north European specimens. 

Compared with the Kolyma subspecies, the Khatanga blue hare is grayer 
and lighter, without black color on the ears and the upper part of the back. 

It differs from the Taimyr -Pyasina form in its small size, in some 
color characters such as the absence of bluish gray tones on the back, and 
in the color of the head. 

If the very large hares from the Gyda Peninsula (described by S. P. 
Naumov, 1931) are identical with the Pyasina and Taimyr blue hares, 
L. t. begitschevi should also include hares from near the village 
Dudinka (lower reaches of the Yenisei) a specimen of which is in the ZMAN 
collection. Specimens from the Taz and Kureika rivers are not identical 
with L. t. begitschevi. 

In my collection are specimens from the Taz river collected by R. E. 
Kol's. The skull of these specimens is not large, and closely resenables 
that of specimens from the Kolyma river. F.F. Shillinger caught a 



* All specimens of the series examined were either in summer or in transitional fur (at the beginning of the 
autumn molt). I did not see any winter specimens of this form. 



223 



specimen near the Kureika river (2 5 Jul. 192 5), Although it was caught in 
late summer, it had much white fur on its head, ears and back. Black 
spots of summer fur could be seen among the white fur. The inner side 
of the ears was completely black. 

The full summer fur of this form would certainly be very dark. A, 
specimen caught by R. E. Kol's on 30 May 1927 near the Taz river had 
retained almost completely its white winter fur. The dark summer fur 
showed on the ears and on some parts of the back. The ears were also 
black, but lighter than those in the specimen from the Kureika river. 




FIGURES 130-131. Taimyr blue hare, Lepus timidus begitschevi Koljuschev. Specimen 
No. 18786, 4 June 1933. Taimyr District, Kardo Cape, Khatanga Gulf, E.G. Yakovlev, ZMAN 
collection 

Drawing by Yu. A. Kostylev 

New material will have to be examined in order to decide which 
2c-n subspecies occurs along the Taz, Yenisei and Kureika rivers, and south of 
the Lower Tunguska. 

GEOGRAPHICAL DISTRIBUTION. This form occurs on the Geidal 
Peninsula, throughout Taimyr, in the lower reaches of the Yenisei near the 
village Dudinka and in the east along the Khatanga Gulf coast. 



224 



A different form possibly lives along the lower reaches of the Khatanga. 

349. Lepus timidus t schukt schorum Nordquist (1883). Chukchi 
blue hare 

1883. Lepus timidus L, v ar. tschuktschorum Nordquist O. , Antecknigar och studier till Sibirska 
sshaiskustens Daggdjursfauna Vega Expeditionens Vetenskapliga lakttagelser, Vol.11, pp. 84-90, figs. 8, 
9-10; p. 88. 

1903. Lepus tschuktschorum Allen 1. , Report on the Mammals Collect, in Northeast Siberia. Bull. 
Americ. Mus. Nat. History, XIX, p. 159-160 (near Cape Chaplin — Indian Point). 

1926, Lepus tschuktschorum Ognev S. 1. , Mlekopitayushchie severov-vostochnoiSibiri (Mammals of 
Northeastern Siberia), p. 77. 

1927. Lepus tschuktschorum (partim .' ) Sokol'nikov N. P. , Okhotnich'i i promyslovye zveri 
Anadyiskogo kraya (Game and Commercial Animals of the Anadyr Territory), Byulleten' Moskovskogo 
obschestva ispytatelei prirody, XXXVI, p. 133. 

TYPE LOCAUTY AND DEPOSITION. Chukot Peninsula. Type in the Stockholm Museum. 
MATERIAL EXAMINED: 10 specimens. 

DIAGNOSIS. In summer fur, color of anterior part of muzzle, cheeks 
and forehead dull brownish gray with a slight straw -rust tinge (buff -brown 
with a slight rust tinge). 

Color of upper part of forehead, vertex and region between ears much 
darker because of the large admixture of completely black hair, and black 
bands on hair with straw -yellow subapical bands. 

Color of back dull straw -gray -brown with a black tinge which forms a 
dull mottling with straw shades. 

Color of flanks and lower part of back grayer (more ash colored) than 
middle part of back. 

Anterior inner margin of ears with marked black tone; straw mottling 
little marked. 

A dull gray stripe across the chest. 

Winter fur and undercoat white. Hair unusually luxuriant, delicate, 
silky and long. Hair of back 62 mm long, hair of flanks and underparts 
80 mm. 

Skull massive, with broad nose and zygomatic arches. Length of skull 
98-106 mm (116 mm according to Nordquist) (M. 103 mm); zygomatic width 
51-54. 5mm (57mm according to Nordquist) (M. 52.8mm). See Figure 125. 

Length of body 610-680 mm; hind feet 150- 165 mm. 

SYSTEMATIC NOTES. The range of this subspecies is adjacent to that 
of the Kolyma subspecies. It differs from the latter in its larger size, and 
its larger and more massive skull. Summer color of eastern Chukchi hare 
lighter and grayer than that of the Kolyma blue hare. Winter fur of 
tschuktschorum more luxuriant and longer. This subspecies differs 
clearly from the Taimyr blue hare (L . t . begits che vi) in its color. 
259 GEOGRAPHICAL DISTRIBUTION. It occurs only in the northeastern 
periphery of the Chukchi Peninsula. A form which is apparently identical 
with L. t. gichiganus occurs along the river Anadyr near Markov. 
D.G. Elliot ("Synopsis of the Mammals of North America", Publ. of the 
FieldNat. Hist. Museum, Zoolog. Ser. , Vol.11, Chicago, p. 271) writes 
that Lepus tschuktschorum occurs in northern Alaska from the 
estuary of the Kuskokvim river up to the Arctic Ocean. The occurrence 
of the Chukchi blue hare in Alaska has not been confirmed in later 
publications of American zoologists . In Alaska occur Lepus othus 



225 



Merriam and L. poadromus Merriam. The relation of these species to 
L. timidus, especially to the Chukchi subspecies, will have to be 
determined by comparison of material. 



Genus Allolagus Ognev 

1922. Lepus Ognev S.I. , Materialy po sistematike nisskikh zaitsev (Materials on the Systematics of 

Russian Hares), Ezhegodnik Zoologicheskogo Muzeya Akademii Nauk, Vol. XXIII, p. 485. 
1929. Allolagus Ognev S.J. , Zur Systematik der rusf .schen Hasen, Zoologlscher Anzeiger, Bd. 84, H. 1/4, 

S. 72. 
1936. Lepus (C aprol agus) Zolotarev N. T. , Mlekopitayushchie basseina reki Imana (Mammals of the 
River Iman Basin), p. 69. 

Ears (measured from occiput) about as long as, or slightly longer than, 
head. 

Dorsal side of tail black -brown, clearly different from general gray 
color of the other part of tail. 

Hind legs short. When the animal stands, the hind legs seem little 
longer than the forelegs. 

Mesopterygoid fossa about as wide as the length of the bony palate. 

Adults without interparietal. 

Postorbital processes elongate triangular. The anterior angles 
relatively short. 

Width of interorbital space posterior to the postorbital process (in its 
narrowest place) 60.4 % of the width of both nasals in their posterior third. 

Length of nasals along the line connecting the outer posterior corner 
with the inner anterior corner about as long as, or slightly shorter than, the 
frontal suture. 

Width of posterior nasal foramen about as long as the bony palate. 

Width of incisive foramina (in their widest part) about as great as the 
length of the bony palate. 

Incisive foramina about as long as the upper diastema. 

Type species: Lepus mandshuricus Radde. 
261 GEOGRAPHICAL DISTRIBUTION. Southern Ussuri Territory, 
Manchuria, and the Hondo [Honshu] and Kyushu Islands. 

161. Allolagus mandshuricus Radde (1861). Manchurian hare 

1861. Lepus mandshuricus Radde G. , Neue Saugethier-Arten aus Ost-Sibir. , M61. Biol. Bullet, de 
I'Academ. Imp. de St-Petereb. , III, S. 684-686. 

1861. Lepus mandshuricus Maximowlcz H. , Zoologisch. Nachricht, M6l. Biol., Ill, S. 693. 

1862. Lepus mandshuricus Radde G. , Reisen im SUden von Ost-Sibir. , S. 215-223. 

1870. Lepus mandjuricus Przheval 'ski N. M. , Puteshestvie v Ussuriiskom krae (A Journey to the Ussuri 

Territory), Sankt Petersburg, pp. 257-258. 
1922. Lepus mandshuricus Ognev S. I. , Materialy po sistematike russkikh zaitsev (Material on the 

Systematics of Russian Hares), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, Vol. XXIII, 

pp. 485-491. 

1922. Lepus mandshuricus subphasa melanonotus Ognev S. I. , op. cit. p. 489. 

1923. Lepus mandshuricus Sowerby A, de C. , The Naturalist in Manchuria, II-IIl, pp. 171-172. 
1929. Allolagus ( m andshurtcus) Ognev S. J. , Zur Systematik der russischen Hasen, Zoologisch. 

Anzeiger, Bd. 84, H. 1/4, S. 72. 
1936. Lepus (C aprol agus) m andshuricus Zolotarev L. T. , Mlekopitayushchie basseina reki Imana 
(Mammals of the River Iman Basin), pp. 69-70. 



226 



260 




FIGURES 132-135. Skull of 
Manchurian hare, Allolagus 
mandchuricus, female 
2 May 1914, Peter the Great Gulf, 
A.I. Cherskii, S.I. Ognev's 
collection 

Drawing by Yu. A. Kostylev 



227 



PLATE IV 




Allolagus mandshuricus Radde. Winter fur 




Allolagus mandshuricus Radde. A melanic specimen in winter fur 

Drawing by V. A. Vatagin 



228 



261 



262 



NAMES: Manchurian Hare (English) ; Mandchurischer Hase (German). 

LOCAL NAMES: Tolo, taula, tolau (Birar Evenki-Tungus); borto-gormako (Nanai-Gold); borta- 
gurmakhung (Manchuria). 

TYPE LOCALITY AND DEPOSITION. Bureya Mountains. Type in ZMAN collection. 

PRINCIPAL FIGURES. Radde G. , Reisen im Siiden von Ost-Sibirien, 1862, S. 215-223, Taf. VIII, 
Fig. 1 a-d (entire animal in color, skull). 

IvlATERLAL EXAIvlINED: 19 pelts and skulls. 

DIAGNOSIS. Head relatively large. 

Outstretched ear (measured from occiput), as long as or slightly longer 
than head. 

Tail (including terminal hair) 60-70% as long as hind foot (without claws). 

In summer and winter fur, dorsal side of tail black -brown, merging with 
the general gray background of the other part of the tail. 

Hair does not become white in winter, but grayer than the summer fur. 
Winter fur of melanic specimens black -brown. 

Fur soft. Guard hair soft, straight and thicker than in the desert hare. 
Guard hair not curly. Undercoat wavy and curly, 

I could not determine the number of teats from the material at my 
disposal. 

Length of nasals, measured along the line connecting the inner anterior 
corner with the outer posterior corner, about as long as, or shorter than, 
the frontal suture. 

Length of postorbital processes about 51,5% of maximal width of both 
nasals measured in their widest part. 

Posterior third of incisive foramina widened. Their lateral margins 
straight, without protuberances. 




FIGURE 136. Manchurian hare, Allolagus mandchuricus Radde. 

Drawing by A. N. Komarov 

Bony palate measured lateral to anterior process about as long as the 
incisive foramina measured in their posterior third are wide. 



229 



Bony palate about as long as the mesopterygoid fossa in its widest part. 

Lower diastema about 86% of the length of the lower tooth row, that is. 
shorter than the tooth row. This was not found in any of the other hares. 

Anterior lower premolar slightly inclined backwards in profile. 

Mandibular condyle massive, slightly inclined backwards. 

Posterior-ventral plate of mandible from corner of angular process to 
notch at beginning of horizontal mandibular process very large. Distance 
between angular process to notch of horizontal mandibular process 1 V2 
times as large as distance between anterior alveolus of premolar to tip of 
lower incisor. 
263 Bullae osseae very small, little inflated and slightly protruding 
forward (Figures 132-139). 

Length of skull 82,l-89mm (M. 85.4 mm); zygomatic width 40-44 mm 
(M. 41.9mm); length of nasals 32 .8-37.9 mm (M. 35.5 mm); width of nasals 
14-18mm (M. 15.7mm). 




FIGURES 137-139. Skull of A llol a gus mandchuricus Radde, No. M. 3069, 
Maritime Territory, S. 1. Ognev's collection 



230 



Length of body and head 420-540mm (M. 473.5mm); hindfeet (without 
claws) 117-130mm (M. 123.4mm); ear (measured from occiput) 84-95mm 
(M. 88.9mm); tail with terminal hair 100-160mm (M. 115.2 mm). 

Weight 1650-2600 gms. 

ADDITIONAL DESCRIPTION. The diagnosis of the genus and the 
description of the species show that the skull of the Manchurian hare has 
264 some specific features of which the following are the most characteristic: 
1) width of bony palate; 2) narrow incisive foramina; 3) small size of 
bullae osseae; 4) short diastema compared with the massive and long rows 
of molars; 5) nnassive naandibula, especially its posterior- ventral plate; 
6) relatively small orbits. 

COLOR. WINTER FUR. Color of back with a mixture of light rust- 
brown and a marked black tone (plate IV, upper figure). 

Color of occiput like that of back. Forehead and anterior part of nose 
darker because of the greater admixture of black hair tips. Area around 
eyes rust -white. Cheeks below the eyes with a diffuse blackish pattern. 
Sides of cheeks whitish. Muzzle dull brown-rust-gray. Two irregular 
white spots below the nostrils. 

Neck rust -chestnut -brown, more rust colored than fawn (Ridgway, 
plate XL). Throat white with a light straw-bay tinge. Chest pale straw 
colored with vinaceous tinge (closely related to avellaneous and vinaceous- 
buff, Ridgway, plate XL). This color also covers the forelegs,- flanks, 
lateral parts of groin and part of the hind feet. Belly and groin white. 

Ears short. Anterior inner margin of ears variegated with a admixture 
of blackish and rust -brown hair. Inner sides of ears with a rich rust-ocher- 
brown tone. Tip of ears (about 3 5 mm on inner and outer sides) black or 
black -brown. 

Tail with luxuriant whitish gray hair. Dorsal side of tail blackish brown. 

This type of color of winter fur is the most common. There also occur 
grayer specimens with an ash tinge. The general color of the back has a 
variegated mixture of a whitish ash-gray tone with a blackish tinge. Gray- 
white tone predominates on the lower part of back. 

Anterior part of forehead with a well marked rust tinge. Area around 
eyes white. Cheeks gray -white. Region behind the eyes pale whitish rust. 
Occiput with a dull rust -brown tinge. 

Throat white. Chest with some pale rust hair and black hair giving a 
grayish shade. Belly and groin white. 

Color of anterior inner margin of ears rust -straw -gray. Inner parts of 
ears whitish. Tips of ears rust-straw colored, margins black. Ears near 
the base rust -gray. 

Forelegs rust-straw colored. This color is weakly marked on flanks 
as two spots partly entering the inguinal region. Hind legs dirty straw- 
gray. 

SUMMER FUR. This description is based on a young specimen. Summer 
hair shorter and coarser. Its general color is intense rust -brown with a 
straw -black tinge. Color of forehead and occiput almost like that of back. 
Eyes surrounded with rust-gray hair. Cheeks with much black hair. 
265 Neck dull rust -brown, grayer than sayal -brown (Ridgway, 1912, plate 
XXIX). Belly whitish. 

Anterior inner margin of ears with short brownish gray hair. Outer 
fringe of ears rust-ocher. Outside of ears pale rust -brown. Tail gray 
with blackish brown dorsal side. 



231 



Melanic specimens also occur. These are rare in the Ussuri Territory, 
but farther south in Manchuria (the Kirin Province) they are as common as 
specimens of normal color (A. Sowerby, The Naturalist in Manchuria, 
II-III, p. 171-172, 1923). 

I had the opportunity of studying two partly melanic specimens. One of 
them (No, M. 3081 female, in my collection) was sent to me by the late A.I. 
Cherskii, who bought it in the Vladivostok market. The other specimen 
was given to the Moscow Zoological Museum by V. V. Bogomolov who shot 
it on the Sikhote-Alin Range (Maritime Territory) 15km from the village 
Sol'skii (No. S.4037). Both specimens belong to the same color type (pi. IV, 
lower figure). 

Color of top of head, area around eyes, cheeks, back, and flanks shining 
black with a slight brownish tinge. 

Throat and lips dirty whitish with a slight straw tinge. A well marked 
transverse black stripe, about 30 mm wide, beneath the throat. Chest pale 
pinkish straw (between light pinkish cinnamon and vinaceous-buff, Ridgway, 
pi. XXIX and pi. XL). Belly white. A diffuse black stripe across the lower 
part of the belly. Groin white. Inner and outer sides of ears black. Outer 
fringes of ears dull straw -rust -ocher. 

Forelegs black -gray-brown. Hind feet with a mixture of pale straw -ocher 
and black -brown hair. Tail dark gray with blackish dorsal side. 

Melanic specimens are characterized by the presence of some very long 
silver-straw guard hair contrasting with the general black background of 
the back and on the flanks. 

The melanic specimens are smaller than specimens of normal color. 
This was confirmed by A. Sowerby (1923), 

SYSTEMATIC NOTES. The systematic position of the Manchurian hare 
is not clear. In the past it was placed in the genus Lepus or in the genus 
C ap r ol agu s . 

After careful examination I came to the conclusion that the genus 
Caprolagus is well characterized. It should not be confused with the 
genera Lepus and Allolagus. 

In his book "Fossil and Recent Lagomorpha" (1898) C.J.Forsyth Major 
includes the following genera in the "Caprolagus group": 1) 
Caprolagus; 2)Nesolagus; 3) O ry c t o 1 agu s ; 4) Sy 1 vil agu s . 
He includes in Sylvil agu s the following genera: Limnolagus, 
Romerolagus, Tapeti, Sylvilagus pr.dict. 

The Caprolagus group thus includes all relatively short -legged and 
short -eared hares with a primitive, elongate bony palate. 

This grouping seems quite artificial. In addition to wide geographical 
separation, each of the species of this group has its specific characters. 
For instance, the genus Oryctolagus differs sharply from Caprolagus 
and Sylvilagus . 

The tajxonomic relationship between the genera Caprolagus and 
Allolagus is best demonstrated in table form. 

C. hispidus Pearson (1848) (from the southern Himalayas) belongs to 
genus C ap rol agu s . According to Glover Allen (192 7), the following 
species should be placed in the genus Caprolagus: C. sinensis 
Gray (1833-1834) from southern China, and C . s i ne n s i s flaviventris 
G.Allen (1927) from Fukieu (southeastern China). 

The monotypic genus Pentalagus Lyon which includes only the 
species P ent al a gu s furnessi Stone (1900) from the Ryukyu Islands is 
closely related to Caprolagus . 

232 



Genus AUolagus Genus Caprolagus 

Eare measured from occiput about as long as Eais much shorter than head, 

the head or longer. 

Fur soft. Fur coarse. 

Skull slightly convex dorsally in profile. Highest Skull flattened dorsally in profile. Highest 

point of cranium behind supraorbital space. point of cranium in the occipital region. 

Postorbital processes with anterior and Postorbital proces with only a posterior 

posterior angular processes. Anterior process angular process. Anterior processes completely 

sometimes poorly developed. fused with frontals. 

Rostrum long. Nasals measured along oblique Rostrum very short. Nasals measured along 

line about as long as frontal suture. oblique line markedly shorter than frontal suture. 

Incisive foramina about as long as bony palate. Incisive foramina much narrower than length 

of the bony palate. 

Interorbital space posterior to postorbital Interorbital space almost as wide as both 

process about 60.4 % of the width of both nasals in nasals in their posterior third, 
their posterior third. 

Pentalagus and Caprolagus have the following characters in 
267 common: 1) ears very short; 2) fur coarse; 3) bony palate primitive, 

i. e. , very long; 4) incisive foramina narrow; 5) nasals shorter. The skull 
of Pentalagus is not as flat as that of Caprolagus. Its dental formula is 

2 3 

m. — instead ofm. — . Pentalagus is uniformly colored, dull black - 

brown both dorsally and ventrally. Its claws are unusually large especially 
on the forelegs. Pentalagus is, in our opinion, a well characterized 
genus. 

We place the following species in the genus AUolagus: AUolagus 
mandshuricus Radde, A.brachyurus brachyurus Temm. (1845) 
from Japan (Kyushu, Shikoku — southern part of Hono); A. b. okiensis 
Thos, (1906) from Dogo Island; A. b. etigo Abe (1918) — northeastern 
part of Hondo Island; and doubtfully A . b . angu s t id ens Hollister (1912) 
described from one specimen fron Hondo (Tateyama). 

J.Abe ("A Synopsis of the Leporine Mammals of Japan" (J. of Sci. of 
Hirosh. Univ.", Vol.1, p. 54-55, 1931)), attempted to analyze the 
characters of AUolagus mandshuricus andLepus brachyurus. 
He came to the conclusion that L. brachyurus does not belong to the 
genus AUolagus, and that it is more closely related to Lepus than to 
AUolagus. It is closely related to the blue hare in his opinion. 

We completely disagree with this. J.Abe (1931) compares AUolagus 
mandshuricus and b r a chy u r u s *, and concludes that there are great 
differences between the two species. 

As described in "Zoologischer Anzeiger", 192 9, the most characteristic 
feature of AUolagus mandshuricus is that the nasals are about 
twice as wide in the middle as the interorbital space posterior to the 
postorbital processes. 

After a new study of skulls I came to the conclusion that this was not 
exact. It should be formulated as follows: 

1) Interorbital space posterior to postorbital processes about 60.4% as 
wide as both nasals in their posterior third. 

We then obtain the following measurements (in mm): 



* These data appear in the "Zoologischer Anzeiger". 



233 



Allolagus A. brachyurus 
mandshuricus 

Width of nasals in the middle 14.5 - 18 16-19* 

Width of interorbital space 10.2 - 13.2 11-13 

Width of nasals in the posterior third 16.3—19.2 — 

2) Nasals about as long as frontal suture or little shorter: 

Length of nasals 32.8 - 37.9 36 - 43 

Length of frontals 33 - 41 35.5 - 41 

269 3) Width of posterior nasal foramina about equal to length of bony palate. 
J. Abe and I obtained different measurements of this foramen because 
J.Abe measured the anterior nasal foramina and I measured the posterior 
foramina. If we compare m and s hu ri cu s and b r ac hy u r u s according 
to J.Abe's measurements, there is no difference between the two species: 

Allolagus , , , 

° A. brachyurus 

mandshuricus 

Width of posterior nasal foramina 11 —12.8 12 — 14 

Length of bony palate 7. 3 — 9. 2 7. 2 — 9 

4) Maximal width of incisive foramina about equal to length of bony palate: 

Width of incisive foramina 7.2—9.9 8. 1 — 1 1 

Length of bony palate 7.3-9.2 7.2-9 

5) Incisive foramina about as long as upper diastema: 

Length of incisive foramina 19 — 21.8 20.5 — 22.5 

Length of upper diastema 20.2—23 21 — 25 

This comparison shows that there are no significant differences between 
the skull of A . mandshu ricus and A . b r a c hy u r u s in these characters. 

The taxonomic position of the black Manchurian hare. Only a few data 
on this form have been published. N. M. Przheval'skii mentions the black 
Ussuri hare in "A Journey to the Ussuri Territory" (Puteshestvie v 
Ussuriiskom krae), Sankt-Peterburg, p. 258, 1870. Th. Noack ("Wild und 
Wald in Ost-Sibirien", Der Weidmann, Bd. XXII, No, 48, S.420, 1891) 
recorded C aprol a gu s brachyurus niger Noack, from the lower 
reaches of the Amur. He did not give a full description, but mentioned only 
briefly that a small hare resembling the Japanese Caprolagus 
brachyurus Temm. occurs in this region, but it is black. 

In my article "Material on the Systematics of Russian Hares" ((Materialy 
po sistematike russkikh zaitsev) Ezhegodnik Zoologicheskogo muzeya 
Akademii Nauk, Vol. XXIII, p. 489, 1922) I named the black Manchurian 
hare provisionally Lepu s mandshuricus subphasa m e lano nt u s . 
I was induced to do this by an interpretation of lower systematic 
categories given by V. L. Bianki in "The Species and Infraspecific Taxa", 
(("Vid ipodchinennye emu taksonomicheskie formy") Russkii zoologicheskii 
Zhurnal, I (9-10):292. 1916.) 

It does not seem justified at present to give the black-backed Manchurian hare 
a specific name. It represents an aberration with rather constant color 
characters of partial melanism. This form seems to acquire a 
geographical range. It apparently occurs more frequently in southern 
Manchuria than in the Ussuri Territory. 

This is apparently analogous to the case of the black hamster 
(Cricetus cricetus) which also predominates (or occurs frequently) 



* According to J. Abe. 

234 



268 







FIGURES 140-143. Skull of C aprol agus hispidus Pearson 



235 



only in some geographical regions. A geographical natio may possibly 
develop from an aberration. 
271 Together with the black -backed melanic form there occur also very rare, 
completely melanic specimens. 

GEOGRAPHICAL DISTRIBUTION. L.mandshuricus occurs in the 
north as far as the Bureya Mountains (to about 50°N). It occurs 
throughout the Ussuri Territory to the Korean border. It is apparently 
widely distributed in northern Manchuria, e. g., in the Kirin Province 
(A. Sowerby, 1923). It seems to occur at least in North Korea. According 
to A.I.Cherskii (in litt. 1914) the Manchurian hare occurs frequently near 
Vladivostok. V. V. Bogomolov (in litt. ) often found this species in the 
Sikhote-Alin Mountains. According to N. T. Zolotarev (1936), the following 
specimens are in the ZMAN collection: from the Bureya Range (collected 
by Radde); two specimens from near Khabarovsk (presented by M. A. 
Menzbir); one specimen from Pokrovskaya railroad station (presented by 
Bykov); one specimen from Chernigovka (Emel'yanov); two specimens 
from Sidemi (Yankovskii). 

Accordingto G. D. Dul'keit (1938, in litt. )the northernboundary of the distri- 
bution of the Manchurian hare follows the bank of the Kop' river*, continues 
along the Amur river near Verkhnee Tambovskoe, Srednee Tambovskoe. 
In the west it follows the Bureya Range, Little Khinganand Great Khingan. Its 
range also includes the basins of the Sungari and Tumen-Ula rivers. 

MODE OF LIFE. We know very little about its biology. 

A.I.Cherskii (1914) gave the following data: "The habitat of 
L.mandshuricus is southern slopes of low mountains, i.e., ridges 
overgrown with forests of oaks, elms, lindens, and various species of 
maple with shrub undergrowth. One always sees tracks, and wide beaten 
tracks in such places. " 

According to N. M. Przheval'skii (1870) this species mainly inhabits 
islands in the Ussuri river which are covered with osier, and forests in 
mountain valleys. It never approaches human settlements, even in winter. 

G.I. Radde (1862) gives the following data on this species in the Bureya 
Mountains, "It is remarkable that this species, unlike related species, does 
not have open lairs. It settles in holes in tree trunks which are open on 
one side. I found beaten tracks to such a cavity. This animal is very shy 
during the day. It leaves its shelter when man is still far away. It 
prefers dense underbrush and osier beds. Its course is as straight as that 
of the Tolai hare. Its meat is very savory. " 

Genus Oryctolagus Lilljeborg (1873) 

1790. Cuniculus Meyer F. A. , Magaz. fiir Thiergeschichte, I, S. 52-53. 

1873**. Oryctolagus Lilljeborg, Sveriges og Norges Ryggradsdjur, I, p. 417 (as a subgenijs of the genus 
Le pus '. ). 

Ear measured from occiput shorter than head. 

Dorsal side of tail black, its color sharply contrasting with that of the 
ventral side. 

Hind legs short. When the animal stands, they seem little longer than 
the forelegs. 



272 



G. D. Dul'keit found only one blue hare north of the valley of the Kop' river. 

The year 1874 appears on the title page of the book. However, it was published in issues, and the issue 
with the description of Oryctolagus was published in 1873 (T. S. Palmer, Index generum Mammalium, 
p. 484, 1904). 

236 



270 





FIGURES 144-147. Allolagus brachyurus Temm. No. M.4518. Female ad. , 27 January 1935, 
Nagoya, Hondo, Japan, N. Kuroda, S. I. Ognev's collection 

Drawing by Yu. A. Kostylev 



237 



273 




FIGURES 148-151. Skull of Oryctol agus cuniculus L. Specimen No. S.3319, from 
near Kalish, MSZM* collection 

Drawing by Yu. A. Kostylev 



* [MSZM — Moscow State Zoological Museum. ] 



238 



Mesopterygoid fossa much narrower than the length of the bony palate. 

Interparietal well developed in adult and old specimens. Postorbital 
pr-ocesses narrow, thin, swordlike, not triangular as in many Russian hares. 

Ratio of width of interorbital space measured behind the postorbital 
processes to width of nasals measured in their posterior third about 81%. 
This is a sharp difference between the skull of Oryctolagus and 
Allolagus''=. 

Nasals longer than frontal suture, frontals measured to the anterior 
process between the nasals. 

Width of posterior nasal foramen less than length of bony palate. 
Anterior nasal foramen about as wide as the length of the bony palate. 

Incisive foramina in their posterior part about as wide or slightly less 
than the length of the bony palate. 

Incisive foramina about as long as, or slightly shorter than, upper 
diastema. 

Type species: Lepus cuniculus Linn. 

GEOGRAPHICAL DISTRIBUTION. North Africa and the southern and 
central parts of western Europe. Since about 1850 the rabbit was introduced 
and multiplied in many countries of the warm region in both hemispheres, 
e.g., Australia, South Africa, New Zealand and islands of the Pacific 
and Atlantic Oceans. 

162. Oryctolagus cuniculus Linn. (1758). The rabbit 

1758. (Lepus) cuniculus Linnaeus C. , Systema Naturae, VoL 1, ed. 10, p. 58. 

1837. Lepus vernicularis Thompson I. , The Athenaeum, p. 468 (nomen nudum.' ). 

1843. Lepus vermicula Gray I. , List Spec. Mammal. British Museum, p. 128 (nomen nudum'. ). 

1857, Lepus cuniculus Blasius L , Naturgesch. , S. 426-428. 

1867. Cuniculus fodiens Gray I. , Annals and Magazine of Natural History, ser. 3, Vol. XX, p. 225 

(instead of cuniculus'. ). 
1904. Oryctolagus cuniculus Lyon M. W. , Classification of the Hares and their Allies. Smiths. 

Miscell. Collect., VoL XLV, 1, part. 3-4, pp. 402-406. 

NAMES: The Rabbit or Cony (English); Das Kaninchen (German); Le Lievre lapin ou le Lapin (French). 

TYPE LOCALITY AND DEPOSITION. C. Linne's "Systema Naturae" (1758) states: "Habitat in Europa 
australi". G. S. Miller (1912) tries to make this more accurate, and considers southern Germany as type 
274 locality. The type is apparently lost. 

Principal figures: 1) Schreber I. Ch. , Die Saugethiere, Erlang. , IV, 1792, S. 891-897, Taf. CCXXXVI, 
A (entire animal in color); 2) Bingley W. , Memoirs of British Quadrupeds, 1809, p. 309 (color, pi. ); 

3) Blasius I. , Naturgesch. der Saugethiere Deutschl. 1857, S. 426-428, Fig. 230; S. 426 (structure of teeth); 

4) Brehm A. , Zhizn' zhivotnikh (Animal Life), II, 1874, p. 257 (half tone plate); 5) Trouessart E. L. , 
Histoire Naturelle de la France, Mammiferes, Paris 1885, p. 189, fig. 79 (entire animal); p. 182, fig. 17 
(skull); 6) Fokht K. , MIekopitayushchie (Mammals), Sankt Petersburg 1885, p. 404 (drawing of entire animal); 
7) Brehm A. E. , Zhizn' zhivotnikh (Animal Life), II, 1893, p. 671 (biological scene); 8) Tullberg Tycho, 
ijber das System der Nagethiere, Upsala 1899, Taf. I, Fig. 1-21 (skull, inner organs); 9) Beddart Fr. E. , 
Mammalia, London 1902, p. 503, fig. 246 (skull lateral and ventral); 10) Millais I. G. , The Mammals of 
Great Britain, Vol. Ill, 1906, pi. 52 (color), pi. 48 (biological scene), pi. 50 (photograph) ; 1 1) Meerwarth H, , 
Lebensbilder aus der Tierwelt, I, Leipzig 1908, S. 45, 47, 49-51, 53-57, 63, 67 (photograph of animals, 
nests and burrows); 12) Schaff E. , Die wildlebenden Saugethiere Deutschlands, Neudamm 1911, S. 122, 

Fig. 35 (skull ventral); 13) Barrett-Hamilton G. , A History of British Mammals, X, 1912, p. 173, fig. 32 
(skull); p. 174, fig. 33 (structure of interparietal bone); pi. XII (head of male and female rabbit); pi. XIII 
(same anteriorly); p. 203, fig. 35-37 (diagrams of burrows); pi. XIV (biological scene); p. 207, fig. 38 
(diagrams of burrows); p. 209, fig. 39 (same); fig. 40-42 (same); p. 220, fig. 43 (tracks) ; p. 230, fig. 44 
(skull lateral); fig. 45 (2) (skull); pi. XIX (I) (feet); pL XX (6) (feet); fig. 48, pi. XIII, p. 315 (4), 



In Caprolagus the ratio of width of the interorbital space to that of the nasals is about the same as in 

O rye t o 1 ag us 

239 



275 



(mandible); 14) Miller G. S. , Catalogue of the Mammals of Western Europe, London, 1912, p. 488, fig. 101 
(skull lateral, ventral and dorsal); 15) Thorbum Archibald, British Mammals, Vol.11, 1921, pi. 36 
(colored); 16) Brauner A. A. , Sel'skokhozyaistvennaya zoologiya (Agricultural Zoology), 1923, p. 79, 
fig. 76 (occipital foramen and structure of palate); 17) Shillinger F. F. , Zaitsy — rusak i krolik (Hares — 
The Common Hare and the Rabbit), 1929, p. 51, figs. 30, 31; p. 52, fig. 33; p. 54 (biological scene); 
18) Kuznetsov B. A. , Dikii krolik na Ukraine (The Ukrainian Wild Rabbit), "Pushnoe delo", No. 6, 1929, 
pp. 64-66, figs. 1-3 (diagrams of burrows); 19) Kuznetsov B. A. , K voprosu o khozyaistvennom znachenii 
dikogo krolika na Ukraine (The Economic Value of the Ukrainian Wild Rabbit), "Ukrains'kyi mislyvets ta 
ribalka" No. 7, 1929, p. 18, fig. (entire animal); 20) Heptner W. G. , Zaitsy (Hares), Moscow, 1933, p. 29, 
fig. 5 (entire animal); 21) Manteifel' P. A. , Zaitsy SSSR (Hares of the U. S. S. R. ), in "Biology of Hares and 
Squirrels" (Biologiya zaitsev i belok), Moscow, 1935, p. 26, fig. 1 (il-day old hare). 
MATERIAL EXAMINED: 15 pelts and skulls. 

DIAGNOSIS. Head relatively large. 

Length of outstretched ear measured from occiput less than maximal 
length of head. 

Tail with terminal hair about 94% of the length of hind foot without 
claws. 

In summer and winter fur, dorsal side of tail black, contrasting with 
white background of the other part of the tail. 

Hair not becoming white in winter, remaining brown. 

Fur soft. Guard hair soft and straight. Guard hair and hair of 
undercoat not curly. Teats 6: p. 1 - 1; a. 2 - 2. Total 6. 

Nasals measured along line connecting anterior inner corner to outer 
posterior corner longer than frontal suture. Postorbital process much 
longer than maximal width of nasals in their posterior part. Width of both 
nasals about 83% of length of postorbital process. 

Posterior third of incisive foramina with straight sides, widened, without 
protuberances. 

Ratio of length of bony palate (lateral to anterior process) to maximal 
width of incisive foramina 100-115%. 

Ratio of length of bony palate to maximal width of mesopterygoid fossa 
100-110%. 

Ratio of length of row of lower molars to that of lower diasteina about 
87%. Row of lower molars shorter than diastema. 

Anterior lower premolar strongly inclined backwards in profile as in 
the common hare. 

Mandibular condyle massive, strongly inclined backwards. 

Posterior-ventral plate of mandible (from corner of angular process to 
notch at beginning of mandibular horizontal process) weakly developed. 
Distance between corner of angular process to notch at beginning of 
mandibular horizontal process longer than distance between anterior 
alveolus of first premolar to tip of lower incisor (not because of the small 
size of the plate, but because of the short diastema). 

Bullae osseae large and inflated, larger than those of Allolagus , and 
smaller than in desert hares. See Figures 148-150. 

Length of skull 76-81 mm (78-82 according to G. S. Miller (1912)); 
zygomatic width 36-39 mm (37.2-41.4 according to G.S. Miller); length of 
nasals 33-37.6 mm (35.4-38.6 according to G.S. Miller); width of nasals 
11. 6-12. 6mm. * 



* In the middle. According to G.S. Miller, maximal width of nasals 14- 17.2 mm. 



240 



Length of body and head 362-375 mm (392-437 according to G. S. Miller, 
1912); hind feet without claws 81-96 mm (87-92 according to G. S. Miller); 
ear measured from occiput 60-72 mm (65-70 according to G. S. Miller); 
tail 52-69 mm (47-68 according to G.S. Miller). 

ADDITIONAL DESCRIPTION. The cranial characters of the rabbit have 
been sufficiently described to distinguish it clearly from related species. 

COLOR. In summer fur, color of back brownish gray with a slight rust 
tinge, long and numerous black guard hairs. 

Color of top of head generally like that of back but darker because of the 
greater admixture of blackish hair. Neck dull grayish rust colored with 
an ocher tinge (between cinnamon (pi. XXIX, Ridgway, 1912), wood- 
brown and fawn color (pi. XL)). 

A broad, light brown stripe across chest, (between avellaneous, pi. XL, 
and light pinkish cinnamon, pi. XXIX). Flanks, forelegs and two stripes 
from the flanks to the groin of the same color. Middle of throat and belly 
white . 
276 Color of anterior inner part of ears brownish gray, similar to that of 
top of head. Outer fringe of anterior inner margin of ears lighter, dull 
whitish gray. Inner part of ears gray. Posterior (outer) part of ears 
brownish gray (between drab and hair -brown, Ridgway, 1912). 

Black fringe at tip of ears little marked. 

Tail white, with contrasting black dorsal side. 

Winter fur more luxuriant than summer fur. Color hardly different 
from that of summer fur, but richer and more rust colored. 

SYSTEMATIC NOTES. Two species of rabbits are known: 
Oryctolagus cuniculus L. and O. crassicaudatus I. Geoffroy, 
with the following subspecies: O. cuniculus cuniculus L. (Germany), 
O.c.septentrionalis Franga (Northern Europe), O.c. 
meridionalis Franga (Portugal), O. c. huxleyi Haeckel (Porto-Santo, 
Madeira), O.c.habetensis Cabrera (Morocco), O.c.oreas Cabrera 
(Morocco), O . c . b r achyo tu s Trouessart (estuary of the Rhone), O. 
crassicaudatus crassicaudatus I. Geoffroy (Transvaal), O.c. 
curryi Thos. (Orange river), O. c. nyikae Thos. (Niassa, Nyika 
Plateau). 

GEOGRAPfflCAL DISTRIBUTION. The wild rabbit was introduced in 
European Russia, and spread and multiplied in its southwestern part. 

A. A. Brauner (1923), in "Agricultural Zoology" (Sel'skokhozyaistvennaya 
zoologiya) writes: "About 25 years ago gray-brown domestic rabbits were 
liberated in plantations near Odessa. They multiplied and spread along the 
Black Sea coast, especially in summer resorts such as Lanzheron, 
Bol'shoi Fontan, Srednii Fontan and Malyi Fontan, Lustdorf, etc. They 
settled in the cliffs of the Khadzhibei, Kyal'nitskii and Tiligul'skii limans, 
as well as in plantations along railways. From here they passed to groves, 
parks, and orchards near the places mentioned. They are now widely 
distributed between the Bug and Dniester, and also east of the Bug 
(Voznesensko -Ratsynskoe Forestry)." 

B. A. Kuznetsov (1929) writes that A. A. Brauner told him that the 
rabbit occurs in the Ukraine as far as the Kodyma river in the north (near 
Balta, 48°N). 

The rabbit apparently spreads eastwards. A. A. Migulin (1927) observed 
rabbits in the Nikolaev District north of Kherson. The Board of the Kherson 
Hunting Association informed B. A. Kuznetsov (1929) that wild rabbits occur 



241 



277 



in the Kherson, Kakhovka, Snigirevka and former Aleksandrovsk districts. 
At present Oryctolagus cuniculus is found in the right bank Ukraine 
between Dniester and Dnieper, reaching Aleksandrovsk in the east and the 
Kodyma river (Balta) in the north. 

History of acclimatization of the wild rabbit in the Ukraine. B. A. 
Kuznetsov (1930) gives some circumstantial data on this subject: "In the 
spring of 1929 I studied rabbit colonies near Odessa. Local hunters told me 
that the ancestors of these rabbits were some pairs of western European 
wild rabbits introduced at the beginning of the century by Graf Malokhovskii 
from Austria. They were set free on his estate near the Khadzibei estuary. . . 

"I received different information from the former manager of the 
hunting collective of the Kherson Hunting Association, Golubenko. He 
told me that the ancestors of the wild rabbits in the vicinity of Kherson 
were seven pairs imported from Switzerland in 1894 or 1895 by the 
landowner Pin'kovskii. He set* the animals free on his estate, 7 versts 
from Kherson. The Ukrainian rabbit colony possibly developed from 
several lots of animals introduced from western Europe by different 
landowners at various times." 

The native country of the rabbit is central Europe (reaching the 
Mediterranean in the south). The rabbit has been widely acclimatized by 
man throughout the world. However it could not become acclimatized in 
northern countries which are covered by snow in winter, and in which 
ecological conditions are unfavorable. According to H. Winge and G. 
Barrett -Hamilton, it did not become acclimatized in Denmark in spite of 
a number of attempts. The same happened in Norway. In northern Europe 
the rabbit is distributed as far as Ireland which has a mild maritime 
climate. 

The rabbit is common on the Azores, Madeira, the Canaries, Porto- 
Santo and on Selvagens. 

Some authors assumed that the rabbit was indigenous to some of these 
islands. Some of these rabbit colonies were introduced long ago by the 
Portuguese. For instance, they were introduced to Porto-Santo and 
Madeira in 1418 and 1419, respectively. The Porto-Santo and Madeira 
rabbits differ so sharply from the typical wild rabbit that Charles 
Darwin considered them as a new species formed by insular isolation. 
The island rabbits are certainly a well differentiated subspecies. 

The North African rabbits are probably not indigenous. According to 
F. Lataste the rabbit does not occur in Tunisia, Algeria or the Sahara 
but is common in the littoral islands. According to A. Cabrera, the 
rabbits in Morocco are not indigenous but are descended from animals 
which were acclimatized long ago. Rabbits also occur in the Chafarinas 
Islands, in the north Moroccan littoral. 

Rabbits were introduced in South Africa (where a separate species 
developed), inNew Zealand and Australia. Several pairs were introduced 
in Victoria, Australia (1859). They multiplied enormously. Large 
colonies occupied the pastures of sheep (which constitute the main source 
of income of the farmers), and destroyed the grass. This enormous 
multiplication of rabbits became a calamity. Statistics of the Department 
of Agriculture show that at the beginning of this century there were about 
20 million rabbits in Australia. The Australian government expends large 
sums for the control of this pest. 



242 



278 MODE OF LIFE. B. A. Kuznetsov gives descriptions of habitats 
favored by wild rabbits near Odessa, Kherson and other towns where 
rabbits survived, became acclimatized and multiplied. In southwestern 
Russia rabbits live in various habitats. Their typical habitat are large, low 
cliffs of loose shell-limestone layers. For instance, large piles of 
broken stone are scattered near the Arkadia summer resort near Odessa, 
after exploitation of limestone deposits (Kuznetsov' s data applied to 1929), 
These stone piles are used as shelters by the rabbits which live in holes and 
cracks among the stones. 

Rabbits occasionally inhabit cemeteries, where they burrow under the 
tombstones, for instance near Odessa and Kherson. 

Rabbits also settle in orchards on cliffs near the seashore, e.g. , on 
Bol'shoi Fontan in Odessa. 

Rabbits are especially numerous near the Luzanovka Health Resort on 
the sandy slopes of the Zhivakhova Mountain, overgrown by stunted grass. 
The animals live m.ainly among the stones of a crumbled wall. Near 
Berezovka, rabbits occur along ravines of brooks, tributaries of the 
Tiligul river and in forests. According to A. A. Brauner, rabbits inhabit 
cliffs at the seashore, banks of estuaries, near railway embankments, 
ravines, and in steppe forestries. The animals occasionally take shelter 
in the straw fences around farms. 

The forester in the Berezovskii forestry informed B. A. Kuznetsov that 
rabbits live under large heaps of reeds on his farm. 

Rabbits live under special conditions in the outskirts of Nikolaev. B. A. 
Kuznetsov found a large colony in the yards of the plant imeni Marti. 
B.A. Kuznetsov writes: "These large yards are between the plant 
and the bank of the Bug. The upper part of these yards consists of sterile 
desert covered by coarse alluvial sand with stunted thickets of grass or 
weeds. 

"In the lower part of the yard the vegetation becomes denser and more 
varied, passing into bulrush and sedge thickets of the coastal belt. 

"The whole yard, and especially mounds near the buildings, are 
covered with scattered heaps of iron, broken machines, heaps of slag, 
planks, etc. Numerous colonies of wild rabbits find shelter under these 
heaps of iron and slag. Burrows of these animals are frequent. The 
burrows pass under blocks of slag of many tons or between parts of broken 
machines. Burrows are never found on free ground. The sandy ground of 
the free spaces is covered by a network of tracks. Everywhere there are 
shallow pits where the rabbits urinate. Some places are completely covered 
by their feces, as rabbits always defecate in a definite place. 

279 "In spite of the proximity of the rattling plant, rabbits are very common 
in its yards. . . " 

A peculiar rabbit colony exists in caves and old mines of the Verevoch- 
naya Ravine near Kherson. B.A. Kuznetsov writes: "The Verevochnaya 
Ravine consists of two approximately equal parts. The upper part is 
narrower and its slopes steeper. The lower part is wider and its slopes 
are less steep. Strata of loose shell-limestone outcrop in the upper part 
of the ravine, especially on the left slope. These were formerly quarried, 
and are therefore full of caves and debris, of various sizes. Some of the 
caves are so large that a man can walk upright in them. These caves 
(which are called "skel'") are inhabited by large rabbit colonies which dig 
their burrows in cracks in the walls. Limestone strata do not outcrop in 



243 



the lower half of the ravine. The stone quarries therefore consisted of a 
labyrinth of underground passages connected with the surface by vertical 
shafts. Many of these underground passages have collapsed, creating 
deep pits. Shafts into the underground passages open at the bottom of these 
pits. In these shafts the rabbits are even more numerous than in the 
"skel's" in the upper part of the ravine. " 

The rabbit thus does not avoid the neighborhood of man. Owing to its 
secretive mode of life, its alertness and caution, it knows how to evade 
danger. 

The rabbit is a completely nocturnal animal. According to B.A. 
Kuznetsov's observations in the Berezovskii Forestry the rabbits emerge 
from their burrow at about 11.00 p.m. in spring and feed until sunrise 
(5-6.00 a. m. ). According to hunters, the rabbits are active in winter from 
midnight to daybreak. Where the rabbits are little disturbed, the young 
occasionally emerge from their burrow during the day to lie before the 
entrance and bask in the sun. However, rabbits never go far from their 
burrow. B.A. Kuznetsov writes: "When rabbits settle in orchards, 
ravines or cliffs on the seashore they usually live in colonies and their 
burrows are concentrated in a small area. Rabbit colonies are especially 
numerous in heaps of stones, in stone quarries or under buildings. 
Rabbit burrows are evenly distributed throughout the territory of the 
Berezovskii Forestry. 

"Near Odessa rabbits dig their burrows in crevices in quarries, under 
heaps of stone, and under buildings. I was therefore unable to excavate a 
single burrow in this area. On the other hand I excavated a number of 
burrows in the Berezovskii Forestry. These burrows are of two types 
Those of the first type are very simple. They consist of a rounded, widened 
central chamber, 30-60 cm below the surface, which is connected with 
the surface by one, generally two (rarely three) galleries 1-2 m long. The 
size of the central chamber varies. It is usually 40-60 cm wide and 25- 
40 cm high. 

"Most burrows of this type did not contain any bedding. Only in two 
chambers did I find nests of dry grass and moss. The exits usually open 
directly onto the surface, and are not at the bottom of broad, funnel-shaped 
pits as are those of the second type. The mounds of earth outside the 
exits are very small. 

"The burrows of the second type are more complicated. They have 4-8 
or more exits. Some of them are usually caved in and blocked with soil. 
Their exits open at the bottom of a deep and broad, funnel-shaped pit which 
has formed as a result of the repeated collapse of the opening. This proves 
that these burrows have been used by many generations of rabbits. 

"The existence of these two different types of burrows can probably be 
explained by the fact that burrows of the first type belong to single animals 
(females and young) while burrows of the second type are used by females 
to raise their young. This can be decided only by excavating a large number 
of burrows during the summer. In the Berezovskii Forestry rabbits are 
often absent from their burrows during the day, hiding under grass or tree 
trunks where they make "lairs". These consist of small oblong hollows 
with bedding of dry grass and leaves. They are usually well covered by 
overhanging branches. The rabbits lie very quietly sometimes letting a 
man approach to 3-4 steps. " 



244 



Numerous fecal pellets are usually found near the burrows on small 
nmounds, for example, on molehills. Such "latrines" are also found in 
the feeding plots. 

The food of the rabbit in summer differs sharply from that in winter. 
From spring to late autumn rabbits feed almost exclusively on green grass. 
When they live near a vegetable garden they feed on cabbages, carrots, 
young shoots of cucumbers, melons and watermelons. Rabbits apparently 
do not touch grain. 

In winter they feed mainly on dried weeds and roots of plants. Shallow 
funnel-shaped hollows on weedy grass plots are often found in winter. 
These are made by rabbits who dig roots from the ground. 

In forest nurseries rabbits feed on bark and young branches of trees. 
Injuries to bark of trees made by rabbits are easily distinguished from those 
made by hares. The teeth marks in the bark are always smaller than those 
of the common hare. Only the tips of branches are bitten off by rabbits. 

According to German authors (e.g., the new edition of Brehm revised 
by Ludwig Heck), the gestation of the rabbit lasts 30 days. This corresponds 
with data obtained in the Moscow Zoological Garden (P. A, Manteifel', "Hares 
of the U.S.S.R. " (Zaitsy SSSR), 1935, p. 26). Accordingto A. A. Brauner, 
the rabbit in southwestern Russia has 4 or 5 litters yearly, and each litter 
consists of 4-8* young. According to B. A. Kuznetsov (1929), each litter 
281 consists of 8 young. It is not known whether copulation takes place in the 
burrow or outside. The young are born naked, blind and completely 
helpless. Compared with newborn hares they look like embryos. Birth 
is followed immediately by copulation. This results in more intensive 
reproduction than that of the hare. If all 8 young of each of the 7 litters 
per year were to survive and reproduce, the population of rabbits would 
reach 1,274,840 in four years. This of course never happens in nature, as 
many are killed by enemies. 

The young become sexually mature in their fifth to eighth month, 
according to the locality. In hot climates they become mature earlier and 
in the temperate zone later. 

Rabbits do not hibernate and are active at night. 

They cause much damage to vegetable gardens and melon fields where 
they are numerous. 

In forests the rabbits gnaw young trees and shrubs. The forester of the 
Berezovskii Forestry told B. A. Kuznetsov that the rabbit is a much more 
dangerous enemy of the forest than is the hare. Rabbits are especially 
harmful in tree nurseries in which they may destroy large plantations of 
poplar, ash, locust trees, maple and fruit trees. 

The rabbit has many enemies. B.A. Kuznetsov (1929) explains the 
reduction of rabbits in southwestern Russia as follows: l) intensive 
hunting; 2) exploitation of quarries (stone heaps are a favorite habitat of 
rabbits), 3) multiplication of enemies of the rabbit, e.g., the fox. In addition to 
foxes the rabbit ishunted by Marte s foina, Mustelaputorius 
eversmanni Lesson, domestic cats and dogs. Young rabbits are 
frequently caught by buzzards and sometimes by eagles. B.A. Kuznetsov 
considers the fox the greatest enemy of the rabbit near Odessa. 



* In the last edition of Brehm, Heck gives the figure as 4-12. 



245 



282 



The pelts of the wild rabbit are beautiful and are in considerable 
demand in Russia and abroad. The rabbit has little commercial value 
because of its narrow range in the U.S.S.R. and its small population. 

The rabbits in southwest Russia should be protected: it should be 
prohibited to hunt them in summer. The damage caused by rabbits can be 
controlled, and it is inconceivable that we should exterminate them, as 
some people have requested. 



SECOND SUBORDER . SIMPLICIDENTA TA 

Maxillae with only one pair of large, curved, rootless incisors covered 
with a thick layer of enamel only anteriorly. Number of premolars 

i or i ; molars i to -. 

11 2 3 

The molars of some species have roots. In other species they have no 
roots and grow throughout life. The molars are either brachydont or 
hypsodont. The crowns are either bunoid or lophodont. The distance 
between the two upper rows of molars is smaller than that between the 
lower rows. Simplicidentata grind food by longitudinal movements of the 
rows of molars; lateral movements also take place. The glenoid cavities 
of Simplicidentata are narrow. As a result the mandible can slide along 
its axis. The two halves of the mandibles are usually loosely connected by 
the transverse mandibular muscle. The optic foramina are separate. The 
size of the infraorbital foramina is variable. They are large in most 
forms. When the central part of the masseter passes through the 
infraorbital foramen, its diameter may be very large, almost as large as 
that of the orbit (e.g., in Hystricidae). 

Bony palate complete, consisting of premaxillaries, maxillaries and 
palatines. Palate with incisive (or palatine) foramina. 

Soft palate with only a few folds. Cecum without spiral valve. 

Os penis well developed ih most forms but sometimes very small. 

The extinct family Ischyromyidae is considered intermediate 
between the Duplicidentata and the Simplicidentata. They lived from the 
Lower Eocene to the Oligocene, 



KEY TO THE FAMIUES OF THE SUBORDER SIMPUCIDENTATA 

1 (2). Broad fur-covered fold of skin (parachute) between fore- and hind legs which is supported by 
a thin bony process as the base of the wrist. The parachute is used for gliding and for leaping from one tree 
to another. 

Pteromyidae 

2 (1). Parachute between fore- and hind legs absent. 

3 (4). Back and flanks covered with long, hard spines. 

Hystricidae 



4 (3). The whole body covsred with more or less soft hair. 

5 (8). Toes of hind legs connected by a bare broad web. Tail squamous. 



1254 



246 



6 (7). Tail very broad, flat. Anterior upper molar about twice as wide as posterior molars. Skull 
without postorbital processes. 

Castoridae 

7 (6). Tail round in transverse section. Posterior upper molar almost twice as wide as anterior 
molar. Postorbital processes present. 

Capromyidae 

8 (5). Webs between digits of hind legs absent. Tail more or less densely covered with hair. 

5 4 
283 9 (12). Molars (including premolars) 7 or — . 

5 4 

10(11). Molars (including premolars) usually — , onlyinTer.es (Caucasian-Persian Squirrel) — . 

Tail always without homy scales, shaggy, often depressed doisoventrally. Tail of arboreal forms with 
long hair. 

Infraorbital foramen small, rounded or rounded-oval. Tubercles on crowns of molars form elevated 
transverse enamel folds. On intact molars these tubercles are well marked, especially in the marginal parts. 

Sciuridae 

(Squirrels, chipmunks, susliks, and marmots) 

4 

11 (10). Molars (including premolars) always — . 

Tail without homy scales in the majority of species and densely covered with hair. The tail of the 
genus Myomimus is covered with scant, very short hair through which the scales may be visible. 
Infraorbital foramina narrow, laterally compressed, elongate vertically. Crowns of molars with numerous 
transverse enamel folds, without well marked tubercles. 

Myoxidae 

4 

12 (9). Molars including premolars less than — . 

4 

13 (16). Molars — ; first molar very small. 

14 (15). Hind legs not more than 2-21/2 times as long as forelegs. Hind feet with 5 normally developed toes. 
Only the first, inner toe of the forefeet shortened, its tip hardly reaching the base of the next toe. Metatarsals 
of hind legs free, not fused. Tail squamous, covered with scant hair, very thin and long, without terminal 
tuft, prehensile, capable of coiling around branches. 

Subfamily Sminthinae 

15 (14). Hind legs about 4 times as long as forelegs. Hind foot with 2 shortened lateral toes, the tips 
of which do not reach the bases of the 3 middle toes, or with only three toes. Three central metatarsals 
fused. Tail without scales, densely covered with short hail, with a well developed tapering terminal tuft 
("flag"). Tail not prehensile. 



Dipodidae 



3 
16 (13). Molars - , 



3 

17(18). Molars very small, rudimentary; length of upper molar row about 1.5 mm. Molars covered by 
gums, not taking part in chewing. Food is cut by the enormous, sharp, curved incisors. Anterior surface 
of incisors with a deep longitudinal groove. 

Seleviniidae* 

18 (17). Molars normally developed and protruding from the gums. 
„Q . 19 (20). Hind feet with two short lateral toes the tips of which do not reach the basis of the three 

central toes, or with only three toes. Tail long, with a terminal tuft (15) ("flag"). 

Dipodidae 



* The vernacular name of this family is derived from the "bayalych" plant (Salsola arbuscula). 
Species of this family are found in thickets of this plant in the Bet-Pak-Dala desert. 

247 



20 (19). Hind feet with five normally developed toes; tips of lateral toes always reaching the basis of 
the three central toes. 

21 (22). Eyes rudimentary, covered with skin. No external ears. Tail absent, not visible externally. 

Spalacidae 

22 (23). Eyes developed but very small in some species (e. g. , in Ellobius). External ears present, 
sometimes small and reduced to folds of skin, forming elevated ridges along the margin of the auditory 
meatus. Tail present, but very short in some species, e.g., Ellobius. 

23 (24). Clavv'S of forefeet very large, thick and curved, particularly those of the three central toes. 
Claws longer than toes, 

Myospalacidae 

24 (25). Claws of forefeet normal, sometimes slightly lengthened. 
Muridae 

Family PTEROMYIDAE, FLYING SQUIRRELS 

1855. Pteromyinae Brandt I. F. , Untersuch. iiber die Craniologisch. Entwicklungstufen und Classification 
der Nager. Memoires mathem. , physiques et naturelles, tome VII, St-Petersburg, p. 151. 

1879. Pteromyidae Anderson, Anatom. and Zoolog. Researches, Expedition of Junnan, p. 278. 

1904. Pteromyinae Trouessart E. L. , Catalogus Mammalium, Supplement, p. 297. 

1912. Pe t auris t i dae Miller G.S., Catalogue of Mammals of Western Europe, p. 940; Pocock R. , On 
the Classification of Sciuridae, Proceed. Zoolog. Soc. of London, 1923, p. 240. 

1930. Pteromyidae Vinogradov B. S. , Mlekopitayushchie SSSR. Gryzuny (Mammals of the U. S. S. R. , 
Rodents), Izdatel'stvo Akademii Nauk, p. 22. 

These animals resemble squirrels in general. The head is rounded and 
the muzzle blunt. The ears are protruding and well developed, the tail is 
covered with long hair. The typical character of flying squirrels is the 
hairy parachuting membrane. This consists of a fold of skin between the 
fore- and hind legs. At the outside of the forefeet there is a bone directed 
laterally and posteriorly, which is as long as the antebrachium in Russian 
flying squirrels. These bones support the parachute membrane. They 
are very short inEupetaurus. 

A small antebrachial fold of skin in front of the forelegs. A more or less 
developed interfemoral fold between the hind legs, which is strongly 
reduced in Eupetaurus . 

Dental formula: i-; c— ; p— ; m- = (22). 

285 GEOGRAPHICAL DISTRIBUTION. Forest zone of the northern 

hemisphere. India, Ceylon, Burma, South China, Sumatra, Java and the 
southern parts of the U.S.A. =■' In the north, the range includes a wide area 
from Scandinavia to Chukchi (Anadyr Basin). In North America the northern 
boundary of the range of the flying squirrel runs from the Yukon Basin and 
the lower reaches of the Mackenzie river to the Great Bear Lake and further 
south to the Great Slave Lake. The boundary continues eastwards along the 
southern part of Mackenzie State, turning south through northern Manitoba, 
Ontario through Quebec, endingnorthof Cape Harrison. The northern 
boundary of the range of the flying squirrel in Alaska and Canada turns 
sharply south from west to east. 

The family Pteromyidae consists of eleven genera some of which may be 
regarded as subgenera. The genus Eupetaurus Thos. should be more 



It reaches the northern part of southern California and Utah in the west, and southeastern Texas and Florida 
in the east. 

248 



correctly considered as a subfamily, Eupetaurinae, separate from the 
subfamily Pteromyinae. O. Thomas * showed that the Eupetaurinae 
should be considered as a subfamily because of their different skull 
structure and hypsodont teeth with a peculiar form of crown. 

The subfamily Pteromyinae includes the following genera--: l) 
JomysThos. (type species J . h o r s fi e 1 d i Waterh. )- Java, Borneo, 
Malacca; 2)Belomys Thos. (type species B.pearsoni Gray) - 
Himalayas, Sikkim, Yunnan, Assam, Formosa; 3) P t e r o m y s c u s Thos. 
(type species P. pul ve r u le nt us Gunth. ) - Malacca, Borneo; 4) 
Pet aurillus Thos. (type species P . h o s e i Thos. )- Borneo; 5) 
Pteromys G. Cuvier (type species P.volans L. )- distribution given 
below; 6) Glaucomys Thos.t (type species G. volans = Mus 
volans L.) - North America; 7) Hylopetes Thos. (H. everetti 
Thos. )- Nepal to the Malay Archipelago; 8)Petinomys Thos. (type 
species P . lugens Thos.) -India, Ceylon, Malay Archipelago; 9) 
Eoglaucomys Pocock tt (subgenus) (type species E . fimbriatus 
Gray) - Gilgit, Himalayas; 10) Trogopterus Heude (type species T. 
xanthipes A.M. Edw. ) - South China. 

Genus Pteromys G. Cuvier (1817) 

1800. Pteromys Cuvier G. , Lejons d'anatomie comparee, recuillies et publiles sous ses yeux, I, tab. I 

(Nomen nudum'. ) X 
1817. Pteromys Cuvier G. , Le Rfegne Animal, Vol. I (description of the genus); See also Editio II, 1829, 

p. 194. 
286 1822. Pteromys Flemming I. , Philosoph. Zoolog. , H. p. 190 (See G. S. Miller, Proceed. Biological 

Society of Washington, 1914, p. 216). 

1825. Sciuropterus Cuvier F. , Dents des Mammif eras, p. 255. 

1826. Pterom ys Cuvier F. , in: Despreaux L. , Dictionnaire des sciences naturelles, dans lequel on traite 
methodiquement des differents etres de la nature, XLIV, pp. 40-41. 

The animal resembles a squirrel in general but the body is flattened 
and a parachuting membrane extends between the legs. This will be 
described in more detail below. There is no parachuting membrane 
between the hind legs and the base of the tail. 

The skull has a large, rounded braincase. The rostrum is short. The 
sides of the rostrum are nearly parallel in dorsal view. The nasal 
processes of the premaxillaries are narrow anteriorly and widened 
posteriorly. The zygomatic arches are widened anteriorly. The zygomatic 
processes of the maxillaries are broad. Seen from the front they form 
plates which are slightly narrowed dorsally. Middle parts of zygomatic 
arches thickened dorsoventrally, forming narrow, horizontal plates. 



* O.Thomas, Eupetaurus, a new form of Flying squirrel from Kashmir Journ. of the Asiatic Society 
of Bengal, Vol. LVII, part II, No. 3, 1888, pp. 256-260. 
** O.Thomas, The Genera and Subgenera of the S ciuropte rus Group., Ann. and Magaz. of 
Natural History, ser. 8, Vol.1, No. 1, 1908, pp. 1-8. 

t This and the following genus are considered by O. Thomas (1908) as subgenera of Pteromys 
tt R. Pocock, Classification of the Sciuridae, Proceed. Zoolog. Soc. of London, part I, 1923, p. 241. 

X I could not obtain the first edition of "Lecons d'anatomie comparee. There is no description of the 
Flying squirrel in the second edition (t. II, p. 152, 1837). Cuvier writes as follows: "D'autres 
mammif^res, savoir, les gal^opiteques, les polatouches ou Icureuils volants, et les phalangers 
volants, ont des membranes entre les pattes, mais sans alongement des doigts; elles ne peuvent 
pas servir a les elever, mais elles les soutiennent asse^ bien en I'air pendant quelques temps de tres 
grands sauts en descendant aux quels on ne peut point donner le nom de vol", According to 
T. S. Palmer ("Index generum Mammalium", 1904, p. 595) the Flying squirrel is also a nomen 
nudum in Cuvier's Comparative Anatomy. A description of the genus appeared in "Regne Animal". 

249 



287 






FIGURE 152. Skull of flying squirrel, Pteromys volans L. , 18 November 1930, Shar'ya District, 
Gorki Territory, A. N. Formozov (specimen No. C) 

Drawing by Yu. A. Kostylev 



250 



Frontals shortened posteriorly, reaching the interorbital space by a little 
more than the length of the postorbital process. Anterior palatine 
foramina large. Mandible m.assive. Angular process very large. 

Molars of complex structure. The posterior enamel folds of the upper 
teeth (from the second premolar) have deep oblique -transverse grooves on 
the posterior side. 

The glans penis of the flying squirrel is very peculiar. It has a slender, 
fingerlike process which is slightly inclined downwards in profile. The 
glans bears an oblique row of blunt, horny denticles dorsally. 

The OS penis is 5.2 mm long. A large lateral denticle at the base of the 
OS penis. Apex of os penis directed downwards. 

The structure of the os penis in the American genus Glaucomys differs 
sharply from that ofPteromys; it has no sharp basal denticle and has 
a slightly elevated fringe at the side. Its tip is turned upwards. These 
differences are well shown in figures 153-156. 

There are also great differences in the skull and in the structure of the 
teeth between the palearctic Pteromys and the American Glaucomys*. 
The sides of the rostrum converge anteriorly in Glaucomys: the rostrum 
is thus markedly narrowed anteriorly. The nasal processes of the 
premaxillaries are slightly narrowed anteriorly. The zygomatic arches 
_„ are not widened anteriorly and are comparatively narrow. The zygomatic 
processes of the maxillaries are narrow. Seen from the front, they form 
plates which are narrower dorsally. The middle parts of the zygomatic 
arch are not dorsoventrally depressed. They are vertical, narrow plates. 
The frontals are lengthened posteriorly and reach beyond the orbital 
constriction by more than the length of the postorbital process. Anterior 
palatine foramina small and narrow. Mandible weak. Angular process 
relatively much smaller than in Pteromys . 

Structure of molar teeth simplified. Posterior enamel ridges of upper 
teeth (from second premolar to last molar) without grooves. 






FIGURE 153, Os penis of Pteromys volans L. 

1 — lateral; 2 — ventral 

From a specimen collected by I, S. Slesarev in the 
Central Reserve (former Belyi County, former 
Smolensk Province). Highly magnified. 

Drawing by Yu. A. Kostylev 



FIGURE 154. Glans penis of Pteromys 
volans L. 

1 — lateral; 2 — dorsal. 



* Comparison of material confirmed the observations of A. Howell (Revision of the American Flying 
Squirrels, North Amer. Fauna No. 44, 1918, p. 15, 



251 



Type species: Sciurus (Pteromys) volans Linnaeus. 
288 It is difficult to decide how many species belong to this genus. 

Pteromys biichneri Satun. from Kansu is in our opinion a good 
species. The position of Pt e r o m ys filchenerinae Matschie from 
Sining (southeast of Kuku-Nor) is not quite clear. It was described from 
only one badly preserved skin, without skull. 




FIGURE 155. 



Denticles on glans penis of Pte rom ys volans L. (lateral) 

Drawing by Yu. A. Kostylev 




FIGURE 156. American flying squirrel, Glaucomys volans L. , Os penis 
1 — ventral; 2 — lateral. 

Pteromys momonga Temm. from Japan is probably a subspecies 
of P. volans L. 

GEOGRAPHICAL DISTRIBUTION. Eurasia. The southern boundary of 
the range is shown on Map V. In the east Pteromys occurs in 
Manchuria, northern and central China, Korea and Japan. 



289 



163. Pteromys volans L, (1758). Common flying sqxiirrel 

1758. Sciurus volans Linnaeus C. , Systema Naturae, ed. X, p. 64. 

NAMES: Flying squirrel (English); Flatterhomchen, das fliegende EichhSmchen (German); Polatouche 
(French). 

LOCAL NAMES: Mysh' parikha, parusha, parikha ^Russian hunters); palyaur (Komi-Zyryan); kiznir 
(Permyak); pulo, or pulokh (Votyak); bobontol (Man'sy, Vogul); tulmneine (Khanty, or Ostyak along the 
Ob); ive-lanki (along the Irtyn); pailanglangi (along the Surgut); poeze (along the Narym and Ket'); 
sartyin (Bashkir) ; nyar-brod (Komi-Zyryan); maskal an (Yakut); netyagi, or detyagi (Yakut name of 
Russian origin); ummukki (Tungus); tumr, tumarsh (Nivkhi, Gilyak of the western coast of Sakhalin Island); 
alvilanga (Nivkhi, Sakhalin); Khongmo (Mangun, Nanai-Gold, Samagii); umugi (Udehe on the river Khor); 
omke (Nanai-Gold on river Goryun); omoki (Kil and Kuril); umki (Birar and Manyagr); notaga (Nani, 
Orochis); olbo (Mongolian) ; khilov (Buryat) ; tznugo (Ainu); tieke (Yukagir, Odul); ot'to-napral'gyn 
(Muravetlan, Chukchi); yambe-khull, umbekhull, fei-shu [flying rat] (Chinese). 

PRINCIPAL HGURES: 1) Schreber J. Ch. , Die Saugethiere, Erlangen, IV, 1792 S. 813-818, Taf. CCXXIII 
(entire animal in color); 2) Dwigubski I. , Opyt estestvennoi istorii vsekh zhivotnykh Rossiiskoi imperii 
(A Natural History of the Animals of the Russian Empire), 1829, pi. 19 (entire animal); 3) Simashko Yu. , 
Russkaya fauna. II (Russian Fauna. Vol. II) 1851, pp. 597-599, pL 133, fig. 1 (in color); 4) Blasius J, , 
Naturgeschichte der Saugethiere Deutschland's, 1857, S. 269-271, Fig. 152; S. 268 (half tone plate); 
5) Brehm A. , Zhizn' zhivotnikh. II (Animal Life), Vol. II, 1874, p. 64 (half tone plate); 6) Kobel't V. , 



252 



Geograficheskoe raspredelenie zhivotnikh (Geographical Distribution of Animals), 1903, p. 75 (half-tone 
plate of entire animal); 7) Miller G. S. , Catalogue of the Mammals of Western Europe, 1912; pp. 941-946, 
fig. 195; p. 943 (skull lateral, dorsal and ventral) ; p. 945, fig. 196(molars); 8) Meerwarth H. and Soffel K. , 
Lebensbilder aus der Tierwelt, III, Leipzig, 1912, S. 305, 307, 309 (photograph of animals); 9) Rajewski 
W. W. and Kaplanoff L. G. , Ubersicht der Sslugetiere der oberen Volga, Zoologischer Anzeiger Bd. 84, 
H. 5/6, 1929, S. 148-150, Fig. 1, S. 149 (photographs of winter and summer skins); 10) Serebrennikow M. K. , 
Ein Flatterhomchen aus dem Kirgisengebiet, Ztschr. fiir Sslugetiere, Bd. 4, H. 3, 1929, S. 142-143; 
Taf. XIII (photograph of entire animal); 11) Argyropulo A. , Opredelitel' gryzunov Ural'skoi oblasti (Key to 
Rodents of the Ural Region), 1931, p. 66, fig. 80 (dorsal view of animal); 12) Vinogradov B. S. and 
Obolenskii S. I, , Vrednye i poleznye v sel'skom khozyaistve mlekopitayushchie (Harmful and Useful Mammals 
in Agriculture), Moscow, 1932, p. 38, fig. 21 (drawing of entire animal); 13) Kuznetsov B. A. , Tovarovedenie 
pushno-mekhovogo syriya SSSR (Marketing of Undressed Pelts in the U. S. S. R. ), Moscow, 1932, p. 377, 
fig, Ti (photograph of dorsal side of pelt). 

MATERIAL EXAMINED: 211 specimens (112 specimens in the ZMAN collection; 75 specimens in my 
collection and from material put at my disposal; 20 specimens in the MSZM collection). 

DIAGNOSIS. Generally resembling a squirrel but its body is more 
flattened. Body covered with very soft, silky hair. Tail also more 
flattened than that of squirrel. Parachuting membrane between fore- and 
hind legs covered with dense hair. Parachuting membrane supported 
anteriorly by a thin, pointed, swordlike bone at the side of the base of the 
forefeet which is directed laterally and backwards^!'. When the parachute is 
stretched, it therefore protrudes markedly laterally. 

Head small, rounded. Muzzle blunt. Eyes black and relatively large. 
Tips of ears without long hair which forms tufts in the winter fur of 
squirrels. 

Ears resemble that of squirrel but are more convex anteriorly, their 
tips narrower and the outer dorsal margin has a broader notch. The ears 
of the flying squirrel are broader but shorter than those of squirrels 
(Figure 158). 

Tail without terminal hair, about 3/4 to 2/3 of the length of body and 
head. Hair on the tail much shorter than in squirrels. These hairs are 
soft, silky and luxuriant. 

Legs shorter and thicker than in squirrels. Forelegs almost bare in 
summer. Covered with soft, silky hair posteriorly and laterally only. 
Forefeet in winter densely covered with silky hair. Ventral sides of toes 
also covered with hair. Soles of hindfeetboth in summer and winter 
covered with dense, soft and silky hair. Bare calluses at the base of 
toes in summer. Ventral sides of toes also bare. In winter specimens 
calluses at base of toes of hind feet densely covered with soft hair. Ventral 
side of toes also covered with soft hair. Hind feet much larger than 
forefeet, 
291 Forefeet with three calluses at the base of the toes and two calluses in 
the posterior part of the palm. The inner callus is larger, the outer 
relatively small (Figure 159, left). Tips of toes with small tubercles 
which are smaller than in squirrels. Forefeet with 4 toes. Inner toe 
shortest, slightly shorter than the outer toe. The 2 middle toes are of 
about equal size, the inner slightly larger than the outer. Both middle 
toes much longer than the lateral toes. Claws of forefeet short. Claws 
of middle toes about 3 mm long (not measured along the curvature). 

Four rounded oval calluses at the base of toes in the hind feet. Tips 
of toes with large tubercles. Hind feet with 5 toes. Inner toe short, much 



290 



This bone is almost as long as the antebrachium. 

253 



shorter than outer toe, which is shorter than the 3 middle toes. The length 
of the 3 middle toes increases gradually from the inner to the outer toe. 
Length of claws of middle toes 2.5-3 mm. 




FIGURE 157. West Siberian flying squirrel 

Drawing by K. K. Flerov 





^\\\\\ \ 



FIGURE 158. Head of flying squirrel, Pteromys 
V o 1 a n s L. 

Drawing by A. N. Formozov 



FIGURE 159. Ventral side of feet of flying 
squirrel 

left — forefoot; right — hind foot. 

Drawing by A. N. Formozov 



254 



292 



293 




Belly white with a more or less marked straw or rust tinge. The white 
color of the belly mixed with gray in young specimens. Color of summer 

fur on back between straw-gray-yellow to 
blackish gray. Winter fur of pale specimens 
light silvery-gray, sometimes with a yellow- 
straw tinge. Color of back of more deeply 
colored subspecies dark ash-gray with a 
light straw tinge. 

Fur silkier and softer than that of 
squirrel. Margins of parachuting membrane 
with a well marked fringe of soft hair. Color 
of tail variable, but less so than in squirrels. 
In subspecies with dull-colored tail, the 
general tone of the dorsal side pale whitish 
gray. Subspecies with straw-rust-colored 
tail occur in East Siberia. The dorsal side 
of the tail of dark flying squirrels is dark 
brownish gray. 

Vibrissae in several groups. The main 
group consists of long black vibrissae on 
the muzzle in front of the eyes. They are up 
to 60 mm long and there are 12-14 hairs in 
each group. Above this group there are 
3-4 vibrissae on each side of the forehead. 

A few smaller vibrissae are present 
above the eye. One group near the external 
side of the eye and behind it. 

Rostrum very short. Nasals rounded 
anteriorly. They seem slightly elevated and terminally inflated in profile. 
Margins of rostrum parallel. 

Frontal processes of premaxillaries narrowedanteriorly and considerably 
widened posteriorly. 

The frontal processes of the premaxillaries in their posterior part are 
slightly wider than the nasals in the posterior third. 

Zygomatic process of the maxilla widened and projecting in dorsal view. 
Interorbital space narrow, its width 1/3-1/4 of the condylobasal length. 
Bullae osseae always longer than wide, strongly inflated, especially 
anteriorly. Auditory meatus large. Bullae laterally compressed in front 
of the auditory meatus. 

Upper tooth row much shorter than bullae. 

ADDITIONAL DESCRIPTION. Rostrum very short. Zygomatic arches 
rounded, markedly widened anteriorly by the zygomatic processes of the 
maxilla. 

Interorbital space narrow. Braincase large and relatively long. 
Distance between the narrowest place posterior to the postorbital processes 
and the highest point of the occiput much greater than that between the 
constriction and the tip of the nasals. This ratio is the opposite in squirrels. 

The anterior half of the nasals widened, rounded and slightly inflated. 
This is clearly seen in profile. Lateral margin of each nasal descending 
to the premaxillary in a rounded lobe. The anterior part of the nasal is 
therefore wedge-shaped. Premaxillary frontal processes much widened 
posteriorly, narrowed anteriorly. The anterior part of the frontals begins 



FIGURE 160. Hind foot of Sakhalin 
flying squirrel, Pteromys volans 
athene Thos, Post Koisakovskii 
[Korsakov], Sakhalin 

Photograph by S. I. Ognev 



255 



to taper at about the anterior margin of the maxillary zygomatic processes, 
i.e. , where these processes border the premaxillary frontal processes 
laterally and anteriorly. 



292 





FIGURE 161. Skull of Pteromys volans 
volans L. , 18 November 1930, Sharya District, 
northern Gorki Territory 

Photograph by S. I. Ognev 



FIGURE 162. Skull of Pteromys volans 
athene Thos. , Post Korsakovskii [Korsakov], 
Sakhalin Island 

Photograph by S. I. Ognev 



293 This constriction results in almost parallel margins of the rostrum. 

Zygomatic arches evenly rounded. Anterior part of zygomatic arches 
not quite as wide apart than the posterior part. Maxillary zygomatic 
processes markedly widened, making the anterior part of the zygomatic 
arches massive. -This is particularly marked if the skull is seen from 
the front. 

The frontal and sagittal sutures are almost straight or convex 
anteriorly. There is sometimes a backwards directed, angular 
protuberance lateral of these sutures. A similar protuberance is sometimes 
present in the middle of these sutures. 

Upper molar row about as long as diastema. 

Bullae osseae inflated, almost twice as long as wide, markedly widened 
and rounded anteriorly, compressed laterally in front of auditory meatus. 

Molars with transverse enamel folds alternating with grooves. Inner 
margin of each molar with a small, curved elevation. 

First upper premolar much larger than in squirrels. It generally looks 
like a small pillar near the anterior margin of the second premolar. Base 
of anterior premolar slightly inwards of the tooth row. Crown of pm 
simple, with one denticle. Inner side of surface of pm with curved or 
pectinate elevation. Together with the elevations of the other teeth, they 



256 



294 



form an almost continuous denticulate crest along the inner side of the 
tooth row. 

Outer side of anterior premolar with three elevations. The middle 
elevation larger than the lateral ones. Transverse enamel ridges 
corresponding to this elevation run to a small inner curved elevation on the 
inner side of the tooth. Posterior transverse enamel ridge broken by an 
oblique groove which divides it into two cusps. 

The first premolar is almost as large as the first upper molar which 
resembles the second premolar, but there is a depression on the second 
transverse enamel ridge at the part of the ridge joining the small inner 
curved enamel elevation at the inner side of tooth. Structure and size 
of second molar like that of first molar. Last molar smaller, narrower 
towards the inner margin. The first transverse enamel ridge of the last 
molar is less well developed. Second enamel ridge identical with that of 
the preceding tooth. A depression in the posterior part of the last enamel 
ridge (Figures 161-162). 

Structure of lower molars. Outer side of premolar with two denticles. 
Anterior denticle higher than posterior. Both denticles connected at the 
inner side by a small enamel bridge. A very small denticle in the center 
of the bridge. 

Inner side of premolar with two denticles. Anterior denticle larger 
than posterior. An additional denticle inwards of the posterior denticle. 
Structure of the two first molars generally like that of the premolar. The 
molars however, are generally larger. A well -marked depression between 
the outer and inner denticles, in the center of the tooth. This depression 
is particularly large in the long posterior molar. The posterior denticle 
(posterior to, and inward of, the second inner denticle) is better developed 
than on the preceding teeth. 

A description of the color has been given in the diagnosis of the species. 
Details and specific color characters will be given in the description of the 
subspecies. 

GEOGRAPHICAL DISTRIBUTION. The northern boundary of the range 
ofPteromys volans crosses Russian Lapland, turning south towards 
the east. According to I. F. Brandt ("Mammals of North European Russia" 
(Pozvonochnye severa Fvropeiskoi Rossii), 1856, pp. 31-32) the flying 
squirrel was distributed as far as Finnmark. However, this has not been 
confirmed by later authors. F. D. Pleske (1884) writes that this species 
does not occur near Lake Imandra. Fellmann observed the flying squirrel 
in Kuusamo, near 67° N. The northern boundary follows 66° N in Karelia, 
and 68° N near Kem. It touches the northern Arkhangelsk tundra reaching 
the Mezen and Kuloi rivers. The flying squirrel is rare in forests along 
the Kuloi river (B. M. Zhitkov, 1904). Farther east the boundary is not 
clear. It crosses the Pechora at 66°N reaching the Urals. The boundary 
apparently turns south in the northern Urals. Small numbers of this 
animal occur in the basin of the Lyapin. It rarely reaches the Synya river, 
It is common along the Sosva river (K.K. Flerov, 1933). Farther east 
the boundary runs to Berezov and continues in the forests along the Kazyma 
river. The northern boundary of the flying squirrel in the east is not 
known. However it does not enter the stunted forests of the tundra. The 
boundary turns south in an arc reaching Yanov Stan (near 66° N, where the 
flying squirrel was recorded by S. P. Naumov, 1931). Near the lower 
Yenisei the boundary passes near 68-69°N. It then continues eastward at 



257 



this latitude through Lake Zhesei and the upper reaches of the Anabara, 
and reaches the Olenek river (R. Maak, 1859). The boundary then crosses 
the Lena and Yana rivers at about 70° N. It turns south along the Indigirka 
and Kolyma rivers to 68° N. In the Anadyr Territory the flying squirrel is 
found along the Maina and Anadyr rivers (at about 65-66°N). 

The southern boundary includes Lithuania passing north of Vilnius, 
passes Minsk in the north and reaches Mogilev and Polotsk. Farther east 
it turns south to Dorogobuzh, Yelnya and Roslavl. It then turns to include 
the Belyi, Rzhev and Dmitrov forests. 

From Moscow the boundary turns south to Yegor' evsk, includes the 
Murom forests and reaches Arzamas, Kazan, and Yelabuga. It then follows 
the left bank of the Belaya river, turning sharply south to 52° N on the Urals 
where it passes between the Sakmara and Ik rivers. Farther east the 
boundary turns again to the birch groves of Shadrinsk and Chelyabinsk. It 
is difficult to say where the southern boundary passes between Petropavlovsk 
and Omsk. Flying squirrels apparently do not occur in the Borovoe 
Forestry (south of Kokchetav) . They occur farther east in the isolated birch 
groves north and west of Pavlodar. Farther southeast the flying squirrel 
is found in "ribbon" pine forests on the right bank of the Irtysh (e.g., in 
the Shulbinskii, Srostinskii forests, etc. ), that is, in typical habitats of 
Sciurus vulgaris exalbidus Pall. 

Farther south the boundary includes the Altai and Sayan mountains, 
runs to the southeastern shore of Lake Baikal, and includes northern 
Mongolia (vicinity of Mondy, Sungur Ravine). The flying squirrel does 
not occur in the Daurian Steppe. The range is interrupted on the Hentey 
Mountains. Farther east the boundary passes between the Shilka and 
Argun rivers (latitude of Nerchinsk), includes the northern part of the 
Khingan Range and curves to include the forests of Manchuria (from 
Tsitsihar andKirin). Farther south the boundary crosses the upper reaches 
of the Liao Ho and Lwan Ho rivers, reaches Peiping (Peking), and the 
Chinese provinces Shansi, Shensi, and Kansu, as well as Chihli [Hopei] 
and Shantung. The range of the flying squirrel covers Japan (islands 
Honshu, Yezo, [Hokkaido]) and the Russian and Japanese parts of Sakhalin. 

According to E. Lubicz v. Niezabitowski (1934) the flying squirrel 
formerly occurred in north Poland, near Wilno [now Vilnius], but it 
disappeared from this region long ago. According to L. Taczanowski (1887) 
it occurs in the Lithuanian forests and was caught near Borisov (former 
Minsk Province). It occurs in small numbers in the Mogilev forests. J. 
Domaniewski recently recorded it from Lithuania and Kurland. G. Schweder 
(1881) recorded it from forests near Kokenhusen [Koknese] and P. 
Wasmuth (1908), from former Kurland. 

According to A. V. Fedyushin (1929), it survives in small numbers in the 
southwestern territory only in the Vishera and Polotsk districts. In the 
Mogilev District it was found only in the Belynicheskoe forest. 

Farther east we find the flying squirrel in forests of the former Smolensk 
Province. According to V. A. Melander (1926) it now occurs only in the 
northern part of this province, in the Belyi and Demidov counties, and 
sporadically in other counties (Dorogobuzh County). M.I. Pogodin found 
here a colony of about 20 flying squirrels in a mossy marsh with small 
pine trees near the village Roslavka. It was also found in the southern 
part of the former Yelnya County near village Zabolot'e (54°15'N). This 
is the most southern record in this province. G. L. Grave (1926) writes 



258 



296 



that small numbers occur sporadically in the northwestern part of the 
former Belyi County of the Smolensk Province. 

In a later publication ("Commercial Hunting in the Western Region" 
(Okhotnich'i promysly v Zapadnoi oblasti), Smolensk 1933, p. 63) G. L. 
Grave writes that the flying squirrel is sometimes common in forests of 
the northern part of the Western Region. It was also recorded fronn central 
regions such as Rudnya, adjacent to the Demidov, Saratov, Yelnya, 
Yekimovichi districts. It occurs rarely and sporadically in forests of the 
southern parts of the Western Region. G. L. Grave recorded it here from 
the following places: Akulichskaya and Kleshnyanskaya wood lots of the 
Lyudinkovskoe Commercial Forestry, the Svenskoe and Beloberezhskoe 
forestries, some forests of the Roslavl, Sukhinichi, and Yershichi districts. 
The flying squirrel was found in 1913 in forests of the Korochizhsko- 
Krylovskoe Forestry. South of these districts the flying squirrel apparently 
occurs in forests of the former Karachev County of the Orel Province. 
K. F. Kessler (1858) writes that a flying squirrel was caught in October 
1851 in the Karachev County. S.N. Gorbachev (in "Nature of the Orel 
Territory" (Priroda Orlovskogo kraya), 1925, p. 452) writes that he found 
in the collection of the former Orel Military School a specimen with the 
label "Karachev County, 1856". Gorbachev thinks that this specimen was 
obtained by A. S. Tarachkov. According to K. A. Satunin (l895), the flying 
squirrel rarely occurs farther northeast in the former Bogorodsk County 
of the former Moscow Province (near the boundary with the former 
Yegorevsk County of the Ryazan Province). It was once caught in the 
Sokolniki near Moscow. I received one specimen from V. G. Durov from 
the former Dmitrov County of the Moscow Province and gave it to the 
MSZM. According to V. V. Raevskii (1929), a flying squirrel was caught 
near Verbilki (former Dmitrov County), near Rasheniya (former Leninskii 
County, in the Tugolesskoe and Kipriyanovskoe forestries). 

According to V. A. Varentsov (1919), it is rare in the former 
Pereyaslavl County of the Vladimir Province. In my collection are 
pelts from near the village Zabolot'e (between the former Aleksandrovsk 
and Pereyaslavl counties of the Vladimir Province). It was stated that 
flying squirrels are common in this region. 

Flying squirrels are also common in the forests occupied by the hunting 
collective "Dynamo" near the Novye Petushki railway station (former 
Pokrov County of the Vladimir Province). In this area flying squirrels keep 
to definite localities. K.S. Satunin ("Primitiae Faunae Mosquensis", 1892, 
p. 2) writes that the flying squirrel occurs rarely at the boundary between 
2Q,^ the Moscow and Ryazan provinces. S.S. Turov (1925) records it from the 
Yegor'evsk County (from a specimen in the MSZM). According to E.A. 
Bekshtrem (1927), it occurs in the Meshchera forests. L. G. Kaplanov and 
V. V. Raevskii (1928) write that the flying squirrel is common near 
Krivyandino railway station in the Tugoles Forest, former Yegor'evsk 
County and near Lake Svyatoe, former Ryazan Province. According to 
these authors* it occurs farther north in the upper reaches of the Mologa 
and also in the former Bezhetsk County of the Tver Province. L. P. 
Sabaneev (1868) writes that the flying squirrel occurs in the former 
Poshekhon'e -Volodarsk County. It occurs more or less sporadically in the 



* W. W. Rajewski und L. G. Kaplanow, Zoolog. Anzeiger, Bd, 84 H. 5/6, 1929, S. 1^ 



259 



298 



other counties of the former Yaroslavl Province. One specimen was 
caught 30 km from Yaroslavl on the boundary with the Uglich County. 
Another was caught in the Uglich County not far from the boundary with the 
Romanovskii County. Another was caught in the Romanovskii County (along 
the Uglich Highway). According to A. V. Shestakov (1926), about 400 pelts 
obtained in 1924-1925 by hunters of the Uglich County were stored. Flying 
squirrels were also found in the Varegovo Marsh and near the village 
Talovinkino in the Uglich County. According to K. A. Vorob'ev (1927), 
it occurs in the former Kostroma Province. A.N. Formozov (inlitt., 1935) 
writes that the flying squirrel is very common in forests near Ponazyrevo 
railway station, Vetluga County of the Kostroma Province. S.U. Stroganov 
(in litt. ) found it near Lake Peno (former Ostashkov County of the Tver 
Province). It is rare near Vyshni-Volochek. Chistovskii ("The Pskov 
Territory" (Pskovskii krai), 1927) writes that the flying squirrel is rare 
in the forests of the Velikie Luki and Kholm counties. According to K. F. 
Kessler (1885) it occurs in the former Vologda, Novgorod and Petersburg 
Provinces. V.N. Shchegolev (1925) recorded it from forests near 
Cherepovets. 

Specimens from the Leningrad (former Petersburg) Region are in the 
ZMAN collection. K. F. Kessler (1868) states that the flying squirrel is 
found in the former Olonets Province, where it is much rarer than the 
squirrel. 

In the north it occurs in Finland (I. F. Brandt, 1856), in the Kuusamo 
District (Fellmann) and rarely in forests along the Kuloi (B.M. Zhitkov, 
1904). 

Distribution of the flying squirrel in the Volga Region. According to 
A.N. Formozov (in litt., 1935), it is common in the former Balakhna, 
Makaryev and Semenov counties of the Nizhni Novgorod Province. I saw 
a specimen from the bank of the Vetluga, northern part of the former 
Semenov County, Nizhni Novgorod Province. 

According to M.N. Bogdanov (I87l), the flying squirrel occurs in the 
northwestern part of the former Kazan Province (in spruce forests). 
In about 1820 the flying squirrel occurred on the watershed of the Ashita 
and Kazanka rivers (former Kazan County). Bogdanov did not find it in 
the oak forests of the former Laishevo County. According to I. Bashkirov 
and N. Grigor'ev ("A Survey of Commerical Hunting in Tataria" 
(Ocherk okhotnich'ego promysla Tatarii). — Rabota Volzhsko-Kamskoi 
promyslennoi stantsii. Issue I, 1931, p. 35), the flying squirrel occurs 
rarely in the Lubyanskii Forest (on the boundary between the Mamydyshskii 
Canton and the Vote Region [Udmurt SSR]), in the forests of the 
Krasnoborskoye Forestry (Yelabuga Canton), and in the Zelenodolskii 
Forest near Kazan. Farther northeast it occurs in the Vyatka and Perm 
forests (M.N. Bogdanov, "Studies on Russian Hunting" (Etyudy russkoi 
okhoty), in "Okhota i konnozavodstvo". No. 5-6, 1874, p. 119). 

According to A.N. Korsakov (in litt. , 1933), it occurs in the Vote 
Region in forests of the Mozhga District. It occurs rarely in the Mozhga 
Forestry near the village Oshmest', 16-18km southwest of Syuginskaya 
railway station, at 56°15'N and 22° E (near Pulkovo). In the Zyryan Region 
(Komi) in the former Ust-Sysolsk County of the Vologda Province it 
occurs rarely in the upper reaches of the Vychegda. In the Udorskii 
District (Komi Region), in the northern part of the former Vologda Province 
and in the southern part of the former Arkhangelsk Province it occurs 



260 



rarely in the upper reaches of the Mezen (62°30' and 64°40' N and 45°30' and 
50°35'E). 

According to L.P. Sabaneev (1874), it occurs rarely on the Urals in the 
Goroblagodatskii and Bogoslovskii districts. It is connmon in the 
Yekaterinburg [Sverdlovsk] birch forests. According to K.K. Flerov (1933), 
it occurs rarely in the northern Urals along the Lyapin Basin, sometimes 
as far as the Synya river. It is common along the Sosva. It was found by 
N.M. Dukel'skaya (1928) on the southern Urals in the Ilmenskii Range. 
E.E. Eversman (1850) recorded it from pine and spruce forests of the 
Urals, especially from the central and northern Urals. It is found at 52° N 
in forests between the Sakmara and Ik rivers, but not south of this latitude. 
On 14 December 1880, i N. A. Zarudnyi (1897) found a flying squirrel in the 
lower reaches of the Sakmara, in the Karaevskaya Grove. The forests 
along the lower reaches of the Sakmara are separated from the forests in 
the middle and upper reaches of the river by a large area without forests. 

According to N. A. Zarudnyi the flying squirrel was common in these 
districts. 

According to I. Ya. Slovtsov (1892) the flying squirrel is common in 
the Tumen district in the Trans-Ural forests of western Siberia. It is 
numerous in the Turinsk District and rare in the Pelym District. It is 
very rare near Berezov. It occurs on the Konda, occurs regularly in the 
Surgut Territory and rarely in the northwestern part of the Yalutorovsk 
District and the western part of the Kurgan District. It is found regularly 
along the Urals. S.M. Chugunov (1915) recorded one specimen caught in 
the Surgut taiga in the Yugan Valley. In I. Shukhov's opinion (19 28), the 
flying squirrel is distributed throughout the marshy taiga of the former 
Tara District, where it occurs in small numbers. 

According to A.M. Belyaev (1934), the flying squirrel is found in 
"ribbon" pine forests of the former Semipalatinsk Region and inbirch groves 
on the right -bank region of the former Pavlodar County in southwest 
Siberia. M.K. Serebrennikov (1929) recorded it from forests 200 km north 
of Pavlodar. O.K. Holtzmeyer (1935) reports that the flying squirrel is 
common in pine forests of the former Semipalatinsk Region, e.g., in 
the Srostinsk Pine Forest. 

Finch obtained the flying squirrel on the Altai near the village Altaiskaya 
(A.M. Nikol'skii, 1884). It was later found by N. F. Kashchenko (1902) in 
mountains on the right bank of the Chulyshman, on the eastern coast of the 
Teletskoe Lake, and also near the village Chergi (N.F. Kashchenko, 1899). 
Specimens from the upper reaches of the Bukhtarma, from near Barnaul are 
in the ZMAN collection. 

In my collection is a pelt obtained by Razorenova in the valley of the 
Katun. V. Selevin (in litt. , 1935) informed me that the flying squirrel 
occurs in the Kolbinskie Mountains. Farther east it occurs in the Sayan 
Mountains (G. Radde, 1862), and in the former Minusinsk District (on the 
Tukeek-Kem, Suturzhu-Karasuk — B. S. Vinogradov, 1927). It also occurs 
in the mountains near the village Oznachennoe and Monok in the former 
Minusinsk County (N. Dukel'skii, 1930), in the Minusinsk and Khakassian 
districts (M.D. Zverev, 1930), in the former Krasnoyarsk and Achinsk 
counties (I.N. Shukhov, 1925). V. V. Gubar obtained numerous specimens 
in the Troitsk District (northern part of the former Biisk County of the 
Tomsk Province). Two specimens were caught by the expedition of V. P. 
Anikin (1902) in the Narym Territory. G. E, lohansen (1923) reported it 



261 



300 



from near Tomsk and forests along the Chulym river (near the village 
Novokuskovo, Tranikovo Locality). Northeast of these districts the flying 
squirrel occurs in forests along the Yenisei and Tunguska rivers. According 
to N.P. Naumov (1934), it is common along the Lower Tunguska. According 
to A. Middendorf (l86l), it reaches 68°N. along the Yenisei. Tell 
(Nordenskjttld's "Expedition to the Estuary of the Yenisei" (Ekspeditsiya k 
ust'yam Eniseya, 1880, p. 178) saw many pelts in Turukhansk and 
Verkhneibatsk. E.O. Yakovlev (1930) reports that the flying squirrel occurs 
in some parts of the forest tundra in the Turukhansk Territory, e.g., in Lake 
Chirinda Region. 

According to S.P, Naumov (1931), the animal was observed near 
Yanovstan (66°N). Records from the forest tundra (near 67°N) are doubtful. 
East of the Yenisei it reaches about 69° N near the Norilskie Lakes (R. 
Maak, 1886). In the Olenek Valley it occurs as far as 68° N. It is common 
along the Vilyui except in its upper reaches. It is widely distributed in the 
LenaBasin(I. Polyakov, 1873) and on the Aldan (ZMAN collection), is common 
near Yakutsk (S.I. Ognev's collection), and occurs in the Verkhoyansk 
District (A. Bunge, 1887). It certainly occurs along the Indigirka. In the 
Kolyma Valley it was recorded from Verkhe-Kolymsk and Sredne-Kolymsk 
(J. A. Allen, 1903; S.l. Ognev, 1926). According to N. P. Sokol'nikov 
(1927), it occurs in deciduous forests 50-60 km from the village Markovo 
(along the Anadyr river). These forests follow the river for 100 km to its 
upper reaches. The flying squirrel also occurs in forests along the Maina 
river from its left tributaries, i.e., the Algans, L.A. Portenkov (1932- 
1934) obtained a good series near the village Yeropal on the Chukot 
Peninsula. It does not occur in Kamchatka. It was found along the shore of 
the Sea of Okhotsk, near Okhotsk, Ola and Ayan. It occurs in forests in 
Gizhiga and is common on the Shantarskie Islands. According to L. v, 
Schrenck it is found in the valley of the Amur, near Nikolaevsk-on-Amur, 
from where I examined specimens, and also in the Amur Estuary, at the 
coast of the Tatar Gulf, in the De-Kastri Region, as far as Bai-Khadzhi in 
the south. It is widely distributed on Sakhalin where it is sometimes 
common. In my collection is a good series from Argi-Pagi (north 
Sakhalin). The expedition of V.S. Stakhanov and K. A. Vorob'ev (1932) 
caught a large series in the Amur Basin, from the Khungari river. Lake 
Bolon'-Odzhala, Khor river, a tributary of the Ussuri, De-Kastri Gulf, 
and Taba Bay in the Tatar Gulf. It was caught in Olga (Maksimovich — 
ZMAN collection), and also near Petropavlovskoye Lake, 50 km from 
Khabarovsk (ZMAN collection). According to G. N. Gassovskii (1927) 
the flying squirrel occurs farther east in the Gilyui-Ol'doi Mountain Region, 
the Teokit and Koval basins, on the northern slope of the Tukuringra 
Range, along the Bezymennaya river, and ravines along the right 
tributaries of the Ol'doi river. In the country of the Orochis A. A. 
Emel'yanov (1927) found flying squirrels in Bouchi and Kopi. 

According to Radde (1862), the flying squirrel occurs rarely in the 
Burein Mountains and also in the middle reaches of the Onon river, along 
the Shilka and Argun rivers, and on the southern part of the Yablonovoi 
Range (Sokhondo Mountain, and at the source of the Ingoda, near Akshinsk 
and Mogotui). 

According to N. F. Kashtchenko (1910) it occurs in Transbaikalia in the 
former Verkhneudinsk [Ulan-Ude] and Troitskosavsk [Kyakhta] counties, in 
birch forests of the Aga Valley (N, F. Kashtchenko, 1913). B.A. Kuznetsov 



262 



(1929) observed the flying squirrel in the Yablonovoi Range taiga and its 
foothills, near the village Kyker, along the Shantelka, Shantera, 
Shanterkeken rivers and other tributaries of the Nercha river; near village 
Zyuzel'kan, on the Udinskii Range, etc. According to S. Sofronov (1926), 
it is common near Chita. It is not rare in the Barguzin forests (G. P. 
Doppelmaier, 1926) on the Koty Peninsula in Baikal (S.I. Ognev, 1929). 
In the ZMAN collection are specimens from the Hentey Mountains (Baik-Gol 
Area) and from the Sungur Ravine (northwestern Mongolia). 

It was recorded from the Ussuri Territory by N. M. Przheval'skii 
(1870). 

In the ZMAN collection are specimens from the following places: Poset 
Bay (collected by M. Yankovskii); the upper sources of the Kulume 
(collected by V. K. Arsen'ev); vicinity of Cossack village Razdel'naya 
(collected by A.N. Formozov). In my collection is a specimen obtained 
near Okeanskaya railway station (collected by E. M. Vermel'). According to 
Sowerby (1923), it occurs in the western part of the Shansi Province and in 
Chihli [Hopei] northeast of Peking). It also occurs in Korea and Japan (on 
Honshu and Hokkaido). 

It is not exactly known how far south the flying squirrel occurs in China. 

Map V shows the distribution of the various subspecies of the flying 
squirrel. 

MODE OF LIFE. In the central section of the European part of the 
U.S.S.R. the flying squirrel occurs mainly in mixed forests, especially in 
coniferous forests mixed with birch and aspen. 

A.N. Formozov (in litt. , 1936) found flying squirrels in the Gorki Region 
(former Kostroma Province and also former Makar'ev and Semenov 
counties of the Nizhni Novgorod Province) in humid spruce or spruce-fir 
forests mixed with birch, alder and aspen. 

In the southern Trans -Urals flying squirrels are common in birch groves . 

Along the Irtysh they occur in the region of "ribbon" pine forests 
together with S c iu ru s vulgaris exalbidus Pallas. According to 
I. Ya. Slovtsov (l892), flying squirrels occur in mixed forests in the 
Tobolsk Territory where they find pine trees with holes made by the black 
woodpecker. The flying squirrel apparently does not avoid "Urmans", 
i.e., the marshy taiga of the northern Tobolsk Territory. 
301 According to B.A. Kuznetsov (1929), the favorite habitats of the flying 
squirrel in Transbaikalia are high forests with many old, hollow trees. 
The animal is especially fond of thickets along banks of brooks and streams. 
Near the village Kyker the flying squirrel most frequently occurs in the taiga 
along the Shamtelka, Shantera, Shanterkeken rivers and other tributaries 
of the Nercha. In the vicinity of the village Zyuzil'kan it mainly inhabits old 
birch-larch forests in the valleys of the Zyul'za and Zyulzil'tan rivers. 
On the Udin Range the flying squirrel occurs at present only in small parts 
of the taiga with pine and larch forests which still exist in some inaccessible 
localities. 

K, A. Vorob'ev (in litt. , 1933) describes the habitats of the flying squirrel 
in the Amur Region: "The flying squirrel lives in forests of willow, aspen, 
hornbeam, Juglans mandshurica, black birch, spruce and fir. We also 
found it on slopes of hills covered with mixed forests of cedar, fir, spruce, 
black birch, hornbeam and yew shrubs (e.g., along the Khungari river). 
It occurs frequently in forests with numerous old, hollow pine trees in 
which it builds its nest. " 



263 



In the mountains of Siberia, e.g., on the Altai, the flying squirrel is 
fovmd in typical habitats. It was found by A.N. Kolosov in a birch forest 
at an altitude of 500m near the Teletsk Lake. In summer 1935 flying 
squirrels were caught on the Altai by Professor S.S. Turov's .-expedition, 
in a large forest at an altitude of about 1000 m. 

In extreme northeastern Siberia, e.g., in the Anadyr Territory, the 
flying squirrel occurs in larch forests along the Anadyr. 

We know very little of its habits. It begins its activity l/2 -Ihour after 
sunset. Its characteristic voice can then be heard from hollows, or from 
trees. The voice of the flying squirrel resembles the chirping of bats 
like Nyctalus noctula. A. Rekst told me that he heard the voice of 
the flying squirrel regularly on Sakhalin. The best indication of its 
presence on Sakhalin are the large, yellowish pellets usually iound in heaps 
near the base of trees or in the forks of large branches. 

One rarely sees tracks of flying squirrels in the snow. They resemble 
tracks of squirrels but are much smaller and the leaps are very short. The 
tracks are rarely longer than 10- 15 m, beginning at the base of one tree 
and ending at the base of another (A.N. Formozov, in litt. , 1936). 

The most peculiar habit of the flying squirrel is its ability to fly by 
means of its parachuting membrane. According to V. V. Raevskii and 
L. G. Kaplanov (1929), the flying squirrel is capable of leaping from the 
top of a tree and gliding for 40 m. The glide descends, and the animal 
lands by clinging to the bark, halfway up the tree. The flying squirrel 
glides slowly in a straight line. 

During flight the animal spreads its forelegs laterally. The spurs of 
the forefeet facilitate the stretching of the parachuting. membranes. The 
hind legs are kept close together along the tail. The silhouette of the 
gliding animal is thus triangular. 

The animal leaps from the top of a tree and glides down. However, it 
occasionally changes its direction and glides upwards before landing 
on a tree. This ascent may be caused by an air current against 
the spread parachuting membrane. 

The glide of the flying squirrel is noiseless. The flying squirrel appears 
unexpectedly in the twilight in a glade among the trees or in a clearing. 
V. V. Raevskii (1929) reports that in the Tugoles Forestry (former Moscow 
Province) flying squirrels are often seen in spring during courtship of the 
woodcock. The hunters occasionally shoot flying squirrels, mistaking them 
for woodcocks. 

Our knowledge of the daily life of the flying squifrel is very incomplete. 

Most authors state that the flying squirrel is crepuscular and nocturnal, 
hiding during the day. V. V. Gubar' (1930) writes that bright daylight is 
disagreeable to it. 

Exceptionally the flying squirrel is active during the day. A.Cherkasov 
(1867) writes "In Transbaikalia, I often saw flying squirrels unexpectedly 
in the forest in broad daylight. While I was riding slowly along a path. . , 
something would suddenly fly from a tree, swiftly flit near the head of the 
horse and disappear among the branches of trees. . . Taking it for a hazel 
hen I would dismount, take my gun and hide . I would then see a flying 
squirrel sitting on a branch, looking down with its large, black eyes. . . the 
flying squirrels are bold, agile animals. They are easily shot as thev are 
not afraid of man. " 



264 



303 



A.N. Formozov (in litt, , 1936) writes that he killed a flying squirrel 
at noon in the middle of November, The animal was returning to a hollow 
in a tree which had been felled in order to examine the hollow. 

The flying squirrel generally chooses hollows in trees as shelters 
for its nest. In the central part of the European U. S. S. R. flying squirrels 
usually occupy hollows in old aspens. In the Gorki Region A.N. Formozov 
(in litt. , 1936) found nests in holes of the spotted and black woodpecker in 
alders, birch and aspen. These nests were 3 -10 m above the ground. One 
tree would sometimes contain several hollows, each occupied by a nest. 
The nests were built of lichen (Usnea barbata and U s n e a 
longissima). 

In birch groves in Bar aba the flying squirrel usually inhabits hollows in 
birch. In forests of the Tobolsk Territory I. Ya, Slovtsov (1892) found 
nests in pine trees. These hollows had been made by black woodpeckers. 
According to B. A. Kuznetsov (1929), flying squirrels make their nests in 
Transbaikalia in hollows in old birches or larches, sometimes in 
abandoned nests of woodpeckers (D end r o c opu s major andPicus 
martius ). 

V. V, Gubar' (1930) gives the following data for the Novosibirsk flying 
squirrel: "Flying squirrels are usually found in hollows in high aspens 
3-4 m above the. ground. Only once did I find an inhabited hollow very low, 
at a height of 1.5 m. It is simple to determine whether a hollow is 
inhabited. Whenthe trunk is tapped with an ax, the flying squirrel comes half 
out of the hole or jumps out completely. Confusedby the light, its movements 
become very slow. The frightened animal climbs about 1 m along the trunk 
and then stops, clinging to the tree and spreading its parachuting membrane. 
After about a minute the animal slowly climbs higher. Due to its protective 
color it is alm.ost invisible on the bark. At the top of the tree the animal settles 
on a branch, rolls up into a ball and suddenly jumps into the air. The 
parachute spreads and the small glider gradually descends to a nearby tree. 
Flying squirrels never land on a branch. They always land on the trunk 








^^ 



FIGURE 163. Flying squirrel on a bough 



Drawing by K. K. Flerov 



265 



about 1-2 m above the ground. They do not land in the middle of the trunk 
but at the side, i.e. , tangentially, apparently in order to absorb the shock. 
In other cases, the flying squirrel crouches at the top of the tree as if 
preparing for flight, and then stays motionless, ignoring stones and sticks 
thrown at it. " 

Flying squirrels generally build their nests in hollows, but sometimes 
also in starling boxes. This was observed by G.E. Johansen (1923) during 
the expedition to the Chulym, by V. V. Gubar' (1930) in the Ust'-Chemsk 
Legostaevskii District (former Novosibirsk District) and by M. D. Sverevoi 
(see Gubar 's work), who found a nest in a starling box in a pine forest near 
the Berek Station of the Altai Railway. 

A.M. Kolosov informed me that he found a nest in the hollow of a birch 
which had been previously inhabited by wild bees. A. Cherkasov (1867) 
observed in Transbaikalia that flying squirrels usually descend to defecate 
in one place near the roots of the tree. Such "latrines" are a reliable 
indication of a colony of flying squirrels. A. Cherkasov writes: "If one 
finds such a heap of feces one has only to tap the tree with a stick and the 
squirrel will invariably jump out of a hollow. Flying squirrels also often 
inhabit abandoned nests of birds or squirrels. 

K. A. Vorob'ev (in litt. , 1933) studied the structure of nests of flying 
squirrels in the Ussuri Territory and Amur Region. 

He writes: "The height above the ground at which the squirrels make 
their nests varies considerably. All nests which we found in the Amur 
Territory in summer 1932 were in hollows in willow, aspen or hornbeam, 
5-9 m above the ground. Most of these hollows had apparently been made 
by the black woodpecker*. We felled some trees with nests in order to 
examine the material of which the nests were made. They were always 
built of dry grass, small roots and moss. There were sometimes also 
a few feathers of hazel hen, nutcracker or other birds. 

"On 14 August 1932 we found a nest near the Khungara river at the edge 
of water in a wood of willow, hornbeam and many shrubs. The nest was 
in a dry willow 5 m above the ground. The opening was 6 cm wide. The 
nest was at the bottom of the hollow, 63 cm below the entrance. It 
consisted of moss, rotten wood and bird feathers. No remains of food were 
found in it. " 

We have no detailed information on the food of the flying squirrel. V. V. 
Raevskii and L. G. Kaplanov (1929) thought that it fed mainly on nuts of 
spruce. V. V. Gubar' writes: "We dissected the stomach of all flying 
squirrels obtained in order to determine their food. Almost all the animals 
dissected were caught in September. Their stomach contents indicated that 
they fed almost exclusively on pines. This was assumed from the presence 
in the stomach of a green bolus with a strong pine smell. The food was so 
finely chewed that it was impossible to determine its origin. It apparently 
consisted of young shoots and green pine cones. " 

A.N. Formozov (in litt. 1936) found food reserves consisting of alder 
catkins and small twigs in nests and hollows of flying squirrels in the Gorki 
Region. The weight of these stores was 30-400 g. There were also 
occasional birch catkins and spruce cones. 

According to I. Ya. Slovtsov (1892), the food of the flying squirrel mainly 
consists of birch buds. The hunters think that it also eats the pulp of pine 

* Cf. the observations of I. Ya. Slovtsov (1892) in the Tobol'sk Territory. 



266 



305 



306 



cones and frozen mushrooms. I.Ya. Slovtsov writes: "I fed flying 
squirrels in captivity with all three foods mentioned. First of all they ate 
fresh young birch buds. Then they gnawed the mushrooms. Only when 
there was no other food, did they accept pine cones. The completely ignored 
pine nuts. 

"The flying squirrel is rather voracious. One pair consumed ■! .5 lbs 
of buds daily. " 

A. Cherkasov (1867) writes: "The flying squirrel feeds mainly on birch 
and aspen catkins. Hunters think that it also eats small birds which it 
catches at night on trees and in nests. They also stated that it attacks 
even squirrels, suffocating them in their nests. " 

V.I. lokhel'son (1898) reports that Kolyma hunters also think that the 
flying squirrel attacks squirrels in their nests. The flying squirrel 
spreads its parachuting membrane during the fight. The squirrel bites 
through the parachuting menabrane. However, the flying squirrel allegedly 
always wins. It is, however, incapable of using its damaged parachute for 
some time. G. K. Gol'tsmaier (1935) hear similar stories from hunters 
during his studies of S c iuru s vulgaris exalbidus in the Srostinsk 
Pine Forest (south of the former Semipalatinsk Region). 

These reports are doubtful. However the fact that such beliefs are held 
by hunters in different and far removed districts makes it necessary to 
consider them seriously and to verify them. 

Our knowledge of the reproduction of the flying squirrel is very 
incomplete. Stoll' (1906) writes that the Baltic flying squirrel has two 
litters per year, one in late February and the second in June, Each litter 
consists of 2 or 3 young. P. Wesmuth (1908) on the other hand, claims 
that the Baltic flying squirrel has only one litter in June, that its gestation 
is 4 weeks, and that the litter consists of 2 or 3 young. 

A. Cherkasov (1867) writes that the Transbaikal flying squirrel has only 
one estrus, in spring, and that it has 1 or 2 young. 

On 21 July 1929 B. A. Kuznetsov found 4 young and a suckling female in 
a hollow of an old larch on the bank of the Shanter in Transbaikalia. The 
young were already able to run on the trees, jumping from one branch to 
another . 

A. P. Razorenova found a young flying squirrel on 10 July 1930 on the 
bank of the Katuni on the Altai. On 18 August 1932, A. Larovskii found a 
very young specimen near the Khungari river. The fur on the tail was 
little developed and much shorter than in adults. 

N. P. Sokol'nikov (1927) writes that in the Anadyr Territory the flying 
squirrel has only one litter a year (late April-early May) and no more than 
3 young. 

As we have not enough material we cannot determine exactly the molting 
periods of the flying squirrel. The sequence of the autumn molt of 
Pteromys seems to be the opposite of that of the squirrel. In the flying 
squirrel the molt starts at the head, occiput and shoulders, continuing 
towards the tail. The tail of the flying squirrel acquires its luxuriant, long 
winter fur already in late summer. 

We also know little of its enemies, 

V, V. Gubar' (1930) reports that in western Siberia the flying squirrel is 
preyed upon by cats. It is apparently also preyed upon by ermines, 
martens and owls. 



267 



307 



Flying squirrels are not hunted much. According to B. A. Kuznetsov 
(1932), the number of pelts dressed in the U.S.S.R. is very small. This is 
however true only of the U.S.S.R. as a whole. In some districts hunting 
of flying squirrels is intensive, e.g. , in some parts of Transbaikalia 
(B.A. Kuznetsov, 1929). 

According to K. A. Vorob'ev (in litt. , 1933), hunting of the flying 
squirrel is increasing in the Amur territory. It was hunted in the Amur 
Territory only accidentally in the past. According to figures of furs 
collected in 1928-1929, 16,184 flying squirrels were obtained in the Far 
Eastern Territory. 

Subspecies of the flying squirrel 

350. Pteromys volans volans Linn. (1758). Common flying 
squirrel 

1758. (Sciurus) volans Linnaeus, C. , Systema Naturae, editio X, p. 64 (nee Mus volans Linnaeus, 

C. , 1. c. , p. 63); Pallas P. , Novae Species Quadrupedum e Glirium Ordine, 1778, pp. 355-369; 

Pallas P.S., Zoographia Rosso-Asiatica, 1811, 190- 191 (partim.'). 
1808. Pteromys russicus Tiedemann, Fried. Zoologie zu seinen Vorlesungen entworfen. Zoologie 

des Menschen und der Saugethiere, I, S. 451 (Russland). 
1822. Pteromys sibiricus Desmarest A. G. , Mammalogie ou description des esp^ces de 

Mammiferes, II, p. 342. 
1829. Pteromys borealis Dvigubskii I. , Opyt estestvennoi istorii vsekh zhivotnikh Rossiiskoi imperii 

(A Natural History of the Animals of the Russian Empire), p. 32. 
1843. Pteromys vulgaris Wagner A. , in: Schrebers Ch. Saugethiere, Supplementum III, S. 228 

(instead of volans.'). 
1851. Sciuropterus vu 1 ga ri s Simashko Yu. , Russkaya fauna (Russian Fauna, II), pp. 597-599. 
1857. Pteromys volans Blasius J. F. , Naturgeschichte der Saugethiere Deutschland, S. 26; Bogdanov 

M.N. , Etyudy russkoi okhoty. Okhotnich' i promyslovye zveri i ptitsy Evropeiskoi Rossii (Studies on 

Russian Hunting. Game and Commercial Mammals and Birds of European Russia) "Okhota i 

konnozavodstvo". No. 5-6, 1874, p. 119. 
1874. Sciuropterus sibiricus, Brehm A. , Zhizn' zhivotnykh. II (Life of Animals, II) p. 64. 
1898-1899. Sciuropterus volans Trouessart E. L. , Catalogus Mammalium, I, pp.401-402. 
1903. Sciuropterus russicus Allen J. , Bull. Amer. Mus. Natur. History, XIX, p. 132 (partim.'); 

Trouessart E. L. , Faune des Mammiferes d'Europe, 1910, p. 116; Miller G. S. , Catalogue of the 

Mammals of Western Europe, 1912, pp, 941-946; Ognev S. I. , Mlekopitayushchie Moskovskoi gubemii 

(Mammals of the Moscow Province), 1913, pp. 281-282. 

TYPE LOCALITY AND DEPOSITION. The type locality is not known exactly. The holotype was lost. 
Linnaeus states "In borealibus Europae, Asiae et Americae". 

In G. S. Miller's opinion, use of the phrase "in borealibus Europae" means that Linnaeus was describing 
a flying squirrel from Sweden. However, G. S. Miller names the European flying squirrel Sciuropterus 
russicus (see above), accepting the name proposed by Tiedemann. In Tiedemann's opinion, the type 
locality, "Russia" is "too vague". This opinion is shared by Miller. If we accept Linnaeus' name for the 
European flying squirrel, central Sweden should be considered as type locality ofPteromys volans 
volans. 

MATERIAL EXAMINED: 44 pelts, most of them with skulls. 

DIAGNOSIS. General color of winter fur ash-gray with a slight straw 
tinge (darker than drab-gray, Ridgway, 1912), more intense than light 
mouse-gray but lighter than mouse-gray. Tail dorsally dark grayish brown 
in the middle and pale straw-colored laterally. Tail black-gray with 
an irregular lighter grayish pattern ventrally. Summer fur on back blackish 
gray (dark mouse -gray) with straw-gray mottling (between drab and mouse- 
gray). Tail of summer fur smoke -gray dorsally. Middle and apical parts 
of tail with a strong admixture of hair with broad black tips, giving a grayish 
tinge. 



268 



Length of skull in full-grown specimens 38.2-41.3 mm (M. 39.92 mm); 
condylobasal length 34.9-37.8 mm (M. 36.4 mm); zygomatic width 22.8- 
25.2mm (M. 24.02mm); length of nasals 11 .9- 13 mm (M. 12.33 mm); length 
of upper molar row 7.1-7.7 mm (M. 7.28 mm). 

Length of body 149- 172 mm (M. 164.7 mm); tail without terminal hair 
98-130mm (M. 113.1 mm); hind feet without claws 32-39mm (M. 35.9mm). 

ADDITIONAL DESCRIPTION. Color of winter fur based on specimens 
from Vladimir, Kostroma and Nizhni Novgorod. General color of back ash-gray 
with a slight straw tinge (darker than drab-gray, but lighter than mouse - 
gray). Head, entire back and large parts of parachuting membranes of the 
same color as the back. The color is formed from the following elements: 
l) broad, gray roots of hair (deep neutral gray); 2) wide light straw-gray 
subapical parts of hair; 3) black hair tips of almost all hair on the back. 
There is also long guard hair with broad, black tips. Broad dark roots of 
hair clearly show on the margins of the outstretched parachute. The color 
of the margins of the parachute is therefore blackish slate -gray. 

Middle of dorsal part of tail dark grayish brown (between hair -brown 
and mouse-gray). Terminal part of tail with a strong admixture of 
blackish hair. Tail dorsally pale straw-gray at the sides. The color of the 
hairs on the dorsal side of the tail is as follows: l) roots gray (mouse - 
gray); 2) pale straw-gray subapical belts; 3) black tips. Many hairs with 
broad black tips everywhere give a dark color to the dorsal side. Ventral 
side of tail black-gray with an irregular grayish pattern consisting of 
more deeply colored stripes from the sides to the middle. 

Underparts white with a well marked yellowish tinge especially on the 
belly. Roots of hair ash-gray. 

SUMMER FUR. Back blackish gray (dark mouse -gray) with straw- gray 
mottling (between drab and mouse -gray). Color of hairs as follows: l) 
roots broadly black-gray; 2) a narrow gray band; 3) a straw-gray 
subapical band; 4) hair tips blackish. 

Top of head like that of back. Parachutes black- gray, grayish only at 
the margins. Dorsal side of tail smoke -gray at the sides (between pale 
mouse -gray and palid mouse -gray). Middle and apical parts of tail consisting of 
hair with gray (neutral gray) roots, whitish gray subapical parts and broad 
black tips, which give a generally dark color. Color of ventral side of tail 
whitish gray with an irregular black pattern. 

Dorsal side of feet black-gray mixed with whitish gray hair. Ventral 
side yellowish white. Outer posterior margin of feet with very long hair. 

Belly dirty whitish with a slight straw-yellow tinge. 

Roots of hair on belly pale gray. 

Hair of lower part of back 17-18 mm long in winter fur, in summer 
fur 12- 14 mm. 

Summer fur thinner and less luxuriant than winter fur. 

SYSTEMATIC NOTES. Until recently it was assumed that the whole 
European U.S.S.R. is inhabited by only one subspecies of the flying 
squirrel. However, a series obtained by S.U. Stroganov in the Western 
Region (former Ostashkov County, and from the Velikie Luki District) 
showed that this region is inhabited by a well characterized straw-rust 
colored form (P . v . ognevi Strogan. ). These color characters should not 
be considered as accidental. Near the eastern boundary of the range of the 
nominate subspecies occur specimens which are transitional to P. v. 



269 



gubari . Specimens of this subspecies already occur in the birch groves 
of Bashkiria (southern Trans-Urals). 

GEOGRAPHICAL DISTRIBUTION. The common flying squirrel is 
widely distributed throughout the European U.S.S.R. The northern and 
southern boundaries of its range and details of its distribution are discussed 
in the general survey. P . v . ogne vi Strogan. is distributed from the 
line Rzhev - Staritsa - Torzhok - Vyshni Volochek and farther west 
(apparently as far as the Baltic). The exact boundaries of the range of 
the common and western flying squirrels are not known. The common 
flying squirrel is replaced by P. v. gubari Ogn. in Bashkiria (southern 
Trans-Urals). 

351. Pteromys volans ognevi Strogan. (1936), Western flying 
squirrel or Ognev's flying squirrel 

1936. Pteromys ognevi Stroganov S. U, , Novye dannye po sistematike nekotorykh gryzunov (New Data 

on the Systematics of Some Rodents). Sbomik Nauchno-Issledovatel'skogo instituta zoologii 

Moskovskogo gosudarstvennogo universiteta No. 3, pp. 111-114. 
1936. Pteromys volans ognevi Stroganov S. U. , Fauna mlekopitayushchikh Valdaiskoi 

vozvyshennosti (The Mammal Fauna of the Valdai Hills), Zoologicheskii zhumal, XV (3): 539. Moscow 

1936. 

TYPE LOCAUTY AND DEPOSITION. Lake Peno in the Western Region (sources of the Volga, former 
Ostashkovskii County of the Tver Province). Type No. M.4158 female adult, 10 Dec. 1933, collected by 
S. U. Stroganov. S. I. Ognev's collection. 

MATERIAL EXAMINED: 10 specimens. 

DIAGNOSIS. Winter fur on back generally ash-gray with well marked 
straw-rust tinge which is most saturated on shoulders, neck and cheeks. 
Tail straw-yellow-gray with well marked black hair tips in the middle, 
dorsally and ventrally, which give a dark color to the whole length of the 
middle of the tail. Sides of tail lighter colored, without black hair tips 
and with a well marked yellow-straw-rust tone. 

MEASUREMENTS. Length of skull 36.7-39 mm; condylobasal length 
34. 7-36. 7mm; zygomatic width 23.9-24.5 mm; length of nasals 12. 4- 
12.9 nnm; length of upper molar row 6.6-7.5 mm. Length of body and head 
160- 163 mm; tail without terminal hair 95- 120 mm; hind feet without 
claws 34-37mm. 

ADDITIONAL DESCRIPTION. Winter fur. Straw-rust tinge, (close to 
avellaneous (Ridgway, 1912. pi. XL)) clearly marked on general gray color 
of back. Straw-rust tinge most saturated on shoulders, neck and cheeks. 
It is weak only on the dorsal side of the parachutes and on the dorsal side 
of fore- and hind legs. Margins of parachutes dark because of the dark gray 
roots of the hair. Ears gray. Tip of ears with a poorly developed whitish 
fringe which forms small tufts on the posterior side of the ears. 

All underparts whitish gray with a slight rust -straw tinge which becomes 
nnore intense in the inguinal region and on the ventral side of the parachutes. 

Tail yellowish straw-rust colored (near avellaneous, pi. XL, with pinkish 
buff tinge, pi. XXIX). 

Dorsal and ventral side of tail darker due to a considerable admixture 
of black caused by the broad black roots of the hairs. 

The summer fur resembles that of P . volans volans in its general 
dark color but differs in the pale straw-rust tone of the hair tips on the back. 
The tail also differs from that of the typical form in its more straw-rust 
color. 



270 



SYSTEMATIC NOTES. This subspecies differs clearly from the 
nominate form. Pteromys v. ognevi is characterized mainly by the 
well marked straw-rust tinge of both winter and summer fur, and the 
straw-rust colored tail. Dorsal and ventral side of tail with well developed 
black hair tips. 

P. V. ognevi resembles P. v. betulinus much more closely than 
the common flying squirrel. P. v. ognevi differs from the West 
Siberian flying squirrel in its dirtier gray color of winter fur. Its belly is 
not white as in betulinus but darkened by a gray- straw- rust tinge. 

Summer fur of P. v. ognevi very similar to that of P. v. betulinus 
310 P . v . o gne vi differs from P . V . at he ne as follows: l) winter fur 
not so light; 2) light hair tips on back of summer fur much less developed; 
in P. V. athene these hair tips are light straw-yellow. 

GEOGRAPHICAL DISTRIBUTION. Not known exactly. The eastern 
boundary of this form apparently runs along the line Rzhev-Staritsa - 
Torzhok - Vyshni Volochek - Novgorod. This subspecies apparently occurs 
in the former Pskov Volochek Province and the Baltic Region. More 
material should be examined in order to decide this. 

352. Pteromys volans gubari Ogn. (1934). West Siberian flying 
squirrel 

1902. Sciuropterus volans Anikin V. P. , Otchet o komandirovke v Narymskii krai (Report on an 

Expedition to the Narym Territory), 1902, p. 67. 
1923. Sciuropterus russi cus Johansen G. E. , Po chulymu (Along the Chulyma), Izvestiya Tomskogo 

gosudarstvennogo universiteta, p. 63; Gubar' V. V. , Zametki po biologii letyagi (Notes on the 

Biology of the Flying Squirrel), Izvestiya Sibirskoikraevoi stantsii zashchity rastenii, 1, 1930, pp. 78-79. 
1934. Pteromys volans gubari OgnevS.I. , Obzor letyag (Pte rom ys), Vodyashchikhsya v SSSR 

(A Survey of Flying Squirrels (Pteromys) of the U. S. S. R. ), Byulleten' Moskovskogo obshchestva 

ispytatelei prirody, otdel Biologii X, III (2) : 304-305. 

TYPE LOCALITY AND DEPOSITION. West Siberia, Troitsk District (northern part of the former Biisk 
County), No. M.4036 female, Dec. 1930, V.V. Gubar', type in S. I. Ognev's collection. 

MATERIAL EXAMINED: Holotype and 1 2 paratypes. 

DIAGNOSIS. Winter fur differs from that of the European flying 
squirrel in its lighter silver-gray color on the back with a slight straw 
tinge (which may be absent), and by the dull whitish gray tail. In 
Pteromys volans vo la n s the middle of the tail on the dorsal side 
is dark gray-brown and the lateral parts straw-gray. Summer fur differs 
from that of the common flying squirrel in its very pale dirty gray color 
with a straw tinge. 

Length of skull 39.2-41 mm (M. 40.2 mm); condylobasal length 35.7- 
36.9mm (M. 36.5mm); zygomatic width 23.1-24.5 mm (M. 23.8mm); 
length of nasals 12.1-13.1 mm (M. 12.5mm); length of upper molar row 
7-7. 5mm (M. 7.2mm). 

Length of body 135.5-171 mm (M. 155.3 mm); tail 109-140 mm ) 
(M. 119.3mm); hind feet 33-39. 3mm (M. 35.3mm). 

ADDITIONAL DESCRIPTION. Color much lighter than that of the 
common flying squirrel, more silvery, with a less marked straw tinge 
(color of back pale mouse -gray, Ridgway, 1912, pi. LI). 

Hair on back 15- 17 mm long. Color as follows: l) roots deep gray 
(neutral gray), broad; 2) tips of hairs short, very pale gray- straw -white. 
Head, parachuting membranes and back of the same color. 



271 



Dorsal side of tail dull whitish gray (between pale mouse -gray and pale 
drab- gray) with a brownish black tinge caused by the dark tips of some 
hairs. 

Belly white, without any yellow shade. Roots of hair on belly pale gray. 
311 Summer fur of adults pale dirty gray on back with a straw tinge (between 
drab gray, light drab and pale mouse gray). Top of head and occiput 
whitish gray. Anterior part of muzzle, cheeks and area behind the ears 
with an admixture of white. Straw tones of back best developed in lower 
part. Parachuting membranes with more intensely colored deep mouse- 
gray hair roots. This ground color appears lighter because of the silver- 
straw hair tips. 

Dorsal side of tail whitish, with a slight straw tinge in one specimen. 
Middle of ventral side of tail with a marked gray tinge on the whitish ground 
color. 

Feet whitish. Ventral side of feet white, sometimes with a slight straw 
tinge. Hair roots pale gray. 

Color of young duller and more gray. Tail with brown- gray hair tips 
through which the whitish roots of the hairs show. 

SYSTEMATIC NOTES. P . v . gubar i and P . v . volans show marked 
differences. 

The West Siberian flying squirrel differs from P. v. betulinus 
Serebr. in its winter fur. The gray color of P . v . b e t u 1 i nu s is much 
purer and its tail is whitish gray without any straw tinge. 

The winter fur of the West Siberian flying squirrel differs from that of 
P.v.incanus as follows: l) hair shorter and less luxuriant; 2) tail 
without straw-rust tones. Its summer fur differs from that of betulinus 
as follows: l) pale yellow color of back much less well developed; 2) 
tail whitish without straw-rust tinge. Fur of young P.v, gubari also 
grayer than that of betulinus . 

GEOGRAPHICAL DISTRIBUTION. A large area in western Siberia 
from the birch groves of the Shelyabinsk District, to the Baraba Region, 
Novosibirsk, Barnaul and Biisk in the east. Its northern boundary is not 
known exactly. Specimens from the Narym District and from the Chulyma 
river possibly belong to this subspecies. 

It is not known which subspecies occurs in the north of the Tobol'sk 
Territory. 

A pelt from the Vershinskie area near the Kazyma river is in the ZMAN 
collection (collected by M.I. Shukhov on 1 April 1914). Color of this 
specimen rather intense, very similar to that of Altai specimens. 

353. Pteromys volans betulinus Serebr. (1929). Semipalatinsk 
flying squirrel 

1929. Pteromys volans be t uli nus Serebrennikov M. K. , Ein Flatterh8mchen aus dem Kirgisengebiet, 

Ztschr. f. Saugetierk. Bd. 4, S. 142-143. Taf. XIII. 
312 ^53^- Pteromys volans Belyaev A. M. , Spisok gryzunov Kazakhstana (A List of Rodents of 

Kazakhstan), Trudy zoologicheskogo instituta, 2 (1) : 32 ("Ribbon" Pine Forests of the Semipalatinsk 

Province, Birch Groves in the Former Pavlodar County). 
1934. Pteromys volans be 1 u 1 inus Ognev S. I. , Obzor letyag, vodyashchikhsya v SSSR (A Survey of 

Flying Squirrels in the U. S, S. R. ), Byulleten' Obshchestva ispytatelei prirody, otdel Biologii, 10, 

3 (2): 305-306. 

TYPE LOCALITY AND DEPOSITION. Tosta, Pavlodar County, Semipalatinsk Province (200km north 
of Pavlodar). Type No. 18013, male, 18 Aug. 1927, collected by A. Dobrinin. ZMAN collection. 

MATERIAL EXAMINED: Holotype and 22 paratypes. 

272 



DIAGNOSIS. Winter fur on back light silver-gray with a clear straw- 
yellow tinge (between drab gray and pale mouse -gray). Dorsal side of 
tail with a yellow-straw-rust tinge (more yellowish than avellaneous, 
Ridgway, 1912). Summer fur with well marked dull straw-yellow tinge. 
Dorsal side of tail straw-yellow with a slight rust tinge*. 

Length of skull 39.2-41.3 mm (M. 40.4 mm); condylobasal length 35- 
37.3mm (M. 36.58mm); zygomatic width 22. 9-25. 4mm (M. 24.07mm); 
length of nasals 12-13.1 mm (M. 12.5mm); length of upper molar row 
7-7. 3mm (M. 7.19mm). 

Length of body 116-172 mm (M. 157.1mm); tail without terminal hair 
98-119mm (M. 113mm) (l22accordingtoM.K. Serebrennikov); hind feet 
without claws 34-39 mm (M. 35.8 mm). 

ADDITIONAL DESCRIPTION. Winter fur. Back light silver -gray 
with a well marked yellow -straw tinge (between drab -gray and pale mouse - 
gray). 

Color very similar to that of P. v. athene Thos. (see below) but more 
yellowish. Fur of P. v. athene from Shantar and Yakut more pinkish. 
Top of head and back of the same color. 

Hair on parachute long, more silvery, with less marked straw tinge. 
Dorsal side of tail straw-yellow rust colored (between pinkish buff and 
avellaneous). 

The black-brown tips of hairs contrast clearly with the ground color, 
especially in the terminal third on the dorsal side. These hair tips give a 
grayish tinge to the ground color. 

A broad medium whitish gray stripe on the ventral side of the tail, 
lateral parts straw-gray. 

Dorsal side of feet with mixed gray and whitish silver hair. Toes with 
short straw-white fur. Soles, especially the inner parts, with dense 
whitish hair. 

Belly white. Sides of belly in some specimens with a slight pale straw 
tone. 

In adults, summer fur on back with marked straw -yellow tone caused by 
hair tips of this color. Color of summer specimens of P. v. betulinus 
very similar to that of P . v . athene from Sakhalin. General color of 
summer fur of P. v. betulinus lighter and more yellow. Dorsal side 
of tail pale straw -yellow. 

Summer fur of young duller. Yellowish tone on top of head and 
parachuting membranes paler than in adults, without reddish and rust tones. 

Yellowish tone of tips of hair on back absent in some specimens and 
color of back closely resembling mouse-gray with a slight gray-straw tinge. 

SYSTEMATIC NOTES. The ranges of P . v . b e t u li nu s and P . v . 
gubari are adjacent. The color of P . v . b e t u 1 i nu s closely resembles 
that of P. V. athene. The winter fur of the Semipalatinsk flying squirrel 
differs from that of athene as follows: l) more yellowish shade mixed 
with the silver -gray ground color, as against the pinkish straw shade of the 
winter fur of some specimens of P . v . at he ne ; 2) fur of back of P . v . 
betulinus not so luxuriant and soft as in P. v. athene; 3) tail duller 
and more yellow. 

Summer fur of P. v. betulinus similar to that of P. v. athene, but 
paler and duller. 



This diagnosis does not agree with that given by M. K. Serebrennikov (see below). 



273 



314 



M.K. Serebrennikov (1929) certainly described a partly albinistic 
specimen ofP.v.betulinus. 

The summer fur of all forms ofPteromys is darker than the winter 
fur as a rule. In the holotype ofP.v.betulinus (which has a complete 
summer fur) the dark gray pigment is very weak, but the straw-rust-yellow 
color is partly preserved (especially on the back and on the tip of the tail). 
The gray color is better preserved on the dorsal surface of the parachuting 
membranes and on the haunches and shanks of the hind legs. The feet are 
white. The whole underparts are also white, without any gray hair roots. 

This distribution of colors results in a very light whitish gray color of 
the back, with a straw -rust -yellow tinge on the back. 

The fact that M.K. Serebrennikov examined three very pale specimens is 
no proof that there is a whitish geographical form. It should be stressed that 
these three specimens were all caught in August 1927 in the same locality. 
There seems no doubt that they belong to the same family of partly albinistic 
specimens. M.K. Serebrennikov states (1929) that P . v . be t u 1 i nu s has 
a larger skull and body than the European and Sakhalin forms. Our data 
(see measurements in Tables 33 and 34) do not support this conclusion. 

GEOGRAPHICAL DISTRIBUTION. Birch groves of the forest steppe 
north and west of Pavlodar. 

The same subspecies occurs in ribbon pine forests in the south of the 
former Semipalatinsk Region, e.g. , in the Srostinsk Pine Forest (former 
Zmeinogorsk County). It is not known how far east this pale form occurs. 
It occupies the southern outskirts of the range ofPteromys in western 
Siberia. North of Barnaul and Biisk it is replaced by a grayer and very 
pale form, P. v. gubari. 

354, Pteromys volans turovi Ogn. (1929), Central Siberian flying 
squirrel 

1858. Pteromys volans Schrenck L. , Reisen und Forschungen im Amur-Lande, I, S. 1 16- 1 18; Maak R. , 
Puteshestvle na Amure (A Journey Along the Amur), Sankt Petersburg, 1859, Obzorzhivotnykh( A Survey of 
Animals), pp. 103-104, Radde G. , Reisen im Siiden von Ost-Sibirien, 1862, S. 131-132. 

1922. Sciuropterus volans var. sibiricus Dybowski B. , Arch. Tow. Nauk. Lwowie, III, p. 8 
(Dauria, Amur). 

1926. Sciuropterus russicus Doppel'maier G. G. , Sobolinyi promysel na poberezh'e Baikala (The 
Sable Trade on the Shore of Lake Baikal), p. 38. 

1929. Pteromys volans turovi Ognev S.J. , Uber einige Nagetiere Ostsibiriens, Zoologischer 

Anzeiger, Bd. 83, H. V4, S. 75; Ognev S. I. , Mlekopitayushchie Shantarskikh ostrovov (Mammals of 

the Shantar Islands), Izvestiya Tikhookeanskoi nauchno-promyslovoi stantsii, 2 (5) : 1929, p. 1. 

TYPE LOCAUTY AND DEPOSITION. Bol'shie Koty (near Lake Baikal). Type No. M. 1241, 6 Oct. 1923, 

collected by Professor S. S. Turov. S.I. Ognev 's collection. 

MATERIAL EXAMINED. Holotype, 19 specimens from Transbaikalia and 33 specimens from various 

localities in eastern Siberia. 

DIAGNOSIS. Winter fur dark, dirty straw-gray with marked black 
mottling. Ground color of dorsal side of tail dull straw-gray or dull gray 
(without any straw tinge) with well marked black-brown hair tips giving 
a dark color to the tail. 

Back of summer specimens gray-straw-rust-colored. Rust tones 
sometimes very weak. This color appears darker because of dark gray 
hair roots on the back. Color of dorsal side of tail from very dull straw- 
yellow-gray (more yellow and duller than drab, Ridgway, 1912) almost 



274 



315 



without black hair tips, to yellow-rust-red (near cinnamon, Ridgway, 
1912) with a black-brown tinge caused by such hair tips. 

Length of skull 38-41 mm (M. 40.3 mm); condylobasal length 31.2-37.6 mm 
(M. 35.8 mm); zygomatic width 22.2-24.7 mm (M. 23.68 mm); length of 
nasals 12-13 mm (M. 12.39 mm); length of upper tooth row 7-7.7 mm 
(M. 7.20mm). Length of body 170-172 mm; tail 98-110 mm (M. 103.5 mm); 
hind paws 30-36. 2mm (M. 33.0mm). 

ADDITIONAL DESCRIPTION. Winter fur generally of dark, dirty 
grayish straw color with marked black-gray mottling caused by black 
guard hair and the black-gray tips of many hairs on the back; general color 
close to mouse -gray with a drab tinge. 

Top of head usually lighter with a straw tinge. Small patches of whitish 
fur behind the ears. 

Dorsal side of tail dull straw -gray or dull gray with a marked dark 
shade caused by many black or black-brown hair tips. Roots of hair on 
tail deep dark gray (deep neutral gray), with straw-gray subapical parts 
and black-brown tips. Ground color of ventral side of tail pale straw- gray 
covered with a black or black-brown pattern caused by dark hair tips. 

Feet gray with marked silvery mottling caused by admixture of whitish 
hair. 

In summer fur of fully grown and old specimens, ground color of back 
gray -rust -straw. Rust tones developed to a variable degree. The general 
color consists of the following elements: l) broad dark slate -gray (deep 
neutral gray) hair roots; 2) broad rust- straw colored hair tips, (between 
wood-brown and cinnamon) which are sometimes duller, straw-gray 
(between drab and buff-brown). Top of head of about the same color as the 
back, but usually more intense. 

Color of tail between dull straw-gray-yellow (more yellow and duller 
than drab) to yellow -rust -red (close to cinnamon). Black-brown hair tips 
cause a dark, well marked tinge in the ground color. Ventral side of tail 
often ash-gray, lateral parts rust colored with a black or black-brown 
tinge, while the middle of the tail remains gray. 

Dorsal side of feet straw-gray. Hair on tips of toes longer and straw- 
white. Soles dirty white, rust-reddish laterally. Two color types were 
found in series of young summer specimens. In the first, most common 
type, the back is dull straw-gray with strong dark-blue gray admixture 
caused by the color of the roots of the hair. 

Color of tail dull straw-gray (between light drab and light mouse-gray) 
with a blackish brown tinge especially in the terminal third of the tail. 
Ventral side of tail dull straw -gray with a black tinge. 

Dorsal side of feet brownish straw-gray with a light silvery tinge. 

Dorsal side of parachutes black-brown. Hair tips with a slight straw 
tinge especially along the margin of the parachutes. 

Fur on the ventral side dirty whitish, with a slight rust tinge laterally. 

Young of the second color type have reddish rust hair tips and short, 
soft fur, which is also well marked on the tail. 

Reddish rust tones mainly on underparts and flanks, contrasting with the 
white background. 
316 Color of very young specimens of the same year (specimens from the 
Teletsk Lake on the Altai, 31 August 1932) as follows: Back with a 
marked straw-gray tinge (drab). Dorsal side of parachutes with soft 
blackish brown fur. Hair of tail very short, 10-12 mm long with well 



275 



marked black hair tips, so that the tail appears completely black, especially 
in its terminal half. 

Straw -gray tone (drab) of hair roots clearly marked in basal part of tail. 
Ventral side of tail almost black, with straw-gray tone only at the base. 

Belly whitish. Hair roots gray. Sides of belly with a slight rust tinge. 

Feet with a mixture of gray and straw colored hair. Tips of toes pale 
straw colored. Soles whitish. Inner side of feet with black margins. 

SYSTEMATIC NOTES. The range of the central Siberian flying squirrel 
extends from the Altai to the Okhotsk Coast and the Amur. It differs clearly 
from the European flying squirrel in its dark, dirty straw-gray color (with 
a black tinge) in winter. Summer fur with reddish rust tones on the back 
and especially on the tail. 

These colors are typical of old summer specimens of the central 
Siberian flying squirrel but are not present in the European subspecies. 
The range of the Sakhalin flying squirrel (P . v . athene ) is adjacent to that 
ofP.v.turovi. P. V. athene is easily distinguished from P . v . 
turovi as follows: l) lighter tone of winter fur; 2) typical rust- 
yellow tinge of summer fur; 3) absence of reddish rust tone in summer 
fur, which appears in fully grown P. v. turovi. 

Specimens from the following localities are identical with typical P . v . 
turovi: upper reaches of the Bukhtarma (2 specimens, ZMAN collection); 
vicinity of Barnaul (Gebler, ZMAN collection); Altai (without exact 
indication of locality); specimens from Professor E. Eversman's 
collection (ZMAN collection). These specimens differ little from typical 
turovi in their lighter, grayer color. Their resemblance to P . v . 
turovi is so great however, that they are considered to belong to this 
geographical form. 

GEOGRAPHICAL DISTRIBUTION. P. v. turovi occurs on the Altai, 
in the Sayans, northern Mongolia (near Mondy, the ravine of the Sungur in 
northwestern Mongolia), on the Hentey, in Transbaikalia, and in the Amur 
Region to the middle reaches of the Amur (Goryun river, LakeBolon' -Odzhal, 
Khor river). 

355. Pteromys volans athene Thos, (1907). Sakhalin flying squirrel 

1861. Pteromys volans Schmidt Fr. , Bemerkungen iiber die Saugethierfauna Sud-Sachaliens, etc., 
Melang. Biol., IV, p. 107; Nikol'skii A. M. , Ostrov Sakhalin i ego fauna pozvonochnykh zhivotnykh 
(The Vertebrate Fauna of Sakhalin), 1889, pp. 122-123; Suprunenko, Pozvonochnye Sakhalina (The 
Vertebrates of Sakhalin), 1890, p. 39. 
31-7 1907. Sciuropterus russicus athene Thomas O. , The Duke of Bedford's Zoolog. Explor. in East 
Asia, IV, Proceed. Zoolog. Society, London, pp. 409-410. 

1929. Pteromys volans al uco Ognev S. J. , Uber einige Nagetiere Ostsibiriens, Zoologisch. Anz. Bd. 
83, H. 1/4, S. 71-76; Ognev S. I. , Mlekopitayushchie Shantarskikh ostrovov (Mammals of the Shantar 
Islands), Izvestiya tikhookeanskoi nauchno-Promyslovoi stantsii, 2(6):11-14. Vladivostok 1929. 
TYPE LOCALITY AND DEPOSITION. Korsakov Station on Sakhalin. Holotype (a young August 

specimen) in the British Museum. 

MATERIAL EXAMINED: 25 specimens. 

DIAGNOSIS. Winter fur with a light silver -gray tone on the back with a 
marked straw tinge. Color intermediate between drab gray and pale 
mouse-gray (Ridgway, 1912). 

Dorsal side of tail with a marked straw-rust tinge. Summer fur with 
well developed straw-yellow color. Dorsal side of tail straw-rust colored. 



276 



Length of skull 39.3-42.2 mm (M. 40.77 mm); condylobasal length 36.2- 
38.2 mm (M. 37.07 mm); zygomatic width 23-25.2 mm (M. 23.74 mm); 
length of nasals 12.3-13,7 mm (M. 12.74 mm); length of upper tooth row 
7-7. 3mm (M. 7.2mm). 

ADDITIONAL DESCRIPTION. Winter fur light silver-gray with a clear 
straw tinge (between drab gray and pale mouse -gray) on top of head and 
back. Long hair on dorsal side of parachuting membrane more silver - 
gray, with a less marked straw tinge. Dorsal side of tail with well 
marked straw-rust (avellaneous) tone sometimes with a grayish tinge caused by 
black-brown tips of hairs. Median stripe on the ventral side of tail gray, 
straw-rust-colored laterally. 

Color of hair of the median stripe on the ventral side of tail as follows: 
l) dull gray roots; 2) broad whitish bands; 3) black-brown tips. 

Dorsal side of feet with gray and silver- white hair. Soles with dense 
white hair. 

Belly white. Sides with a straw-rust tone. Roots of hair lead- gray. 

Summer fur with well marked straw-yellow color. Color of hair tips 
intermediate between avellaneous- buff- brown and clay color. Top of head 
lighter and grayer. Back sometimes with a rust tinge. 

Color of hair on back as follows: l) intense gray (deep neutral gray) or 
even darker roots; 2) straw-yellow subapical bands sometimes with a 
rust tinge. Tips of some hair brownish black. Some long, black guard 
hairs are present among the other hairs, 
318 Dorsal side of tail ye How -straw -rust (between avellaneous, wood-brown 
and buff-brown). In some specimens, dorsal side of tail darker because 
of grayish black hair tips, especially on terminal part of tail. Ventral side 
of tail duller, whitish straw-yellow with well developed black hair tips 
which form an irregular pattern in the middle and terminal parts. 

Dorsal side of feet with gray and straw colored hair. Soles with dense 
whitish hair. 

Color of underparts variable. In some specinaens it is white with a straw - 
rust tinge at the sides. In other specimens nearly the whole belly has a 
straw -rust tinge (between pinkish buff and cinnamon -buff). 

Color of young in summer generally dull. Anterior part of back and 
parachutes without reddish and rust color. General color dull, buff- 
brown, darkened by long guard hairs with black or gray-black tips. 

Dorsal side of tail dull yellowish gray (between drab and buff-brown) 
with well marked gray-black hair tips, especially in distal third of tail. 

Belly, especially at the sides, dirty white with rust-ocher shades. 

SYSTEMATIC NOTES. The winter fur differs from that of P.v,volans 
L. as follows: l) lighter straw shade on back; 2) marked straw-rust 
tones on dorsal side of tail. 

The color of the tail of six Sakhalin specimens in the ZMAN collection 
shows considerable variation. In two specimens the yellow-straw tinge of 
the tail is very weak and is replaced by gray tones. 

The differences between P,v,athene, P.v.ognevi and P . v . 
turovi have been described above. The color of the winter fur of P . v. 
athe ne closely resembles that of P. v. betulinus. 

In conclusion, the confused taxonomy of the Japanese flying squirrels 
will be discussed. Most authors who have dealt with this question had only 
limited material, which has produced confusion. For example, Nogamichi 
Kuroda (On three new Mammals from Japan, Journ, of Mammalogy, Vol. 2, 



277 



319 



No. 4, 1921, pp. 208-209) described a new subspecies, Sciuropterus 
russicus orii, from six specimens from Hokkaido (Yezo). According 
to him the skull of this subspecies is larger than that of P . v . athene 
and its zygomatic arches are wider. The only data on P . v . at he ne at 
Kuroda's disposal were measurements of the single young specimen 
described by Thomas. 

Kuroda gives the following measurements for Sciuropterus 
russicus orii: length of skull 39.5 mm; basal length 30.5 mm; 
zygomatic width 24 mm; postorbital width 9 mm; interorbital width 7. 5 mm; 
width of rostrum in front of nasals 6.5mm; length of nasals 13 mm; 
diastema 8mm; length of bullae 10.5mm; length of upper tooth row 6.5mm. 

These measurements clearly show that the skull of Sciuropterus 
orii is identical with that of P. v. athene. (see measurements in 
Table 35). 

Kuroda writes that the color of S . r . o r i i is much lighter than that of 
P. V. athene. However, Kuroda used winter specimens while P . v . 
athene was described from a summer specimen. The description of 
winter fur of S. russicus orii by Kuroda agrees well with that of 
P. V. athene. There is a winter specimen from Hokkaido in the ZMAN 
collection, which has the same color as athene . Its skull is also 
identical with that of P. v. athene. S. russicus orii is therefore 
synonymous with P. v. athene. 

O. Thomas attempted to clarify the systematics of Japanese flying 
squirrels (Proceedings Zoological Society of London, 1905, pp. 344-345). 

After examination of material from the southern and central parts of 
Hondo and the paratype of Pt e romys momongaTemm. (supplied by 
the museum of Leyden) Thomas came to the conclusion that P. momonga, 
which is a small form, was described from one of the southern islands of 
Japan ("will prove to be an inhabitant of one of the southern islands"). 

O. Thomas made a new subspecies, Sciuropterus momonga 
amygdali for specimens from Hondo. These conclusions of O. Thomas 
are doubtful, not being based on exact data. Temminck (Fauna Japonica, 
1850, p. 49) does not indicate the exact type locality of his Pteromys 
(Sciuropterus) momonga. He writes: "On trouve cette esp^ce, due 
aux soins de M. Biirger, dans les for&ts de I'interieur, ou elle vit dans 
les troncs des arbres et se nourrit de noyaux et de semences". 

Finally, the status ofS. momonga as a separate species is doubtful. 
Hondo Island is probably inhabited by a subspecies of P. volans. 

GEOGRAPHICAL DISTRIBUTION. Sakhalin, Hokkaido, lower reaches 
and estuary of the Amur. The specimen from Ayan (collected by 
Voznesenskii, ZMAN collection) certainly belongs to this form. It is 
difficult to decide how far west this form occurs. However, it certainly 
occurs near Lake Petropavlovskii, 183 km from Khabarovsk. Flying 
squirrels from St. Ol'ga Bay are transitional to the Ussuri form in color. 

356. Pteromys volans incanus Miller (1918). Yakut flying squirrel 

1873. Pteromys volans Polyakov 1. , Geograficheskoe rasprostranenie zhivotnykh v yugo-vostochnoi 
chasti Lenskogo basseina (Geographical Distribution of Animals in the Southeastern Part of the Lena 
Basin), Zapiski Geograficheskogo obshchestva, III, pp. 34-35; Maak R. , Vilyuiskii okrug (The Vilyui 
District) II, Sankt Petersburg, 1856, pp. 142-143. 



1254 



278 



1898, Flying squirrels. lokhel'son V.I. , Ocherk zvero-promyshlermosti v Kolymskoin okruge (A Survey 

of the Animal Trade in the Kolyma District), Sankt Peteisburg, p. 67. 
1903. Sciuropterus russicus Allen J. A. , Report on the Mammals, collect, in Northeast Siberia, 

Bull. Amer. Mus. Nat. History, Vol. XIX, pp. 132-134. 
1918. Pteromys volans incanus Miller G.S., A New Flying Squirrel from Eastern Asia. Proceed. 

Biol. Society of Washington, Vol. 31, pp. 3, 4. 
1926. Pteromys volans incanus Ognev S. I. , Mlekopitayushchie Severo-Vostochnoi Sibiri (Mammals 

of Northeastern Sibiria), pp. 89-93 (partim.'). 

TYPE LOCAUTY AND DEPOSITION. Vicinity of Verkhnekolymsk. Type No. 200613, female, ad. , 
24 April 1915. Collected by Copley Amori. Deposited in the U. S. A. National Museum. 

MATERIAL EXAMINED: 21 specimens. 

DIAGNOSIS. Winter fur sharply different from that of P. volans L. 
in the following characters: l) lighter silver -straw-gray tone of back 
(between pale drab- gray and pallid mouse -gray); 2) straw color of tail; 
3) long, luxuriant, silky fur, especially on the tail. 

Summer fur with a straw tone on a dull background on the back and with 
a straw-rust shade on the tail. 

Length of skull 40.1-41,6 mm (M. 41.01 mm); condylobasal length 37.1- 
38.7 mm (M. 37.83 mm); zygomatic width 24,6-25,6 mm (M, 25,03 mm); 
length of nasals 11.1-13.6 mm (M, 12.73 mm); length of upper tooth row 
7. 2-7. 7mm (M.7.4mm). 

Length of body 161-180 mm; tail 100-141 mm; hind feet 31-38 mm. 

DESCRIPTION. Winter fur resembling that of P . v . gub ar i but with 
a more marked straw tone (between pale drab-gray and pallid mouse-gray). 
Top of head, back and dorsal side of parachuting membrane of the same 
color. 

Hair on back very long, up to 24-26 mm. Color as follows: l) roots 
broad, gray (between neutral gray and deep neutral gray); 2) tips pale 
whitish gray with a slight straw tinge. Some hair with blackish tips 
producing a grayish tinge. 

Tail very luxuriant. Color of dorsal side pale straw-gray (drab and 
light-drab), darkened by blackish tips of hairs. 

Straw (drab) shade on tail more marked than in P . v , gubar i . 
Ventral side of tail whitish straw-gray or ash-gray with an irregular 
blackish brown pattern in the middle and terminal parts of tail caused by 
broad black or black-brown hair tips. 

Dorsal side of feet silver -ash -white. Soles with unusually dense and 
long white hair. 

Underparts white with a slight yellowish tinge. Roots of hair of 
underparts gray. 

SUMMER FUR. Underparts mouse -gray with a pale straw -gray (drab) 
tinge. Top of head, forehead and occiput like underparts. More marked 
straw tones on neck. Sides of neck and region behind the ears with two 
white stripes the color of which passes gradually into the color of the 
cheeks. 
321 Hair on back up to 20mm long. Color as follows: l) roots broad deep 
gray (between deep m.ouse-gray and deep neutral gray); 2) well marked, 
wide, whitish straw-gray, subapical bands; 3) tips of most hairs blackish 
gray. Tips of some hairs straw colored (drab). Hair on the sides of the 
back and on the dorsal side of the parachuting membranes darker and 
their roots more intensely gray (dark neutral gray). 

Light colored bands on the hair of the parachuting membranes little 
developed. The color of the parachuting membrane is therefore very dark. 



279 




00 

O S" 



o ■^ 



<; o. 



280 



Throat, chest and belly white with a slight yellowish tinge. 

Hair on tail up to 33 mm long, with mixed whitish, straw-rust and 
blackish tones. 

Feet with whitish and brownish black hair. 

SYSTEMATIC NOTES. The winter fur of this form resembles that of 
P . gubari but differs from it as follows: l) more marked straw tone on 
back; color of hair on back of P. v. gubari more silvery; 2) tail with 
a marked straw tone, which is absent in specimens of gubari examined; 
3) fur much longer and more luxuriant. 

The summer fur of P . V . inc anus differs from that of gub a r i as 
follows: l) straw tones on back less developed; 2) fur duller and longer. 

1 examined a large series of P. v. incanus from various localities in 
Yakutia. Their color was rather uniform. However, the extent of the 
straw tinge on back and tail varied. 

The color of specimens from the Anadyr Territory and the Chukchi 
Peninsula, differs from that of P. v. incanus in some respects. 

GEOGRAPHICAL DISTRIBUTION. The subspecies occurs in the 
valley of the Kolyma as far as Nizhne-Kolymsk (I have specimens from the 
locality in my collection), in the Verkhoyansk Territory, the Lena and 
Aldan valleys. Specimens from the lower Tunguska also belong to this 
subspecies. The western boundary of the range ofP. v. incanus is not 
clear. It is not known which subspecies occurs in the northern Yenisei 
Territory. In the south, the range of P. v. incanus is adjacent to that 
ofP.v.turovi, in the southeast to that ofP.v.athene, in the 
northeast to that of P. v. anadyrensis. The exact boundaries of the 
ranges of these subspecies will have to be determined in the future. 

357. Pteromys volans anadyrensis subsp. nov. Anadyr flying 
squirrel 

1926. Pteromys volans incanus Ognev S. I. , Mlekopitayushchie severo-vostochnoi Sibiri (Mammals 
of NortheastemSiberia), pp. 89-93 (partim.'); Ognev S. I. , Obzor letyag, vodyashchikhsya v SSSR (A 
Survey of the Flying Squirrels of the U. S. S. R. ), Byulleten' Moskovskogo obshchestva ispytatelei prirody, 
otdel Biologii, 10, 3 (2):308-309. 1934 (partim.'). 
oop 1927. Pteromys volans inc anus Sokol'nikov N. P. , Okhotnich'i i promyslovye zveri Anadyrskogo 
kraya (Game and Commercial Animals of the Anadyr Territory), Byulleten' Moskovskogo obshchestva 
ispytatelei prirody, otdel Biologii, Vol. XXXVl, p. 136. 

TYPE LOCALITY AND DEPOSITION. Eropol Village, Anadyr Territory. Type: No. 798, male, 
February 1932, L. A. Portenko's collection. Type in the collection of the All Union Arctic Institute. 
MATERIAL EXAMINED: holotype and 13 specimens. 

DIAGNOSIS. Winter fur differs from that of P. volans volans as 
follows: 1) lighter and purer bluish gray color of back (between pale 
neutral gray and dull gray, plate LIII, Ridgway, 1912); 2) usually luxuriant 
and silky winter fur; 3) hair of tail very long; 4) skull larger and molar 
row longer than in P. v. volans. P. v. anadyrensis differs from 
P. V. incanus as follows: purer bluish silver-gray color of back; straw 
tinge on back and tail of most specimens absent. 

Length of skull 41.2-42.2 mm; condylobasal length 38-38.3 mm; 
zygomatic width 25.3-25.7 mm; length of nasals 13-14 mm; length of upper 
tooth row 7.5-8 mm. 



281 



323 



Length of body 170-207 mm; tail 95-1 10 mm; hind feet 35-37 mm. 

ADDITIONAL DESCRIPTION. In winter fur, back pure bluish gray 
(between pale neutral gray and dull gray). Specimens collected by L. A. 
Portenko's expedition to the Chukchi Peninsula (on behalf of the All -Union 
Arctic Institute) were of this color. In specimens collected by N. P. 
Sokol'nikov, back duller, of a more silvery shade with a light straw tinge. 
Color of hairs on back as follows: l) roots deep gray, near neutral gray 
2) subapical parts pale whitish gray; 3) tips of hair dark gray. Top of 
head and upper part of back with hair of the color described. 

Tail very luxuriant, dull gray with a light straw-white tinge, without 
the straw-rust tone which is typical of P. v. incanus. 

Tail generally dull straw-gray. This color is caused by hair with pale 
whitish gray roots and appears dark because of blackish brown hair, or hair with 
broad blackish brown tips. 

Ventral side of tail whitish gray, darkened by many blackish hair tips. 

Belly white, without any yellow tones. Roots of hair on belly gray 
(neutral gray). 

SYSTEMATIC NOTES. The description of P . v . i n c a nu s byG.S. 
Miller (1918) agrees well with the color characters observed. G.S. 
Miller states that the color of the hair on the back of P. v. incanus is pale 
whitish gray with a buff tinge: "On the upper parts and sides this is 
overlaid by a whitish gray with a barely perceptible tinge of buff, the two 
colors blending into a general effect that is near the pallid mouse -gray of 
Ridgway. . . 

"Tail of the same color as back but with a buff tinge at the sides and at 
the tip above. . . " 

The Anadyr flying squirrel thus differs from the P. v. incanus in the 
absence of the buff tinge on the back and in the darker color of the upper 
part of the body. 

P. v. incanus was described from a specimen from Verkhnekolymsk. 
It is intermediate between the more buffy and rust tailed P. v. incanus 
and the pale s ilve ryP.v. anadyrensis. 

GEOGRAPHICAL DISTRIBUTION. Larch forests along the Anadyr and 
Maine rivers in the Anadyr Territory. 

358. Pteromys volans arsenjevi Ogn. (1934). Ussuri flying 
squirrel 

1870. Pteromys volans Przheval 'skii N. M. , Puteshestvie v Ussuriiskom krae (A Journey to the 

Ussuri Territory), Sankt Peterburg, pp. 254-255. 
1923. Sciuropterus russicus Sowerby A. de C. , The Naturalist in Manchuria, Vol. II-III, 

pp. 144-145. 
1934. Pteromys volans arsenjevi OgnevS.I. , Obzor letyag, vodyashchikhsya v SSSR (A Survey of 

the Flying Squirrels in the U. S. S. R. ). Byulleten' Moskovskogo obschestva ispytatelei prirody, otdel 

Biologii, 10, 3(4): 30. 

TYPE LOCALITY AND DEPOSITION. Sources of Kulume river, Ussuri Territory. Type collected on 
16 November 1929 by V. K. Arsen'ev. 

MATERIAL EXAMINED: holotype and 5 paratypes. 

DIAGNOSIS. Winter fur dirty buff- gray. Yellowish brown shades well 
marked on midline of back, clearly contrasting with the ground color. 
Back mouse-gray. Middle of back hair-brown with marked buff-brown 
and olive -brown tinges. 



282 



Ground color of tail yellowish buff-brown (between buff-brown and clay 
color). In summer fur, color of back gray-rust-buff. Tail rust-red 
(cinnamon). Summer fur of young yellowish brown (between buff-brown and 
sayal brown). Tail of a purer yellow tinge. 

Length of skull (from one old specimen) 38.3 mm; condylobasal length 
36 mm; length of nasals 12.1; length of upper tooth row 7. 2mm. 

ADDITIONAL DESCRIPTION. The characteristic color of the back is 
described above. Yellowish brown (buff-brown) tones well marked on top 
of head. Parachutes dor sally gray, with a weaker buff tinge than the back. 

Ventral side of tail with a marked grayish tinge darkened by black hair 
tips. 

Color of young summer specimens from the vicinity of the Razdol' 
Station of the Ussuri Railway yellowish brown-buff (between buff-brown and 
sayal brown). Tail of a purer yellowish shade than in adults. 

SYSTEMATIC NOTES. The winter fur of the Ussuri flying squirrel has 
a very typical color which resembles that of P. buchneri Satun. from 
Han-Su, in the presence of yellowish brown shades in its summer fur. 

I previously considered the Maritime Territory and Amur flying squirrels as 
P. v. aluco Thos. (Sciuropterus aluco). This subspecies was 
described from a specimen caught in the vicinity of Kaloguai, 55 miles 
northeast of Seoul (Korea) on 25 November 1906. (Duke of Bedford's 
Zoological Exploration in Eastern Asia, V. Second list of Mammals from 
Korea, Proceed. Zoolog. Society, London, 1907, 464-465), The holotype 
is in the British Museum (No. 76318). 

After examination and comparison of the characters of the Korean, 
Maritime Territory and Amur flying squirrels I came to the conclusion that they 
do not belong to the same subspecies. Thomas states that the Korean flying 
squirrel, P. v. aluco Thos,, is small. The measurements of the female 
are as follows: length of body 157 mm; tail 149 mm; hind feet 35 mm; 
ear 19.5 mm. 

Length of skull 39mm; basal length 29.5 mm; maximal width of skull 
(apparently measured at the level of the zygomatic arches) 23 mm; nasals 
12.2X5; interorbital space 7.5mm; width of braincase 17mm; length of 
palate 16.2 nam; length of palatine foramina 5.5 mm; length of bulae osseae 
10.2 mm; length of row of teeth (without p^) 6.5 mm; greatest width of m^ 
1 .8 mm. 

In Thomas's opinion P. v, aluco is smaller than P. v. volans, but 
this statement seems of little value as only one specimen was examined. 

A comparison of the Maritime Territory and Korean flying squirrels can not 
be based on skull structure or measurements of skull and body, owing to lack 
of material. However, an analysis of the color characters gives some 
results, 

Thomas describes the color of the Korean flying squirrel from one 
specimen collected in late November as follows: "General color pale 
buff-drab, not so strong as in momonga nor of such a clear gray as in 
russicus. Upper surface of hands and feet gray, the light hairs on the 
toes cream colour, more extended proximally than in the allied forms. 
Tail pinkish buff (very pale yellowish with a light reddish tinge), the 
intermixed black hairs of the upper layer unusually few in number and 
therefore not hiding the buff. " 

Thomas does not indicate whether the holotype was in summer or 
winter fur. However, the description of the color and the date of collection 
indicate clearly that the specimen was in winter fur. 

283 



325 



The Korean flying squirrel differs from the Ussuri subspecies in its 
very light tail, the color of which is pale yellowish buff- drab, almost 
without black hair. 

A.S. Lukashkin sent me an interesting male specimen from the 
Dzhandungol, a tributary of the Khaidarkhe (Argun'), on the western slopes 
of the Great Khingan (28 December 1934). The specimen has a light 
colored winter fur with buff tones. It is much lighter than P. v. turovi 
and differs clearly from P.v.arsenjevi in its small size. In some 
respects it resembles P. v. aluco Thos. in its color. 

Length of body 162mm; tail (without terminal hair) 103 mm; hind feet 
30 mm; ear 16 mm. 

Length of skull 38.2 mm; condylobasal length 34.7 mm; zygomatic width 
24.2 mm; width of skull at level of auditory openings 20.2 mm. The 
specimen was killed near a hollow in a larch tree. 

GEOGRAPHICAL DISTRIBUTION. Not clear. The subspecies 
apparently occurs in the Ussuri and Sungazi basins, and on the Posyet 
Peninsula. I examined specimens from the following localities: l) No. 
3796, February 1883, Posyet Peninsula. N. Yankovskii, ZMAN collection; 
2) 16 November 1929, upper sources of the Kulume, Ussuri Territory, 
V.K. Arsen'ev ZMAN collection; 3) 14 November 1929, same locality; 
4) No. 20645, 14 November 1929, same locality; 5) vicinity of the Razdol' 
Station of the Ussuri Railroad, 1928, collected by A. N. Formozov; this 
summer specimen was found, according to its label, in February; the date 
is apparently wrong; 6) No. M. 1697, female, September 1926, Okeanskaya 
Station, Maritime Territory, E.M. Vermel' (S.I. Ognev's collection). 

Flying squirrels from the Chinese provinces Shansi (western part) and 
Hopei (northeast of Peiping) mentioned by A. de Sowerby (The 
Naturalist in Manchuria, II— III, 1923, pp. 144- 145), possibly belongto this 
subspecies. 



Family SCIURIDAE 

1821. Sciuridae Gray, I. London Medic. Repos. , XV, 304, Apr. 1. 

Body slender or thickset, adapted to arboreal or terrestrial life. 

Tail always without horny scales, densely covered with hair, often 
dorsoventrally depressed. Tail of arboreal forms with long hair. 

Structure of skull varying, but never adapted to underground life. 
Infraorbital canal long and compressed. Infraorbital foramen very sinall. 
Frontals with well developed postorbital processes. Bony palate wide. 

p -^; m ' ' • Molars with short crowns and long roots. Testes inside 

scrotum. Os penis present. 

GEOGRAPHICAL DISTRIBUTION. Western and eastern hemispheres 
except for the extreme north and south. Not present in Madagascar, New 
Guinea, Australia or the Pacific islands. 

The family includes the following subfamilies*: 



* R, Pocock, The Classification of the Sciuridae, Proceed. Zoolog. Soc. London, 1923, parti, pp. 209-246. 



284 



1) Sciurinae. The subfamily includes the genus S c iur us , subgenus 
Tenes ( = Oreosciurus) and the American subgenera Parasciurus, 
Ar ae o s c iuru s , Neosciurus, He s pe r o s c iur u s , Otosciurus, 
Distribution: Palearctic, Nearctic and Neotropical regions. 

2) Tamiasciurinae with the genus Tamiasciurus - Nearctic Region. 

3 ) Funambulinae with the genera: Funambulus, Tamiodes, 
Ratufa, Protoxerus, Aetosciurus, Funisciurus, 
Paraxerus, Heliosciurus, and possibly also M yo s c iu r u s . 
Distribution: Ethiopian and Oriental Region. 

4) Callosciurinae with the genera: Callosciurus, Menetes, 
Tomeutes, Rhino s c iuru s , Lariscus, Dremomys, 
Tamiops, Nannosciurus. Distribution: Oriental Region from India 
to China and Borneo. 

5) Xerinae with the genera: Atlantoxerus, Xerus, Euxerus, 
Geosciurus. Distribution: Africa from Morocco to the Cape Province. 

6 ) Spermophilopsinae with the genus Spermophilopsis . Distribution: 
Transcaspian Desert, Iran and some parts of Turkestan. 

7 ) Marmotinae with the genera: Marmota, Marmotops, 
Otospermophilus, Callospermophilus, Citellus (Subgenera: 
Urocitellus, Colobotis); genera: Ammospermophilus, 
Cynomys (subgenera: Cynomys s. str. and Leucocrossuromys), 
Eutamias, Tamias. Distribution: Palearctic and Nearctic regions. 



KEY TO THE SUBFAMIUES OF THE FAMILY SCIURIDAE IN THE U. S. S. R. 

1 (2). Toes unusually long and slender. Outer margins of fore- and hindlegswith stiff pectinate fringes 
of hair which may be 5 mm long. This hair is especially long on the outer side and at the tip of the 5th toe 
of the hind feet, where it forms a tuft which is longer than the claw. 

Toes with black claws; longest claw usually longer than 1 cm, sometimes 1.5 cm. 
Molars lophodont, that is, the crowns have plicated enamel loops. 

Spermophilopsinae. 

2 (1). Toes of normal length. Margins of legs without pectinate hair fringe. 

Claws not particularly long. Molars bunodont, that is, tubercles on crowns with transverse elevated 
enamel folds. 

3 (4). Ears long, often with tufts. 

Tail long, luxuriant, with long hair. No cheek pouches. 

S ciurina e. 

4 (3). Ears short, rounded, forming only small folds of skin slightly protruding in some genera. 
Tail with hair of moderate length. Terminal hair of tail longer than other hair. 

Cheek pouches present, sometimes little developed. 

Marmotinae . 



327 Subfamily SCIURINAE, SQUIRRELS 

Arboreal animals which build their nests on trees or in hollows. Fur on 
back of one color. Ears long, often with tufts. 

Tail long, luxuriant, with dense and long hair. Forelegs with four toes 
and hind legs with 5 toes. Fourth digit longest on both fore- and hind legs. 



285 



328 



Claws short, curved, adapted for clinging to the bark of trees. No cheek 
pouches. 

Postorbital processes of frontals well developed. 

Crowns of molars quadrangular, brachydont. Tubercles on molar teeth 
form transverse, elevated enamel folds. 

Dental formula: p — ; m ^' '^ . 
4 1. 2. 3 

Form of glans penis varying. It is usually widened anteriorly and its tip 
inclined backwards as a blunt process (Sciurus)or a pointed process 
(Callosciurus), 

Os penis large, cylindrical, with widened base and flattened tip, which 
forms a rounded plate the upper tip of which is inclined to the right. 
Anteroventral tip of os penis with a pointed, downward-directed process. 

GEOGRAPHICAL DISTRIBUTION and genera of this subfamily are 
given above. 



Genus Sciurus Linnaeus (1758) 

1758. Sciurus Linnaeus C. , System a Naturae, I. editio 10, p. 63. 

1893. Aphrontis Schultze E. , Ztschr. f. Naturwissensch. , Leipzig, LXVI, S. 165 (subgenus of the species 
Sciurus vulgaris), (sic!) 

Mainly arboreal animals which climb well on trees, run and jump on 
branches and build nests in trees. 

Back of one color without stripes. Tail (without terminal hair) 1/4 - 
l/5 shorter than body. 

Hair on tail very long, soft and directed laterally in ventral view. 

Ears long, at least twice as long as broad. Hair on tips of ears very long, 
forming well marked tufts in winter specimens. Cheek pouches not 
developed. 

Vibrissae arranged as follows: l) the longest and thickest vibrissae 
between eye and nose; 2) above eyes; 3) on cheeks; 4) one single 
vibrissa on the chin; 5) on the forelegs, on the outer side of the 
antebrachium; 6) 2 (very rarely 3) pairs on the belly. 

Teats: p 1-1; a 2-2; i 1-1; total 8. 

Premaxillary frontal processes widened, projecting at the sides of the 
nasals like wings. Width of posterior parts of premaxillary frontal process 
equal (rarely a little narrower, e.g., in S. aberti), and often greater 
(e.g. inS.carolinensis) than width of basal part of both nasals . 

Coronal suture rounded (arcuate) or nearly rectangular, occasionally 
with two oblique, rounded processes, backwards -directed laterally. 
Subgenus T e ne s characterized by sharply angular, backwards-directed 
processes at the sides of the coronal suture. 

Interorbital space very wide, 37-50% of condylobasal length. 

Frontal area of Palearctic species flattened or depressed to some extent. 
In American squirrels (S c iur us (Tamiasciurus)hudsonicus and 
Sciurus carolinensis) the frontal area is slightly convex, like that 
of chipmunks. 

Braincase short and broad, angular and irregularly rounded*. The 
anterior part of the zygomatic arch, i.e., the zygomaxillary process, 
forms a vertical plate, not depressed horizontally. 



* The braincase of the American genus Tamiasciurus resembles that of chipmunl<s. 

286 



The infraorbital foramina are very small, laterally compressed, with 
small angular processes at the base. These processes are especially well 
developed in S. vulgaris and in the American S . n i g e r . They are 
absent inTamiasciurus hudsonicus. 

Diastema of moderate length. Diastema about l/5 to l/6 longer than the 
row of upper molars. 

Type species Sciurus vulgaris Linnaeus. 

GEOGRAPHICAL DISTRIBUTION. Species of the genus Sciurus are 
widely distributed in the Palearctic, Nearctic, Neotropical and Oriental 
regions. They are not found in Australia, the Ethiopian Region or in 
Madagascar. 

Two subgenera occur in the Palearctic Region, the subgenus Sciurus 
and the subgenus Tenes (Oreosciurus). 

The following subgenera occur in North America: Tamiasciurus 
(which is in some respects transitional to the chipmunks), Baiosciurus, 
Sciurus, Gue r lin gue t us . The North American species are much 
more numerous than the Palearctic species; there are 34 species and 
102 subspecies, 

329 KEY TO THE SUBGENERA OF THE SUBFAMILY SCIURINAE IN THE U. S.S. R. 

1 (2). Color of fur changing with the season. Belly white throughout the year. Soles of hind feet with 
4 calluses. Teats 8. Two pairs of upper premolars. 

Subgenus S c iurus Linn. 

2 (1). Color of fur not changing or changing very little with the seasons. Belly various shades of 
rust-brown, or buff-yellow. Hind soles with 6 calluses. Teats 10. One pair of upper premolars. 

Subgenus Tenes Thos. 



Subgenus Sciurus Linnaeus (1758) 

Color of fur changing with the seasons. Color of winter fur sharply 
different from that of summer fur. Belly white throughout the year. 
Hind soles with 4 calluses. Teats 8. Two pairs of upper premolars. 

Type species: Sciurus vulgaris Linnaeus. 

The geographical distribution of the subgenus has been given above in the 
description of the genus, 

164. Sciurus vulgaris Linn. (1758). The squirrel 

1758. Sciurus vulgaris Linnaeus C. , Systema Naturae, X, 1758, p. 63 (Habitat in Europae Arboribus). 

NAMES: Squirrel (English); das Eichhomchen (German) ; I'ecureuil (French). 

LOCAL NAMES: belka sosnovka, elovka, berezovka (Russian hunters); belka chernaya, kedrovka, 
Shamanchik (Russian and Siberian hunters); vyvirka, bilka zvychaina (Ukrainian) ; vyvirka (Belorussian); 
wuorwantsch (Lapland); orre (Finmarken); arrew or oarre (Swedish-Finnish parts of Lapland); wanwaris 
(Lettish); orraw (Estonian) ; ur (Permyak and Cheremis-Mari); urne (Mordvinian) ; konue, kionne (Votyak) ; 
neeshs (Vogul— Mansi); leina and lengen (along the Sos'va); naksha (Chuvash- Etel); langi (Ostyak-Khanty 
along the Ob); n'aia (Ostyak-Khanty along the Kete); tabek (Ostyak-Khante along the Narym); tare, tarreku 
(Nenets); along the Yenisei - tareche; ur (Zyryan-Komi); mashi (Mescher); tiin (Bashkir) ; tiin or tikhu 
(Tatar); keermen (Kalmyk); yavunya (Koryak-Nymyllan); oloki (Orochen, Neni); kereme, kirme 
(Mongolian); kirmang (Buryat); lyrmo (Daurian); khumu (Mangun, Goldi, Nanai - along the Amur and the 



287 



Ussuri); kirmo (Kils along the Kura, Goldi and Nanei above the Ussuri, Birars, Manyagrs, Orochen- Neni, 
Daurians); ol'iki (Lamut); kirmi, khirmi (Soiot); koromon (Tungus, Evenk along the Khatanga); kirmun 
(Tungus-Evenki along the Baikal); uluki, ulyoiki (Tungus, Evenki); ulukhi, ilukhi (Manchuria) ; ulyuki 
(Manegr); khulu (Goldi, Nanei); lafkor (throughout the inner part of Sakhalin); lakr (along the western 
coast of Sakhalin); lyakr, lakr, lakrs (Gilyak); yavuny a (Koryak, Nymallan); raulen, uliki, rodlia 
(along the Anad>T); ting (Yakut); pogodzhub (Yurak, Odul); lerchieng (Tundra inhabitants, Yukagirs, 
Oduls) ; khuishu, Khui-dzhu, khui-ku-tzu [gray dog] (Chinese). 

PRINCIPAL FIGURES: 1) Schreber j. , Saugethiere, Erlangen IV, 1792, S. 760, Taf. CCXII (drawing 
of S . fusco-ater); 2) Bingley W. , Memoires of British Quadrupeds, 1809, p. 277, color plate (entire 
animal); 3) Nilsson S. , Illuminerade figurer till Scandinaviens Fauna, I. 1832, pi. 11 fentire animal 
in color — winter fur); 4) Simashko Yu. , Russkaya fauna. II. Mlekopitayushchie (Russian Fauna. II. 
330 Mammals), 1851, pp. 589-594, pi. 27 (skeleton, teeth); pi. 31 (entire animal in color) ; 5) Blasius J. , 

Naturgeschichte der Saugethiere Deutschl. , 1857, S. 272,274, Fig. 153, S. 271 (halftone plate), Fig. 154, 
S. 272 (entire animal); 6) Mewes W. , Holmgren A. E. , Atlas ofver Scandinaviens Daggdjur. Stockholm 
1873, pi. XII, fig. 3; 7) Brehm A. , Zhizn' zhivotnykh. II (Animal Life. II), 1874, p. 67 (half tone plate 
of entire animal); 8) Fokht K. , Mlekopitayushchie (Mammals), 1885, p. 375 (drawing of entire animal); 
9) Trouessart E. L. , Histoire naturelle de la France, Mammif^res, Paris 1885, p. Ill, fig. 52 (half tone 
plate); 10) Brehm A. E. , Zhizn' zhivotnykh. II (Animal Life. II), 1893, p. 429 (half tone plate of entire 
animal); 11) Lydekker R, A. , Hand-Book of the British Mammalia, 1895, pi. XIX (entire animal in color); 
12) Tullberg, Tycho, Ueber das System der Nagethiere, Upsala 1899, Taf. XXI, Fig. 1-7 (skull); Taf. XXIV, 
Fig. 37 (auditory ossicles); Taf. XXVII, Fig. 21-22 (teeth); Taf. XXX, Fig. 33 (scapula); Taf. XXXIII, Figs. 9-10 
(pelvis); Taf. XXXIV, Fig. 25 (skeleton of legs); Taf. XXXV, Fig. 16 (the same); Taf. XXXVI, Fig. 29 
(palate); Taf. XXXVIII, Fig. 16- 18 (tongue); Taf. XXXIX, Fig. 48-49 (hydrobranchial apparatus) ; Taf. XU, 
Fig. 33 (Lungs) ; Taf. XL, Fig. 33 (stomach); Taf. XLVI, Figs. 4-5 (intestine); Taf. LI, Figs. 6-11 (genitalia); 
Taf.LIII, Figs. 15-17 (same); Taf. LVII , Figs. 5-6 (soles of feet); 13) Gaake V. , Zhivotnyi mir (The 
Animal World), 1901, Vol. I, pi. 4 {entire animal in color); 14) Millais J. G. , The Mammals of Great Britain 
and Ireland, London 1906, Vol. Ill, pi. (entire animal in color); 15) Meerwarth H. , Lebensbilder aus der 
Tierwelt, 1, Leipzig 1908, S. 249-250 (photograph of animal); 16) Schaff E. , Die wildlebenden 
Saugetiere Deutschlands, 1911, S. 71-74, Fig. 20; S. 70 (skull), 17) Miller G. S. , Catalogue of the 
Mammals of Western Europe, London 1912, p. 900, fig. 186 (skulls); fig. 187 (structure of teeth); 18) Brehm 's 
Tierleben, Saugethiere, II, 1914, S. 538 (color plate of Siberian squirrels); 19) Thorburn Arch. , British 
Mammals, I, London 1920, pi. 24 (color plate); 20) Barrett-Hamilton Ger. and Hinton Martin A. C. , A 
History of British Mammals, London, 1921, parts XX -XXI, fig. 98, p. 687 (teeth); p. 692 (color plate of 
Sciurus leucurus: seasonal changes); pi. XXXVI, p. 700 (tails); pi. XXXVII (legs) ; p. 703, fig. 99 
(skull structure); p. 709, fig. 100 (diagram of track); 21) Migulin A. A. , Mlekopitayushchie Khar'kovskoi 
gubernii (Mammals of the Khar'kov Province), "Priroda i okhota na Ukraine" 1-2, 1924, pp. 119-125, 
fig. on page 123 (photograph of skulls); fig. on p. 124 (photograph of nest); 22) Vinogradov B. , Material 
for the system, and the morph. of the Rodents, Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, 1926, 
pi. XX, fig. 1 (skull lateral); 23) Wollebaek A. , Norges Pattedyr, Oslo 1927, p. 42, fig. 32 (skull of 
squirrel, lateral); p. 42, fig. 33 (track); pp. 43-44, figs. 34-35 (drawings of entire animal); 24) Formozov 
A. K. , Sledopvt-okhotnik (A Pathfinder-Hunter), Moscow 1928, fig. 79, p. 101 (track of squirrel); pp. 102- 
103, figs. 80-82 (gnawings of a squirrel); 25) Kuznetsov B. A. , Stroenie mekha i lin'ka belki (Fur and Molt 
in Squirrels) Izvestiya Assotsiatsii nauchno-issledovatel'skikh institutov pri Fiziko-matematicheskom fakul'tete 
Moskovskogo Gosudarstvennogo Universiteta, 1 (1-2): 207-249, 1928; Fig. 1 p. 210 (arrangement of vibrissae 
on head and foot of squirrel); Fig. 2, p. 212 (hair); Figs. 3-4, pp. 220-221 (hair); fig. 5, p. 231; fig. 6. 
p. 238 (Distribution of hair); pi. A, pp. 244-245 (diagram of spring molt) ; pi. C, pp. 246-247 (diagram of 
autumn molt); 26) Luhring Rud. , Das Haarkleid von S c iurus vulgaris und die Verteilung seiner 
Farbenvarianten in Deutschhind, Ztschr. f. Morphologie und Okologie der Tiere, B 11, Berlin 1928, S. 667- 
762, Fig.2:S. 674, Figs. 3-6; S. 675-678, Figs. 7-9; S. 680-682 (hair, magnified); Fig. 11a- 11a', S 692- 
694; Fig. 12a-12a', S, 696-698 (molt pattern); Figs. 13-22, S. 704, 705, 707, 708, 710, 711, 713, 716 
(hair); Fig. 23a- 23f (molt pattern); Fig. 24a-24g (pigmentation of hair on tail) ; S. 726-727; Fig. 25a-25cS. 
733-735 (molt pattern); Taf. XVIII-XIX (distribution of color types of squirrels in Germany, geographical maps 
with legends); 27) Serebrennikov M. K. , Vozrastnaya izmenchivos't i protsess rosta cherepa u belki (Age 
Variations and Growth of Skull in Squirrels), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, 1930, 
pp. 399-418, figs. 1, 2, 3 (skulls of animals of different ages); 28) Serebrennikov M. , On the polychromatism 
and albinism of Siberian Squirrels, Proceed. Zoolog. Society London, 1931, pp. 493-495, pi. I (color t>'pes of 
Siberian squirrels, color plate); pl. II (transitions between gray specimens and albinos); pi. Ill (photograph of 
partial albinos); 29) Argyropulo A. , Opredelitel' gryzunov Ural'skoi oblasti (Key to Rodents of the Ural 
Region), .Moskow 1931, p. 45, fig. 45 (skull of squirrel, ventral); 30) Kaverznev V. N. , Belka i belichii pro 
promyse) (Squirrel Hunting), .Moscow 1931, p. 5 (a squirrel near its nest); p. 6 (a squirrel nest); p. 14 

288 



(a swimming squirrel); p. 29 (a squirrel track); p. 32 (a wounded squirrel); p. 44 (a squirrel on a tree); 31) 
331 Vinogradov B. S. and Obolenskii S.I. , Vrednye i poleznye v sel'skom khozyaistve mlekopitayushchie (Harmful 
and Useful Mammals in Agriculture), Moscow 1932, p, 9 (entire animal); 32) Lobachev S. V. , Verkhnevyche- 
godskaya ekspeditsiya (An Expedition to Verkhnevychegod), 1932, p. 102, fig. 13; pp. 104-105, pp. 14-15^ 
p. 109, fig. 16 (genital apparatus); 33) Stakhrovskii V. G. , K izucheniyu biologii belki pri soderzhanii ee 
V vol'ere-parke (The Biology of Squirrels Kept in Open Air Cages), Zoologicheskii zhumal, 11 (1); 82-104, 
1932; p. 83, fig. 2 (photograph of squirrel); pp. 92-94, figs. 3-9 (photograph of young squirrels to compare 
their development); p. 101, figs. 12-13 (nest); p. 102, fig. 14 (diagram of molt); 34) Naumov N. P. , 
Opredelenie vozrasta obyknovennoi belki (Determination of the Age of Common Squirrels); Uchenye 
zapiski Moskovskogo gosudarstvennogo universiteta, issue II, 1934, pp. 275-291; p. 281, fig. 1 (diagram of the 
abrasion of the surface of teeth); 35) Naumov N. P. Biologiya razmnozheniya obyknovennoi belki (The 
Biology of Reproduction of the Common Squirrel), in "The Ecology of the Squirrel" (Ekologiya belki), Moscow 
1934, pp. 80-82, figs. 4-6 (molt diagram); p. 89, fig. 11 (diagram showing direction and sequence of fall 
molt); 36) Bobrinskii N. A. , Ocherki promyslovykh zverei i ptits (A Survey of Commercial Mammals and 
Birds), Moscow 1934, p. 17 (plate with biological scenes); 37) Manteifel' N. A. Easpopov M. P., IsakovYu. A., 
Lyubimov M. P. , Biologiya zaitsev i belok i ikh bolezni (Biology and Diseases of Hares and Squirrels), 
Moscow 1935, pp. 40-42, figs. 3-5 (diagram of sequence of spring and autumn molt); p. 45, fig. 6 
(population of squirrels); p. 46, fig. 7 (swollen teats in a gravid female); p. 48, fig. 8 (changes in the 
female genitalia); p. 53, fig. 9 (arrangement of vibrissae on the body of a newborn); p. 54, figs. 10-11 (young 
squirrels); p. 58, fig. 16 (structure of feet); p. 59, fig. 17 (development of ear, external morphology); 
p. 60, fig. 18 (development of incisors); p. 60, fig. 19 (young squirrels); p. 61, fig. 20 (head of an 1 8- day- 
old squirrel) ; p. 62, figs. 21-22 (40-day-old squirrels); p. 63, fig. 23 (nests of squirrels); p. 67, fig. 24 
(diagram of structure of the squirrel nest). 

MATERIAL EXAMINED: 1456 specimens. 

DIAGNOSIS. Slender animal with rounded, blunt head; a sharp 
constriction between head and body. Ears long, about two or more times 

as long as broad. Tips of ears with long hair 
which forms tufts in winter specimens. 

Tail (without terminal hair) almost as long 
as body and head. Hair on tail long, soft and 
directed sideways. 

Hind legs long (Figure 164). In summer 
specimens, area between heel and paw with 
dense hair along the sides to the middle. Foot 
bare except for heels and margins. Four 
calluses at the base of the toes. The two 
lateral calluses stronger and longer than the 
inner calluses. Forefeet with 5 calluses. 
Anterior middle callus cordiform. The two 
calluses rounded. The 2 posterior outer 
calluses rounded. Inner lateral callus larger 
than outer callus. 

Belly white. Back rust colored or black in 
summer and various gray shades in winter. 
Color of tail variable between rust-red, 
gray, brown and black. 
333 ^^M '91 J^IH Vibrissae present on head (between eyes and 

nose, on cheeks, and an unpaired vibrissa on 
the chin), on forelegs (outer side of 
antebrachium) and on belly (2, rarely 3 pairs). 
Teats: p 1-1; a 2-2; il-1; totals. 

Premaxillary frontal processes widened, 
FIGURE 164. Hind foot of wing-shaped laterally from nasals. Width of 

squirrel, Sciurus vulgaris L. frontal process posteriorly equal or greater 

Photograph by S.I. Ognev than that of both nasals in their basal part. 




289 





FIGURES 165-168. Skull of squirrel, Sciurus vulgaris ognevi Migulin, No. M. 1285 male, 12 
September 1912, Moscow County, S.I. Ognev's collection 

Drawing by Yu. A. Kostylev 



290 



334 



Coronal suture rounded or nearly straight, sometimes with 2 rounded 
posterior processes. This suture, however, has no pointed, angular 
processes characteristic ofTenes anoraalus. 

Interorbital space very wide, 40-50% of condylobasal length. Bullae 
osseae longer than wide. Upper molar row shorter than bullae osseae. 

Small upper anterior premolars present. They are less than one- 
quarter the size of the other premolars. Second upper premolar oblique 
and with a rounded crown. 

ADDITIONAL DESCRIPTION. Rostrum short. Zygomatic arches 
rounded. Interorbital space and braincase broad. 

Nasals much widened anteriorly and narrow at the base. 

Premaxillary frontal processes wing-shaped, obliquely descending along 
the sides of the nasals. Its posterior part as wide as both nasals at the 
base. 

Zygomatic arches rounded. Anterior parts of zygomatic arch slightly 
less apart than posterior parts. Postorbital processes thin, pointed and 
long, directed backwards and sideways. 

Posterior part of the coronal suture nearly straight or slightly rounded 
convex posteriorly). Two sutures run from the coronal suture to the 
postorbital processes. Posterior part of coronal suture sometimes with two 
rounded processes, not pointed and angular as in the subgenus Tenes . 

Tooth row much shorter than diastema. Bullae osseae longer than broad. 

Molars with transverse enamel folds alternating with narrow grooves, 
and with small rounded enamel processes at the inner side. 

First upper premolar very snaall and cylindrical, close to anterior 
inner oblique edge of second premolar. P^ trapezoidal, its long side 
directed outwards and its short side inwards. Inner side of occlusal 
surface of p^ with a rounded or pectinate process which forms a nearly 
continuous crest along the inner side of the tooth row with the processes of 
the other teeth. 

Outer side of p with four processes. The first and fourth are of about 
equal size, and both are markedly lower than the second process. The third 
process is very small. The second and fourth processes are connected 
with the inner process by transverse enamel crests. M^ of the same 
structure. M^ differs clearly from p^ and m^; in addition to the inner 
process it has only one well marked outer anterior process and a slanting 
and wide posterior process. The first outer process is connected with the 
inner process by a transverse crest. The other surface of m bears a 
shallow depression. 

In the lower tooth row p^ has two blunt denticles at the outer side. The 
anterior denticle is larger than the posterior and there is a very small 
process betweenthese two. The inner side of the tooth has a large anterior 
process and a second blunt process of the enamel margin. A deep round 
groove in the middle of the crown. Two rounded denticles on each side of 
m . A marked depression between them in the middle of the tooth. M^ the 
same, but with very small denticles on both sides between the 2 large 
processes. M^ with two rounded processes on both sides of the anterior 
margin, and two small intermediate processes at the sides. A well-marked 
rounded posterior (third) process at the inner margin, which is connected 
with the low inner process by an elevated edge. 



291 



335 



VARIATIONS OF SKULL AND TEETH OF SQUIRRELS DUE TO AGE. 
Two detailed papers on this subject have been written by M.K. 
Serebrennikov* and N. P. Naumov**. 

M.K. Serebrennikov divides squirrels into age groups. He describes 
in detail squirrels up to the age of 6 months, of the stages juvenis 
and subadultus. He divides these two stages into four subgroups. 
The other stages (older than 6 months) are divided into two subgroups, viz. 
adult us and senex. Serebrennikov does not give a detailed description 
of the last. 

Description of cranial characters of different ages of squirrels. 

Juvenis. Tothefirst s t a ge belong blind squirrels a few days old, 
waich feed only on mother's milk. The incisors have only just cut through and 
the molars are completely covered by mucosa. Condylobasal length 30- 
31mm; zygomatic width 21 mm. Skull oblong-oval; zygomatic arches little 
protruding; face very short compared with braincase. Bones very thin; 
fontanel nearly always present Dexween frontals and parietals. The 
separate occipitals form small, occipital condyles in the exoccipital zone. 
Interparietal, parietals and posterior half of the frontals form an almost 
ovoid surface. Nasals very short and wide. 

The mandible differs sharply from that of adults, it is elongate and its 
processes are little developed. It is 23 mm long and about 11mm high. 

The second stage differs from the first in the presence of two 
milk premolars. The rostrum is relatively longer compared with the 
occipital region. The ratio of length of the facial region to length of the 
braincase is 13:27. All bones are connected by more solid sutures than in 
the first stage. The fontanel becomes ossified. The occipital condyles 
grow considerably in size and are situated on the basioccipital. 
Condylobasal length 34 mm. 

Lower anterior milk premolars cut through. Mandible 23.8 mm long and 
11.7 mm high. 

Third stage. The first true molars appear. Condylobasal length 
38 mm. Zygomatic arches 25 mm wide, more widely apart laterally. 
Length of facial region to length of braincase = 14:30. 

Mandible growing more in height than in length. Height to length of 
mandible = 12.9:26. 
336 In the fourth stage four molars (premolars + molars) are present. 
Only the third posterior molars are absent. Condylobasal length about 
41-42mm, zygomatic width about 25-26 mm. Skull generally longer than 
wide. Facial length to length of braincase = 16.6:32. 

Height of mandible to length = 14:27. 

According to M.K. Serebrennikov the juvenis stage continues until 
the growth of the third posterior molar. The period lasting until the final 
replacement of deciduous teeth by permanent molars forms the subadult 
stage . 

Subadult. First stage. This age is easily determined by 
examination of the upper deciduous premolars. The crown of the second 
upper premolar is very small and has a laterally compressed anterior 



N. K. Serebrennikov, "Variations and the Growth of the Skull of Squirrels Due to Age" (Vozrastnaya 
izmenchivost' i protsess rosta cherepa u belki), Ezhegodnik-Zoologicheskogo muzeya Akademii Nauk, 
1930, pp. 399-418. 

N. P. Naumov, "Determination of the Age of Squirrels" (Opredelenie vozrasta belki), Uchenye zapiski 
Moskovskogo universiteta, II, 1934, pp. 275-291. 

292 



root. The anterior limit of this root is usually at the same level as the 
first small premolar. The anterior root of the second premolar usually 
reaches beyond the first premolar. The first premolar has one process and 
is very small. It is lower than the molar row and its inner side is 
inclined against the anterior root of the second premolar. The first 
premolar is loosely attached as its alveolus has only one root (Figure 169, l). 






'-^MJf^' \ 






FIGURE 169. 1 -skull of young squirrel of the first stage; 2 - skull of 
young squirrel of the fourth stage; 3 - skull of adult squirrel; 4 - molars 
of adult squirrel; 5 - molars of old squirrel (after M. K. Serebrennikov) 

The second stage of the subadult starts when the permanent molars 
begin to differentiate under the two deciduous premolars, ending when the 
first deciduous premolar falls out. The condylobasal length at this stage 
is about the same as in adults — 45.7mm. Zygomatic width about 29mm. 
Ratio of facial length to length of braincase = 20:34. 



293 



337 



The third stage of the subadult lasts frr,m the loss of the first 
deciduous premolar until the loss of the second deciduous premolar. The 
boundaries between the interparietals disappear at this stage as the sutures 
close and converging crests form on the braincase. These crests bifurcate 
anteriorly, reach the base of the postorbital processes and form a lyre- 
shaped figure. 

The period from the loss of the second premolar to the appearance of the 
permanent premolar forms the fourth stage of the subadult (Figure 
169, 2). 

The growth of the teeth and the replacement of the single deciduous 
premolar on the naandible which corresponds to the second deciduous upper 
premolar is the same as in the upper tooth row. 

The adult stage is characterized by full development of the tooth row. 
The braincase bears lyre -shaped crests in adults. These crests begin with 
closely placed ridges from the lambdoid crest and continue through the 
interparietal region. They diverge widely anteriorly and continue to the 
posterior base of the postorbital processes, forming the typical lyre-shaped 
figure. About 2/3 of the sagittal suture and the frontal suture become 
fused (Figure 169, 3). 

The skull becomes oblong. The adult skull differs markedly from that 
of the juvenis, being much straighter in profile. The juvenile skull 
in profile has its highest point at about the middle of the frontals; the adult 
skull near the coronal suture. 

The adult mandible is almost 2/3 as high as long. The juvenile 
mandible is about 1/2 (even less) as high as long. 

Senex. The old skull has frontal and sagittal sutures completely 
fused; the lyre-shaped crests are well developed, a lambdoidal crest is 
present, and all cranial bones are relatively thick. The crown of the teeth 
of old specimens is much worn off. 

N. P. Naumov studied the changes of the teeth due to age. Although the 
enamel of the crowns is worn off in a certain order, abrasion is influenced 
by different kinds of food. Solid food accelerates abrasion of the teeth. 
On the other hand, soft food like mushrooms and berries retards abrasion. 
These variable factors make it difficult to exactly determine the age of 
squirrels from the form of the teeth. 

N. P. Naumov (loc. cit, pp. 280-283) gives a good synoptic table from 
which the age of squirrels can be determined according to the structure of 
the upper molars-, (see Figure 170, p. 297). 

I. KEY TO MAIN AGE GROUPS. 

A. Upper tooth row incomplete. Permanent molars either absent or breaking through. The tooth row 
(including molars znd deciduous premolars) does not have a straight upper surface formed by crowns of the 
same size. Cranial bones very thin. Zygomatic arches narrow and hardly protruding laterally - young 
(juvenis), 2-3 months old. 

B. Upper tooth row complete but including deciduous teeth, i. e. , two first premolars. Surface of 
p2 triangular, its cross section about half as wide as m . Anterior root of p2 projecting far anteriorly, 
sometimes reaching beyond alveolus of p : subadult, up to one year old. 

C. Upper tooth row with only permanent teeth. Upper surface of p only little narrower than m . 
Occlusal surface of p trapezoidal. Anterior root of p never reaching beyond alveolus of p : adult. 



' This table has been modified and corrected in some details. 



294 



338 II. KEY TO SUBGROUPS OF SUBADULTS AND ADULTS. 

A. Tooth row not yet with level surface i. e. , p^ much lower than m . subadult less than one year old 

B. Occlusal surface of all teeth at one horizontal level (tooth row fully developed). 

a. Dentine not exposed at inner crest. All molars with "young" (sharply tuberculated) crowns. 
Outer tubercles not worn off, or little worn, so that their height is little decreased. Polished 
"mirrorlike" surfaces absent, or present only in narrow, disconnected areas. Dentine occasionally 
exposed in these areas, rarely visible on the outer tubercles and even more rarely on the transverse 
crests. Dentine not exposed along the inner pectinate tubercles: 6-12 month old squirrels 

(Figure 170, 4). 

b. Dentine exposed on inner pectinate tubercles. Dentine exposed either in small yellow or dark, 
spindle-shaped depressions, or as a continuous dentine network on all projecting parts of the surface. 
Polished, mirrorlike enamel surfaces present on all tubercles and transverse crests forming a network 
all over the tooth. They are easily visible with the naked eye. 

al. Areas of exposed dentine on elevated parts of the surface, disconnected and not forming 

a general, continuous dentine network. 

a^. First or second tooth with areas of exposed dentine along inner crests. Dentine is exposed 

on the transverse crests and on the high external tubercles which are worn and about 3/4 of 

their normal height: specimens more than 1 year old. (Figure 170,5). 

b^. All teeth (except p^ which remains almost unchanged) with areas of exposed dentine along 

inner crests. Teeth considerably worn and their high external tubercles 1/2 - 3/4 of their 

normal height. A polished enamel stripe around the areas of exposed dentine all over the convex 

parts of the occlusal surfaces: more than 2 years old. (Figure 170, 6). 

b^. Areas of exposed dentine either connected and forming a general network over the surface 

or mostly connected along the inner tubercles coimected with areas along the transverse crests. 

c^. Network of dentine absent. Spots of dentine connected along inner tubercles and anterior 

transverse crests of m^ and m (sometimes also of p^). Projecting rings of enamel present along 

the network. The outer tubercles are worn, half their normal height, or less: more than 3 years 

old (Figure 170, 7). 

d2. Network of dentine present all over the tooth and bordered with projecting ridges of enamel. 

The surface of exposed dentine occupies about 1/4 of the occlusal surface. Crowns of the teeth 

with very low (strongly worn) tubercles: more than 4 years old (Figure 170, 8). 

e^. Network of dentine present all over the tooth. The enamel spots near the depressions begin 

to disappear. Exposed dentine covers 1/3-1/2 of the surface. Crowns flat, without tubercles: 

more than 5 years old (Figure 170, 9). 

f^. Exposed dentine covers more than half of the occlusal surface. Surface of the teeth 

depressed: more than 6 years old (Figure 170, 10). 



339 



The color of the squirrels is very variable and the variations are 
described under the various subspecies. 

The belly of all forms of Sc iur us vulgaris is white. However, the 
size of the white area is variable. In Russian- European squirrels and in the 
West Siberian subspecies the white area covers the area from chin and 
throat to the groin and also white spots on the inner side of the legs. It is 
reduced in central and East Siberian squirrels. Dark fur is present on the 
sides of the throat and reaches the chin. As a result, a narrow triangular 
figure of white is present on the throat. The white color is also strongly 
reduced in the inguinal region and on the spots on the inner sides of the 
legs. The inguinal region is often dark. 

Color of back varying with age and season. In east European summer 
specimens, back different shades of red-brown-rust to deeper dark brown 
tones. Back of central Siberian and East Siberian squirrels dark brown, 
often "black. The color of squirrels from mountainous regions is more 
intense than that of squirrels from the plains. 

Winter specimens sometimes show so-called "baldness" of the spine. 
This consists in the appearance of a rust-colored stripe along the spine in 
winter fur. Such pelts should not be confused with late autumn pelts with 
remains of rust-colored hair on the spine. 



295 



The following color types of the tail are found: 

Black-tailed: Tail with completely black hair. 

Dark-tailed: Tail black- gray. The roots of the hair are dark gray and 
the tips black. 

Brown-tailed, tail brownish. The hair has gray roots, rust-colored 
central parts and black tips. This group is divided into two subgroups, 
dark brown-tailed, and light brown-tailed. 

Red-tailed specimens with rich rust-colored tail. The hairs have gray 
roots and rust-colored tips. 

Dark red-tailed: Many specimens of the Yenisei squirrels. Their tail 
is reddish rust-brown with a strong admixture of deep black-brown or 
black tone. Roots of hair smoky bluish gray; middle parts rich rust-brown, 
broad black or black-brown subapical bands and rust-colored tips. 

Gray-tailed specimens: tail generally of a silver-gray tone. 

The vibrissae of S c iuru s vulgaris have a typical distribution. A 
group of small vibrissae present near the lips, and larger vibrissae along 
the sides of the nose, close to those on the lips. A small group above the 
eyes, the longest may be 25 mm long. Four or five vibrissae, about 
35mm long, in a dense group on the cheeks behind the eyes. Several 
vibrissae on the outer parts of the antebrachium, about 25 mm long. B.A. 
342 Kuznetsov (1928) found a group of six short vibrissae on posterior side 
of the foreleg, at the base of the wrist. I. P. Kiris who studied the 
structure and distribution of vibrissae came to the conclusion that the hairs 
at the base of the wrist are not vibrissae. Two separate pairs of long 
vibrissae on the belly. The first pair on the chest posterior to the base of 
the forelegs. The second pair about 2 cm behind the first. These 
vibrissae are 30-35 mm long. They are directed backwards and lie 
close to the body, only slightly projecting from the fur. The vibrissae 
have a rather constant form, they are straight, stiff, tapering at the tip. 
They project clearly from the fur. Their color is constant, and does not 
depend on the age of the animal or the season. The vibrissae of the head 
are dark brown, almost black. The vibrissae on the feet are rust-colored, 
sometimes yellowish. Some of them have dark bands. The vibrissae on 
chin and belly are white. 

The forefoot has four toes. The first is much reduced. The second is 
about as long as the fifth; both are much shorter than the third and the 
fourth. The third is nearly as long as the fourth. 

The first toes of the hindfoot is the shortest. The third and the fourth 
are of about the same length. The second is shorter. The fifth toe is 
thicker and longer than the first, but shorter than the fourth. 

SYSTEMATIC NOTES. Sciurus vulgaris in the U. S. S. R. forms 
16 or 17 subspecies which have transgressive characters. Each subspecies 
is connected with the adjacent subspecies by transitional forms. Some of 
these subspecies are closely related showing a definite composition of 
characters such as a certain combination of colors on the tail, bald-spine 
in winter, density and luxuriance of fur, etc. It is often difficult to identify 
a single specimen because some characters become evident only in 
series. Some subspecies can be easily identified from single specimens, 
e.g., S. vulgaris kessleri Migulin, S.v. bashkiricus Ogn. , 
S.v.exalbidus Pall., etc. 

The following subspecies occur in Western Europe: l) Sciurus 
vulgaris vulgaris L. —Scandinavia; 2) S . v . leucurus Gmel. — 



296 




2-3 months 







More than 2 years 




3-4 months 





More than 3 years 




4-5 months 



^^^Xj^^^^ 




More than 4 years 




6-9 months 




More than 5 years 




More than 1 year 




10 



More than 6 years 



FIGURE 170. 



297 



FIGURE 1 70, Stages in the abrasion of molars ofSciurus vulgaris L. Description of the principal 
age groups of the squirrel after changes in the upper tooth row. 

1. 2-3 months. Upper tooth row incomplete, third molar (m^) not yet cut through or only beginning 
to erupt; second premolar (p*^ deciduous, its crown triangular. Its occlusal surface about half the size of 
that of the first molar (m ). Anterior root of p^ strongly projecting forward. First premolar usually absent. 
Areas of exposed dentine (the bony substance of the tooth under the enamel) present only on the large outer 
tubercles of second premolar. 

2. 3-4 months. Upper tooth row complete, only p^ absent, p^ deciduous. Areas of exposed dentine 

2 1 2 

on outer tubercles of p and m . A fine line of exposed dentine sometimes present on inner tubercle of p . 

2 

3. 4-5 months. Tooth row usually incomplete or not yet level. Deciduous p replaced by permanent 

tooth. Deciduous p sometimes persisting on one or both sides of maxilla, but the new permanent tooth is 
visible beneath it. In this case, the deciduous tooth is raised by the permanent tooth and projects markedly 
above the surface of the upper tooth row, 

4. 6-9 months. Upper tooth row complete and level. All teeth permanent, fully grown. Small spots 
of exposed dentine on the large outer tubercles of all teeth. Short lines of dentine sometimes on the 
transverse crests of m and m , and on inner tubercle of m . 

5. More than 1 year. Areas of exposed dentine on outer tubercles of all teeth oblong-pyriform. Only 
m^ and m^ with spindle-shaped areas of exposed dentine on the transverse crests. Inner tubercles of m and 
m2 with well-marked crescent-shaped areas of dentine. Rarely a faint line of dentine on the inner 
tubercle of m^. 

6. More than 2 years. Spots of dentine on the large outer tubercles fused with dentine areas on 
transverse ridges. Crescent-shaped areas of dentine on inner tubercles of all teeth; that on inner tubercle 
of p2 usually less marked than on the other teeth. 

7. More than 3 years. Small tubercles on outer margin of teeth also worn, with small spots of dentine. 
Areas of dentine on m^ fused into a horseshoe -shaped figure with apex directed inward. Transverse stripes 
of dentine on m and m^ close toenlargedfalcate dentine spots on inner tubercles, but not reaching them. 
Posterior margin of m^ with small dentine stripe. 

8. More than 4 years. Area of exposed dentine markedly increased, covering about a quarter of the 
surface. Transverse dentine stripes coalesced or almost coal'-.sced with dentine spots on inner tubercles of 
all molars, forming horseshoe -shaped figures on m and m^. Dentine area on m^ of the shape of half a 
horseshoe. 

9. More than 5 years. Dentine bare on all tubercles and crests of upper tooth row. Area of dentine 

network covering about 1/3 of the surface. Dentine spots on transverse crests and inner tubercles fused 

P 1 5 3 

with the horseshoe -shaped figures of p , m and m'^. Posterior margin of m strongly worn. 

10. More than 6 years. Dentine area covering about half the surface. Tubercles and crests of all four 
teeth little elevated above surface. Dentine spots on all teeth horseshoe-shaped. A considerable part of the 
concave surface of the crown of m worn (according to I. D. Kiris, 1937). 



298 



343 



343 




FIGURE 171. Clans penis of squirrels 
(highly magnified) 



England; 3) S. v. russus Miller — western and central Europe, from 
Britanny to Holland and central France; 4) S. v. fuscoater Altum — 
eastern and central Europe, Germany to Austria, Hungary and Rumania; 
5) S. V. italicus Bonaparte — Italy; 6) S. v. lilaeus Miller — 
Agoriani, north of Lyakura — Parnassus in southern Greece; 7) S . v . 
ameliae Cabrera — Greece; 8)S.v.alpinus Desmarest — Pyrenees; 
9) S. V. numantius Miller —Spain, Pinares de Quintanar de la Sierra, 
Burgo; 10) S. v. in fuse at us Cabrera — Spain, Las Navas, Avila; 
ll)S.v.segurae Miller — Spain, Molinicos Sierra de Segura; 12) S. v. 
baeticus Cabrera — Spain, Alapis, Sevilla Province, The following 
subspecies occur in the Far East: 13) S . vul ga r i s orientis Thos. — 
described from Hokkaido (Yezo); according to O, Thomas squirrels from 
the vicinity of Seoul in Korea are closely related to this Japanese form. 
Thus, 30 Palearctic subspecies ofSciurus vulgaris are known so far. 
GEOGRAPHICAL DISTRIBUTION. (Map VI). According to F. D. Pleske 
(1884) the northern boundary of squirrels in the Kola Peninsula is connected 
with coniferous forests. North of the Imandre squirrels are rare. Pleske 
obtained a specimen from Raznovolok (north of the Imandre) and saw pelts 

from the Kola Peninsula. In Eastern 
Europe the range of the squirrel reaches 
nearly to the Arctic circle, north as far 
as the forest belt. The northern boundary 
crosses the Urals at about the Arctic 
circle, and turns south between the Ob and 
the Taz (see map), S,I. Orlov (l930) 
writes as follows on the distribution of 
squirrels in these parts of Siberia: "In the 
Taz District north of Syurge (67°20'N and 
49°50'E), many natives never caught 
squirrels except in 1923, duringthe mass 
northward migration of squirrels on the left 
bankof the Taz. In autumn of this year a 
large number of squirrels reached the Taz 
Bay and tried to swim across to the right 
bank. They drowned because they were 
exhausted from the long journey and from 
hunger, as they could not subsist on the 
few dwarfed larch trees in the tundra. In 
spring 1924 I found several nests in which 
the squirrels had frozen to death in an 
alder grove in the estuary of the Kopur 
river. The dead squirrels were very 
emaciated, their stomachs contained only moss and water, and their fur 
showed many bare patches, , , The squirrel is common along the Tur and 
344 Taz rivers (250km from their estuaries), but occurs only in small 

numbers. East of the Taz river the squirrel reaches the sources of the 
Turukhan river and farther east Lake Khantaiskoe where it was caught by the 
Yuraks and the Khantaika Tungus. , . Farther east the boundary apparently 
turns south, as squirrels were not found along the Khatanga Highway, In 
my collection is a specimen from the Rossomazh'ev Station. According to 
the Yakuts squirrels are very rare along the Kutoi river near the estuary 
of the Kotuikan, although there are forests there. Squirrels do not occur 
along the Khatanga where the forest is 4.5 m high. " 





FIGURE 172. Os penis of squirrels. 1- 
lateral; 2 - doisal (highly magnified) 
Drawing by Yu. A. Kostylev 



299 



These data are confirmed by observations of E.O. Yakovlev (1930). The 
squirrel is widely distributed in the Turukhansk Territory, spreading far 
north. It was found near Khatanga along the lower reaches of the Kheta 
river. It is common along the right tributaries of the Kheta river (e.g., 
Ayakla river), at about 70° N. In the Pyasinsk District the squirrel was 
found near the village Tungusy (70°30'N). It rarely reaches the village 
Dudinsk near the Yenisei (about 69°N). Farther west it is found along the 
Malaya Kheta river. It was not found farther north. According to R. 
Maak (1886), it is very rare along the Olenek river (about 68° N). The 
northern boundary of squirrels between the Yenisei and the Lena thus 
reaches 70°N or a little beyond. 

East of the Lena the boundary crosses the Yana and Indigirka rivers 
south of 68°N. According to V.I. lokhel'son (1898), the squirrel disappears 
northwest of Srednekolymsk, near the circiimpolar belt. Numerous 
squirrels occur on the right, "stony" bank of the Kolyma between the upper 
reaches of the river and Anyuev. 

In the Anadyr Territory the squirrel is apparently restricted to isolated 
larch forests. According to N. P. Sokol'nikov (1927), there is such a 
forest, 170 km wide, 50-60 km north of Markov. Another is found on the 
upper reaches of the Maina beginning from its tributaries, the two Algans. 
This forest is smaller than the forest near Markov. These forest islands 
are generally very small, l-3km wide, according to Sokol'nikov. They are 
separated from the east Siberian forests by tundra. In the Anadyr 
Territory the squirrel occurs along the Shchuch'ya, Yablona and Yeropol 
rivers (S.N. Barmontov (in litt., 1933)). 

The squirrel does not occur in the tundra north of Kamchatka. It has 
been generally accepted that squirrels do not occur in the Kamchatka 
forests. The first data on the invasion of Kamchatka by squirrels 
appeared in the 1920's. According to P. T. Novograblenov (1930), the 
migration of squirrels to Kamchatka from the Gizhiga and Penzhino districts 
began in 1923/24. The spread of squirrels was as follows: from the 
Tigil' Valley they passed through the Anaun Valley to the Kozyrevka river. 
They thus reached the Kamchatka valley where they moved in both 
directions. They then passed to the Avacha Valley. According to S. N. 
Barmontov (in litt. 1933), squirrels were found along the Dvugortochnaya 
and Kamchatka rivers (and the tributaries of the latter). They were 
numerous in the Mil'kov — Naniki districts, and gradually moved south to 
the Petropavlovsk District (Koryaki, Elizova, Nalychevo, Nikolaevka 
villages). 
345 The northern boundary at the extreme eastern corner of the range of 
Sciurus vulgaris thus reaches the northern part of Kamchatka. 

The southern boundary of the range. S. vulgaris occurs in the forests 
of northern Bessarabia and in Podolia. It was found in the following 
localities, which are isolated from the main range: the Birzulo-Zherebkov 
forests in the Tiraspol District near the Dniester (A. A. Brauner, 1923); 
the deciduous forest in the former Aleksandriiskii County, the Grinen 
Forestry of the former Tul'chinsk District (A, A. Migulin), the Samara 
Forest along the Dnieper. 

Farther east the southern boundary passes south of the former Kiev 
Province in the Zvenigorodka District, Cherkas (A. A. Migulin, "Mammals 
of the Khar'kov Province" (Mlekopitayushchie Khar'kovskoi gubernii), 
1924). On the left bank of the Dnieper the boundary follows 50°N. 



300 



According to N. Gavrilenko (1928), squirrels still occur in the eastern part 
of the former Poltava Province in the northern part of the former 
Konstantinograd County. According to A. A. Migulin (1924), the range of 
the squirrel includes the Valok and Zmiev districts. The boundary then 
turns sharply north near the Khar'kov and Kura steppes to Sumam, L'gov, 
Dmitriev, including forests of the former Trubchevsk, Briansk and 
Karachev counties. Squirrels were found in the following places, which are 
separated from the main range: near Izyum (A. A. Migulin), the Belgorod 
District (Shebenin, Nizhegol' and Murman forestries), Serebryan Pine 
Forest of the former Artemovskii District. 

According to S.N. Gorbachev (1915), the squirrel occurs near Orel, near 
the village Mylinki (former Karachevsk County). The boundary then runs 
north to include the Bolkhov, Zhizdrinsk, Kozel', and Kaluzh forests and 
the northern forests of Tula Province. Farther east the squirrel is found 
in the forests of the former Ryazan Province, in the former Ryazan and 
Kazimov counties (S.S. Turov, "Mammals of the Ryazan Province" 
(Mlekopitayushchie Ryzanskoi gubernii), 1925, p. 59), in the Morshansk, 
Kadomsk and Temnikov forests of the former Tambov Province. The 
squirrel is widely distributed in the former Penzen Province (F. 
Fedorovich, 1915). According to B. M. Zhitkov (1898), the squirrel was 
once common in the former Simbir Province in forests of the former 
Ardatov County. They are more common in the Lukoyanov and Sursk 
forests. The boundary then turns sharply north, crossing the Volga 
between 55° and 56° N. 

There are several records in the forests of the Chernozem Region in 
the south. Squirrels were also found farther north, isolated from the main 
forest. 

N. A. Severtsov (1855) found the squirrel in the Khrenov Forest (former 
Bobrovsk County of the former Voronezh Province), in the Graf Forestry 
(near Voronezh), in the Shipov Forest (former Pavlovsk County) and in the 
Lipov Forest. 

In "The Fauna of Terrestrial Vertebrates of the Voronezh Province" 
(Fauna nazemnykh pozvonochnikh Voronezhskoi gubernii), 1923, p. 153, 
o^p I wrote "Squirrels have become very rare in the Khrenovsk Forestry. 

During long and regular expeditions we only once caught a young squirrel 
(22 July (4 August) 1921). We also received the pelt of an old winter 
specimen from the forester of the Maidan Forest Belt. S.I. Ognev shot 
a female on 22 September (5 October) 1920 in the Graf Forestry not far 
from the Graf Station. 

"N.N. Spitzyn, the forester of the Graf Forestry states that squirrels 
were formerly very concimon in forests near Voronezh. Their sharp 
decrease is apparently the result of the increase of martens. I.D. 
Tushnev told us that in the Shipov Forest the squirrels are being 
exterminated by the numerous martens. * 

"p. Barants (in litt. , 1895) writes that the squirrel is very common in 
the Khrenov Pine Forest. It occurs both in pine forests and in deciduous 
forests near the river. The squirrels have decreased considerably in the 
last decade. In the late 1880's and the early 1890's they were very common 



* It seems more plausible that the squirrel population in the Khrenovsk Forest decreased as a result of lack 
of food in years with poor crops of cones and nuts. The food supply of squirrels in the Shipov deciduous 
forest is generally poor. 



301 



in the forest. At present (1895) squirrels occur in the forest but only 
in small numbers. " 

Squirrels occur in the Tellerman Pine Forest (former Borisogleb 
County of the Tambov Province, along the Khotra river). A. A. Silant'ev 
(1894) wrote that squirrels are very rare in forests of the former Balashov 
County, Saratov Province. The range of the squirrel in this region was 
apparently isolated from the main range. In the middle reaches of the Volga 
the squirrel apparently occurs south to the line Serdobsk, Petrovsk, 
Syzran and Vol'sk. 

In the Trans-Volga Region the squirrel survived sporadically in the 
central part of the former Kazan Province (e.g., in the Cheremshan Forest 
along the Cheremshan river). It does not occur on the left bank of the 
Belaya river. It occurs rarely in forests on the right bank of this river, 
e.g., in the Angasyak Forestry of the former Birsk County of the Ufa 
Province. The occurrence of squirrels in the Bazuluk Pine Forest is 
isolated from the main range. Farther east the squirrel occurs on the 
southern Urals, in the Zlatoust District, on the Ilmenskii Range (Dukel'skaya, 
1928). According to N.A. Zarudnyi (1897), it is common in forests of 
Orenburg Bashkiria, M.K. Serebrennikov (1929) obtained 2 squirrels from 
the eastern part of the Chelyabinsk District. A.I. Argyropulo found 
squirrels in the Troitsk District. Farther east the southern boundary 
continues through the south of the former Tobolsk Province (the Tara 
District) and, skirting the Barabinsk steppes, enters the central part of 
Tomsk Province. It then turns sharply south to the Altai Region. The 
west Siberian subspecies Sciurus vulgaris exalbidus occurs in 
isolated localities in "ribbon" pine forests along the Irtysh and the upper 
reaches of the Ob. 
3^Y The southern boundary of the squirrel in the southern Altai Region is 
not well known. It apparently occurs in Tarbagatai, the southern Altai, 
the Tannu-Ola Range, the Sayan Mountains, in forests of the Uryankhai 
Territory, and on the Kentei Mountains. The squirrel occurring in the 
Kontey mountains is apparently closely related to the Altai squirrel. The 
southern boundary then reaches the southwestern shore of Lake Baikal. In 
Transbaikalia, the squirrel occurs throughout the forest zone, beginning 
from the Barguzin forests. It occurs throughout the Great Khingan, the 
north Manchurian forests and Korea. 

The squirrel also occurs in Japan and in Sakhalin. 

The distribution of the squirrel in this large area is not uniform. Its 
numbers fluctuate markedly because of food supply and the varying rate of 
reproduction. The squirrel is subject more than other animals to 
fluctuations in numbers in different localities. 

MODE OF LIFE. The squirrel depends on the forest. Absence or 
presence of forests thus results in absence or presence of squirrels. 

In different regions of its range the squirrel prefers different types of 
forest. Its choice depends mainly on the presence of food. When there are 
many spruce cones the squirrels concentrate in spruce forests. When 
there is a yield of pine cones they move to pine forests. When there is a 
crop of hazelnuts they move to the nut trees, etc. However, other factors 
also influence the choice of habitat: density of the trees, the presence of 
fallen and hollow trees, and the density of the undergrowth. These factors 
enable the squirrel to hide from enemies, mainly from birds of prey. They 
also supply sites for nests, e.g., dense undergrowth and tree holes. 



302 



348 



The suitability of a locality for squirrels is determined by the grass - 
shrub layer of the forest, the presence of a dead moss layer, the quality of 
the soil, etc. Of great importance is secondary food such as mushrooms 
and berries. An important factor is the presence of small competitive 
rodents on the ground which eat the fallen cones and nuts. The grass layer 
of the forest also affects the squirrels, as it serves as habitat for these 
small rodents. In spite of the variety of habitats inhabited by the squirrel, 
in any region there are some forests which are most suitable to be 
permanently inhabited by squirrels. 

In the Moscow Region, the squirrel lives mainly in forests in which 
spruce predominates. It usually builds its nest where the spruce forest is 
dense. According to B.A. Kuznetsov (1929), in the Pogonno- 
Losinoostrovskii Forestry the squirrel most frequently occurs in under- 
brush in pine and spruce forests, in linden-spruce forests and spruce 
nurseries. It rarely occurs in young spruce-larch and pure larch 
associations. The squirrel enters this region from the high forest. It is 
especially numerous in pine forests and in underbrush in pine forest with 
spruce in the second layer. It is very rare in old pine trees with reindeer 
lichen which has no undergrowth of spruce. Although such forests produce 
abundant cones, they are less suitable for squirrels as the trees are 
widely spaced, and there is much light and little shelter against predators. 
The cones in these forests are naostly eaten by woodpeckers and crossbills. 

S. V. Lobachev (1932) gives a detailed account on the habitats of 
squirrels in the northern European U. S, S. R. , in the Verkhnevyechegod 
District. Squirrels occur in this region in the following types of forests: 

1) "Parms", i.e., spruce forests on sandy loam, loam, clay and soils 
in humid valleys with a base of moraine clay. "Parms" are the principal 
habitat of the squirrels. 

2) Spruce marshes overgrown with few, low spruce with grass cover 
consisting of sedge, rushes, etc. 

3) Birch forests with spruce underbrush. 

4) Pine forests, e.g., these with moss ground or marshes and pine- 
birch forests. 

The squirrel is never found in birch groves, peat-bogs, pine marshes, 
and burns with new young birch. 

In the south, e.g., the Graf Forestry of the Voronezh Region (former 
Voronezh County) the squirrel occurs in oak forests (where it feeds on 
acorns), and also in mixed forests with pines. According to A. A, Migulin 
(1924), squirrels occur in the former Kharkov province both in pine forests 
and in broad-leaved forests. They are particularly common in pine forests 
mixed with deciduous trees. 

As mentioned above, the squirrel moves to different habitats because of 
fluctuations in the food supply. It therefore spends some time in different 
types of forest. 

A detailed description of the occurrence of squirrels in certain types of 
forest is given in an article by I.D. Kiris (1934), who studied the ecology of 
the central Ob squirrel. 

1. The characteristic habitat in the Surgut Territory is "insular forests". 
These are characterized by: a) separation from the taiga by large meadows 
(called "sors"); b) the small area of each forest island; c) the prevalence 
of stone pine stands over other forest types; d) rich food supply in the rare 
productive years, and almost complete lack of food during the long periods 
of crop failure. 

303 



The squirrel does not nest or reproduce in insular stone pine forests. 
It may spend years there, during years with a rich harvest of nuts. 
349 2. The second habitat may be called "principal inland stone pine 

forests" or "urmans". Large areas are covered by stone pine stands which 
have broken borders making the margins of the forest very long. These 
are the most suitable habitat of the squirrel. There is a rich and constant 
food supply because of the presence of many coniferous trees besides stone 
pine. The seeds of these trees diversify the food of the squirrel. This 
habitat also has a good development of the various stories of the forest, 
providing good shelter. 

The squirrels live constantly in "urmans", reproduce and molt in them. 
According to I.D. Kiris, nests are most frequently found in stone pine 
urmans. He found an average of 0.139 squirrel nests per kilometer. 

3. Large fir and spruce forests, which often pass gradually into stone 
pine forests. Here the squirrel finds good shelters and a constant food 
supply. 

The squirrel is uniformly distributed in fir and spruce forests. However, 
the density of population is much lower than in urmans, except for during 
the molt, when the squirrel is even more numerous here than in urmans. 

4. In pine forests which are separated from other habitats and far from 
urmans, the squirrel does not find good shelter and sufficient food. It does 
not nest here and stays only on its way to other more suitable places. 

5. The squirrel does not occur in pine marshes with large peat bogs 
covered with small pines and birches. I.D. Kiris saw a squirrel only once 
in this habitat. 

6. Birch groves within other forests supply good shelter in the form of 
dense coniferous underbrush of stone pine, fir, pine or spruce. The food 
supply in such a habitat is poor and squirrel nests are therefore absent. 
The squirrel does not usually stay long in birch forests. However, it molts 
in denser parts of the birch forests, as they give good shelter. Squirrels 
are found in birch forests during migi ations. 

In western Siberia the preferred habitat ofSciurus vulgaris 
exalbidus is pine forests on the right bank of the Ob, between Biisk and 
Novosibirsk, and pine forests from the Ob to the Irtysh, across the 
Kulundinsk Steppe. These pine forests are of a humid type. They grow on 
sand dunes. Professor V. V. Reverdatto (1931) describes such a forest, the 
Verkhneobskii Pine Forest: "This enormous forest has a high moisture 
retention. Many streams therefore originate in it and there are several 
large lakes like the Bol'shoe Kamyshnoe, etc. The boreal character of 
this forest, including the presence of peat bogs caused by the high rainfall 
produced sandy podzolic soils (as these forests grow on sand dunes 
deposited by the Ob river). Large areas are occupied by Pinetum 
vaccinosum and other forest associations such as Pinetum 
herbosum (grassy pine forest), Pinetum hylocomiosum (mossy 
pine forest), etc. The grass cover and the vegetation of the marshes have 
a definite northern character. 

The only other coniferous tree in the Verkhneobskii pine forest is larch 
(rare). There are many lakes of various sizes in the Ozerskii forest. 

The "ribbon" pine forests between the Ob and the Irtysh (on the left bank 
of the Ob) are restricted to upper quaternary sand in depressions along rivers. 
These ribbons run from the northeast to the southwest. Most of these pine 
forests are narrow strips, about 15km wide and 106km (Burlin and Kulundin 



304 



350 



forests) to 213 km long (Kasmalin and Barnaul forests). These lie lower 
than the steppe areas between them. In the monograph "The Teleut 
Squirrel" (Belki-teleutki*, 1935), G. K. Gol'tsmaier distinguishes the 
following types of ribbon pine forests: l) green-moss pine forest; 2) dry- 
lichen pine forest; 3) grassy-steppe pine forest. 




351 



FIGURE 173. A squirrel running on a branch 

Drawing by A. N. Formozov 

The Srostino forest is the southernmost of the so-called "central group", 
northwest of Rubtsovsk. Gol'tsmaier describes it as follows: "There is no 
running water in the Srostinsk forest. Lakes and marshes named "sogr" 
constitute the only water bodies. They lie in deep depressions between sand 
banks and are overgrown with vegetation. Free water surfaces are found 
in the deeper parts of the marshes. The shallow parts of the marshes are 
covered with an impenetrable thicket of cattail, cane, bulrushes, sedge and 
other marsh plants. Their rhizomes form a shifting cover on the water. 
The marsh is surrounded by a wall of cane or bulrushes which is replaced 
on the outside by a ring of willow (Salix cinerea L.)ona layer of thick 
moss, overgrown with reeds (Phr a gm it e s communis Trin. ) and 
some grasses. Higher up, the willows are replaced by dense aspen and birch 
underbrush. The soil is completely covered with fallen leaves, and the 
grass cover is sparse. Further inland the ring of deciduous trees is 
replaced by dense young pine forest (40 years old or less). The ground is 



* [Teleut squirrel = S c iurus vulgaris exalbidus Pall.] 



305 



covered here by the leaves of the deciduous trees, pine needles and peat 
moss. The grass cover is dense, including also So li dago virga 
aurea L. , Artemisia campestris forma glabra, Hypochoeris 
maculata L. , Pyrola sp. Yellow acacia trees form underbrush in 
some places. Higher up, the trees surrounding the marsh are replaced by 
old pine forests with thin grass cover in which Fe stuc a ovina L, 
predominates. A few of the following plants also occur among the grass: 
Alyssum desertorum Statt. , Artemisia campestris forma 
glabra, Koeleria glauca D.C., Melandrum album Larcke, 
Euphorebia esula L., Pulsatilla patens Mil., Stipa 
ioannis Czelak. , Allium nutans L. , Phragmites communis 
Trin. , Solidago virga aurea L. , Silene volgensis Bess., S. 
chlorantha Ehrh. , Phleum boehmeri Wibel, Scabiosa 
ochroleuca L., Hypochoeris maculata L., Equisetum 
hiemale L., etc. 

"The grass cover is thin, and the 
soil is visible everywhere, barely 
covered with pine needles and cones. 
In some places, mainly in the small 
depressions, there are small green 
„ shoots, pine underbrush, and lichen. 

j^'Tf (^^ "X Yellow acacia bushes are rare, small 

aspens even rarer. . . The higher 
ground is occupied by steppe pine 
forests." 

This form of pine forests produces 
well developed forest margins around 
marshes and clearings which occupy 
about 25% of the area of the pine forest. 
Trees growing in them give better 
crops. Forest borders are the most 
suitable habitat of squirrels, as food 
is more plentiful in them than in other 
parts of the forest. 

According to K. A. Zabelin, in East 
Siberia, e.g. , in the Barguzin forest 
of Transbaikalia, squirrels occur along 
the shore of Lake Baikal and high in the 
mountains near the limit of vegetation. 
Although squirrels may live in various 
types of forest, they prefer dry, high 
and dark pine forests. Depending on 
the supply of cones, squirrels occur 
either in cedar, pine or larch forests. 

According to G.N. Gassovskii (1927), 
in the Gilyui-Oldoi District (East 
FIGURE 174. Squirrel looking out from behind Siberia) squirrels occur in mixed, 

coniferous or in deciduous forests, 
in broad river valleys and at the limit 
of mountain forests in thickets of creeping cedar underbrush. They are 
especially numerous in coniferous forests, mainly in Jeddo spruce forests 
along streams and brooks, and in pure pine forests on terraces above 
rivers, on mountains and dry, steep slopes. Squirrels rarely occur in pure 

306 




a tree 

Drawing by A. N. Formozov 



larch forest. G.N. Gassovskii states that the occurrence of squirrels in 
different habitats depends on the food supply: cones of larch, spruce or 
cedars, and the mushrooms in deciduous forests. 

Although the squirrel typically inhabits the forest, it also occurs 
exceptionally in the dwarf forest or tundra shrubs. This is due to 
migrations (see below). 

The squirrel is well adapted to arboreal life. Its soles are padded and its 
toes have sharp, curved claws which enable it to grasp branches firmly. 
Its movements are elegant and rapid. It climbs trees quickly, its tail 
outstretched. It maintains its balance well, even on thin, moving branches, 
by lowering or raising its tail. It is able to perform great leaps. It leaps 
2-3 m or more. Its leaps are especially long when it jumps downwards 
(Figure 175); when it jumps from a tree to the ground it may cover 15m. 
When a squirrel sitting on the ground is frightened, it gives a sudden 
squeak, rapidly running up a tree. When threatened it hides by pressing 
oj-o itself tightly to a branch or conceals itself among the dense branches. 




FIGURE 175. A squirrel leaping from one branch to another 

Drawing by A. N. Formozov 

When resting, the squirrel sprawls against a branch, extends tail and 
legs and basks in the sun. 

In winter, it either moves from one tree to another, or more rarely 
moves on the snow, depending on the weather. When snow hangs from the 
branches the squirrel often has to move on the ground, because when it 
lands on the branches it cannot grasp them owing to the crumbling snow. 
When the branches are free of snow, the squirrel prefers to remain above. 
It always moves in the trees when going to the nest. Where squirrels are 
little persecuted, they let man approach. Young squirrels are also very 
trusting. When man appears, the squirrel shows a keen curiosity. It 
settles on a branch, raises and lowers its tail, swings and leans forward. 



307 



354 



giving its typical squeak. When it is frightened it rapidly climbs to the top 
of a tree, stops on a branch, and continues to scrutinize man. 

The usual voice of the squirrel is a typical quick squeak. To express 
joy it may emit a soft whistle. The plaintive cry of a wounded squirrel 
resembles that of a wounded hare. 

Squirrels can swim for a long time. Migrating squirrels swim across 
broad rivers like the Ob and Yenisei. A.K. Zabelin states that squirrels 
occasionally swim across the Chivyrkui Inlet of Lake Baikal which is 10- 
12 km broad in some places. While swimming the squirrel holds its tail 
inclined and raised above the water. The tail of the exhausted animal drops 
and becomes submerged. 

The squirrel is a diurnal animal. Its daily life begins soon after sunrise. 
In July active squirrels maybe seen as early as 5a.m. After leaving the 
nest, the animal plays, climbs on trees, runs on the ground and is very 
active. It then begins feeding. If it becomes tired it may take a short 
nap. It rolls itself into a ball on a branch, tucks its head under its body, 
and covers itself with its tail. In summer, during the midday heat 
squirrels take shelter in their principal or auxiliary nests. They 
sleep rolled up, waiting until the heat subsides. They appear again when 
the afternoon becomes cooler, and continue their activity. They return to 
their nests before sunset. G.K. Gol'tsmaier (1935) writes: "After 
separating from their mother, young squirrels spend the night in any 
shelter they may find or on a thick branch, as they have no nests of their 
own. On 9 September 1930, early in the morning, I found a young squirrel 
which has spent the night in the upper third of a high pine, on branches 
near the trunk. " 

G.K. Gol'tsmaier thinks that the Teleut squirrels sleep about 7-8 hours 
a night in summer. 

The type of food in a locality influences the daily activity. A.N. 
Formozov (1934) writes: " Squirrels in spruce and pine forests spend 
almost the whole day extracting seeds from the cones. On the other hand, 
squirrels in oak forests still their hunger with 3 or 4 acorns. In winter 
they sleep in their nest soon after sunrise. " 

In autumn, as the days become shorter, the animals come out of the 
nest later and return earlier in the evening. The midday rest becomes 
shorter and finally is given up completely. Morning and afternoon feeding 
periods become united into one period of activity. This is typical for 
late autumn and winter activity. 

The short summer rains apparently do not affect the daily activity of the 
squirrel. When it is surprised by rain, it waits in a shelter for good 
weather, but it may sometimes ignore bad weather. I frequently saw 
squirrels active during rain. Rain in autumn and continuous rain followed 
by snow usually drive the squirrel into its nest, or into shelter among 
branches. 

Snowstorms in winter certainly affect the squirrel. The animal 
then sits in its shelter and does not come outside. G.K. Gol'tsmaier 
writes: "On 18 November 1931, my assistant Smakotnin found a 
squirrel nest during a snowstorm. Attempting to drive the squirrel from 
the nest he began shooting over the nest and shaking the tree, but in vain — 
the animal did not appear He therefore thought that the nest was empty, 
pushed it off with a pole and brought it to the sledge. Only then did a 
squirrel jump out, and Smakotnin caught it. It was a healthy female which 



308 



we kept a long time in captivity. Its reluctance to leave the nest was 
certainly not due to injury or disease. " Golts'maier observed that severe 
frosts do not influence the daily activity of the squirrels. He saw, for 
instance, squirrels leaving their nest at temperatures between -30° and -42°. 
This severe frost occurred in clear and calm weather, without snow. 

N.P. Naumov (1930) observed the following: "Near the Stony and Lower 
Tunguska, the squirrel is active in the first half of winter. It comes out 
of the nest in the morning. It does not feed on its food reserves, but digs 
deep under the snow to feed on what it can find. When the snow covers 
the ground in a thick layer and hangs heavy on the branches, it hinders the 
animal's climbing. There is nearly no wind in this season. . .The appearance 
of large lumps of snow on the branches is followed by true arctic frosts 
(-50° to -60°). The squirrel is therefore forced to remain in its nest from 
the second half of December, leaving it only on warm days to feed a little. " 

In the central part of the European U.S.S.R. squirrel nests are mainly 
found on spruce. M.P. Raspopov and Yu. A. Isakov (1935) give a detailed 
description of squirrel nests in the Moscow region. 

They examined 128 nests. Of these 119 were on spruce, 7 on pine, 1 on 
birch and 1 on larch. 




FIGURE 176. Squirrel tracks in the snow near a hollow with a store of nuts 

Drawing by A. N. Formozov 

356 The squirrel generally builds its nest near the trunk, usually at the fork 
of two or three branches. The nest is rarely built in the middle or at the 
end of branches of trees which stand close together. In the Moscow region 
nests are built on trees 15 - 30 cm thick. They are usually 5-15 m above 
the ground. Nests on pine trees are usually higher up than on spruce. 
This may be due to the fact that the stem of pine trees is without branches to 
a greater height. The nest has therefore to be built in the dense upper part 
of the tree. Raspopov and Isakov (1935) observed that in the center of the 
forest nests are built higher up than at the border, because trees grow 
higher in the center than at the border. 



309 



357 



The measurements of the nests of Moscow squirrels are as follows: 
length 20-50 cm (generally 35-40 cm); width 25-50 cm (generally about 
35 cm); height 20-40 cm (generally about 30 cm). Average weight of a dry, 
inhabited nest about 450 g. 

These figures show that the nest is generally spherical. The inner 
space is 12-16 cm wide. The entrance to the nest is round and is about 
5-6 cm wide (rarely 8 cm). The entrance is strictly orientated. Nearly all 
entrances in the Moscow region are facing south, southeast or east. This 
is most probably due to the prevalence of cold northwestern and humid 
western winds. 

Raspopov and Isakov (1935) give the following table: 

Entrance Facing NW W SW S SE E 

Number of nests 1 4 8 27 34 52 

The outer layer of the nest consists of branches of the tree on which it 
is built. Branches of other trees are used rarely, when a different tree 
grows close to the tree on which the nest is built. 

In the Moscow Region, the inside of the nest is lined with soft material: 
moss, lichen, dry grass, bark and leaves of birch, oak, aspen, etc., 
occasionally feathers and fur of the squirrel. The preferred lining is 
apparently bark. It is used even when enough moss and grass are available. 
The squirrel strips the bark in an original way. It bites off the tip of a 
twig and, holding it with its teeth, descends along the trunk, peeling the 
bark until it tears off. Raspopov and Isakov observed that the animal takes 
the strips of bark into its mouth in loops and carries them to the nest. 

Inhabited nests are of two types: the nests of the females are larger 
and better built than those of the males. The males usually build their 
nests in abandoned nests of females or on abandoned nests of thrushes, 
magpies and sparrow hawks. The nest is built as follows: The squirrel 
first makes the bottom by interweaving slender and flexible twigs between 
two branches, as support. Then it builds the sides. The top is made of 
strong branches. The nest is reinforced by interwoven twigs. After 
completing the frame, the squirrel begins collecting material for the inner 
lining. The building of the nest takes 3 to 5 days. As the nest of the male 
is built on an existing nest, its building and repair takes only 1 to 2 days. 

One often finds abandoned and unfinished nests. The squirrel ceases to 
build when frightened. It also may abandon a nest if disturbed. 

In the Moscow Region squirrels often nest in hollows in trees, and even 
in starling boxes. 

A. A. Migulin (1924) observed that the Ukrainian squirrel occasionally 
builds its nest on oak trees, using slender twigs from the top together with 
pine twigs. It lines its nest with moss. 

G.K. Gol'tsmaier (1935) gives the following description of the building 
of the nest of the Teleut squirrel: "The squirrel uses different materials 
found in the vicinity. The solid external frame of the nest is usually made 
of pine twigs, which are so firmly interwoven that it is almost impossible 
to pull out one twigwithout dislodging the rest or breaking the nest. It 
appears from the crumpled pine needles and broken twigs that this firmness 
is not attained without effort. . . The outer frame covers the inner, soft part and 
protects it against injury. The inner lining of the nest is made of moss, 
lichen of the genus Cladonia, dry grass, dry leaves (mainly of aspen). 



1254 3,^j 



bark of birch or willow, fur of hares which is commonly found in forests 
because of the capture of hares with nooses, tufts of reed and feathers, . , 
The inner layer of the nest on which the squirrel rests is carefully plucked, so 
that it is always soft. 

"The nest is usually spherical, sometimes ovoid. Its outer measurements 
are as follows: length 50 cm; width 30 cm; height 35 cm; circumference 
100 cm. The cavity of the nest is small as the walls are very thick. The 
inner measurements are as follows: length 20 cm; width and height 10 cm. 
The length of the cavity is not always in the same direction as the outer 
length, but sometimes lies in the short axis of the next. Every nest has two exits. 
The openings are small, and their width (about 5 cm) is adapted to the size 
of the squirrel. Each entrance is carefully concealed by dry grass, moss, 
etc. 

"The Teleut squirrel usually builds its nest on pine and only rarely 
on other trees. . . 
358 "The Teleut squirrel chooses different habitats for its nests. In the 
Srostinsk pine forest, about 50 % of allnests found were in youngforests, 30% 
in clearings and 20% in other forest. The proximity to water is a decisive 
factor in the choice of a nesting site in this pine forest. V, V. Gubar' 
observed that in the Upper Ob pine forest most of the nests are built on 
high trees or in clearings. Proximity of water is apparently not important 
in this forest, as there are many marshes, lakes and rivers, 

"Nests are always built in the upper part of the tree, at a height of 
3.5 to 20m and higher, , , We once found a nest in a hollow the entrance of 
which was only 2,5m above the ground, 

"We heard from local hunters that nests may occasionally be found in 
unusual places, e.g., hollows in felled trees. A hunter from the village 
Srostino stated that he caught two squirrels in a nest on the bank of the 
Sorga, under broken reeds covered with snow. . , 

"Each female has several nests which it uses at the same time. Each 
nest may be used as principal and the others as auxiliary nests . The principal 
nest is built last. In winter it is used as a resting place and as a dwelling at 
night. In summer the young are kept in it. The squirrel uses the 
auxiliary nests only rarely in winter to spend the night and to rest. 
In summer it sleeps in them at night, visiting the principal nest only to 
feed the young. If the principal nest is destroyed the squirrel moves to 
the auxiliary nests until a new principal nest is built. If the principal 
nest is found, it is easy to observe activities of the squirrel, , . 

"There are many reasons for the possession of a number of nests by one 
squirrel. Some of them are possibly taken over from squirrels which have 
been killed. However, most of them are built by the squirrel itself, so 
that it can transfer its young to a new shelter in an emergency, . . The 
squirrel also probably builds a new nest before giving birth, in order to 
rear its young. 

"Nests of males and females are indistinguishable. * During the rut, 
the male apparently loses connection with its territory and nests. He 
uses the nests of the female in her territory. The male generally 
devotes less time to nest building than does the female. This is supported 
by the fact that in summer, the male may rest in old nests of crows. " 



* This differs from the observations of M. P. Raspopov and Yu. A. Isakov, quoted above, on squirrels of 
the Moscow Region. 



311 



359 



360 



In addition to residential nests, the squirrel also builds so-called "shade 
nests". 

G, K. Gol'tsmaier writes: "Shade nests differ from residential nests in 
the absence of a soft inner lining; they consist only of an outer frame. 
They are not as solidly built as residential nests. The squirrel occupies 
them when it eats pine cones. The shade nests therefore always contain 
remnants of food. As they are built only in summer, their purpose is 
probably to protect the squirrel from the heat of the sun, against which the 
pine needles give little protection. The shade nests also presumably hide 
the squirrel from birds of prey, while it feeds. " 

According to K. A. Zabelin (1926), squirrels do not build their nests in 
birds' nests in the Transbaikal forests. The author explains this by the 
fact that there are no magpies in Podlemor'e*, and the crows nest in places 
where there are no squirrels**. "When the nest has no roof, it is usually 
covered by cedar branches. In this case it resembles a crow's nest. Such 
nests are apparently used by the squirrel only in summer. In winter it 
builds a more solid nest. The winter nest has a dome-shaped roof made 
of interwoven twigs. It is generally spherical and closed from all sides. 
It is of the same size as the summer nest. " 

According to G.N. Gassovskii (1927), in the Gilyui-Oldoi hunting territory 
(Far East) squirrels nest close to sources, mainly in spruce groves common 
in the Gilyui-Oldoi Territory. However, squirrel nests are also often found 
in pine groves. 

Squirrels rarely move far outside their territory. S.I. Orlov (1930) reports 
that squirrels entered the tundra, and also that squirrel nests were found in 
alder groves along the Pura river. Frozen squirrels were found in these 
nests in spring. 

FOOD OF THE SQUIRREL. S. V. Lobachev (1932) examined the 
stomach content of squirrels in the Verkhnevychegod District. He found that 
it consisted mainly of coniferous seeds of the tree species in the forest in 
which the animal was caught. In the absence of other food the squirrel may 
eat the bark of trees, e.g., birch bark. Such a diet causes rapid emaciation, 
and reduction of fat and weight. 

M.P. Raspopov and Yu. A. Isakov (1935) made interesting observations 
on the diet of the squirrel in nature. They examined the contents of tree 
hollows in the park of the Agricultural Academy (near Moscow), and found 
the following: l) 650 acorns in a hollow in a linden tree 3,60m above ground; 
the hollow was 40 cm deep; 2) 214 acorns in a hollow in a poplar, 6 m above 
ground (hollow 25 cm deep). All acorns were fresh and none were spoiled. 
From 30 September onwards the authors observed that squirrels stored 
Boletus luteus in the vicinity of the Petushki station of the Gorki 
Railway. They placed the mushrooms into forks at the end of branches and 
near the stem. Squirrels often eat buds of spruce which are covered by a 
membranous cap (especially in early spring). At this time one finds 
trifurcate spruce branches in the snow under the trees. I frequently found 
them in forests near Zagorsk (Moscow Region). Similar observations were 
made by Raspopov and Isakov. 

Squirrels can smell spruce cones buried under 30 cm of snow. The 
crossbills break off large numbers of cones and throw them to the ground. 



* [Area near Lake Baikal. ] 
** Squirrels often use nests of other birds. For example, in the central part of the European U. S. S. R. they 
may be found in nests of thrushes, jays, etc. 

312 



These cones contain many seeds which have not been eaten. The seeds are 
preserved under the snow until summer. This supplies food for the 
squirrels during the time when the new cone crop is poor and only cones 
from the previous crop are left over. 

M.P. Raspopov and Yu. A. Isakov record that squirrels eat Planorbis 
corneus and V i v i p a r a contecta as secondary foods. Squirrels were 
seen to catch the snails on thawed patches near the pond of the Agricultural 
Academy. Pieces of shells were found in starling boxes. 

Observations and experiments on the food of squirrels in captivity are 
important for the study of its diet. Such studies were carried out by M. P. 
Raspopov and Yu. A. Isakov (l935) in the Moscow Zoological Garden. They 
found that squirrels were particularly fond of walnuts, spruce cones and 
buds, pine buds, oats, hemp, apples, Armillaria mushrooms, etc. 
They also ate pine and hazel nuts, acorns, fresh or dry bilberries, pears, 
carrots, lettuce, wheat, peas, lentils, etc., but less avidly. The 
preference for the above foods varied with the seasons. In autumn the 
squirrels accepted hazelnuts, walnuts and pine nuts. In spring they 
preferred carrots, apples and other vegetables and fruits. 

V. G. Stakhrovskii (1932) studied the food of squirrels in open-air cages. 
He determined which mushrooms the squirrels eat, and their preference 
for each species. They eat and store C ant ha r e llus cibarius. 
Boletus edulis. Boletus rufus. Boletus scaber, Pholiota 
mutabilis, Colocera viscosa, Armillaria mellea, 
Lactarius torminosus. Boletus luteus. Boletus 
subtomentosus, and more rarely, Psalliota campestris. 
Squirrels refuse the following mushrooms: Hypholoma 
fasciculare, Hydnum repandum, Lactarius vellerus, 
Amanita phalloides, Coprinus ephemerus, Coprinus 
fuscescens . 

Stakhrovskii also studied the preference of squirrels for berries. They 
are indifferent to honeysuckle, cherry, gooseberry, buckthorn, elder, 
European bird cherry, lily of the valley, wild rose, spindle tree. They 
eat little or reluctantly: raspberry, bilberry, Alpine bilberry. They 
sometimes eat European mountain ash, cranberries, red bilberries, 
bramble and juniper berries. 

Valuable information on the food of the Teleut squirrel is given in the 
work of G.K. Gol'tsmaier (1935). He came to the conclusion that the most 
important food of the Teleut squirrel is pine nuts and mushrooms. The 
squirrel begins feeding on pine nuts in early July, and eats different 
numbers of pine cones per day depending on the season. G. K. Gol'tsmaier 
observed that the greatest number an animal may eat in 24 hours is 190 
pine cones. The average weight of one pine nut is 8.45 mg, and a cone 
contains 10 full seeds on the average. The animal therefore may consume 
16.1 g of dry pine nuts per day, M. P. Raspopov and Yu. A. Isakov made 
similar studies of the Moscow squirrel. They found that the squirrel 
consumes about 55 g of food per day in summer, 70 g in autumn, 35 g in 
winter and 80 g in spring. 

The Teleut squirrel does not open pine cones the seeds of which are 
empty or infested by insects. It is apparently guided by its smell. 

The Teleut squirrel begins to feed on mushrooms in late summer. In 
autumn its diet consists mainly of fresh mushrooms and pine nuts. During 
this period it begins to store mushrooms for the winter. Gol'tsmaier found 



313 



the first dried mushrooms (Boletus luteus)in early August which had 
apparently been collected in late July. 




362 



FIGURE 177. A mushroom stored by a squirrel 

Drawing by A. N. Formozov 

Gol'tsmaier writes: "The Teleut squirrel collects mushrooms with 
stems. It carefully carries one in its teeth to the nearest tree, where it pushes 
in into a fork of two branches or between the stem and a branch. The 
mushroom is arranged so that its pileus lies above two branches or hangs 
down from them. When it is dry it shrinks and surrounds the branch. It is 
so firmly fixed that neither wind nor shaking makes it fall. . . Most 
mushrooms are stored on pine trees, as these are most numerous. A 
smaller number is found on birch, aspen, willow and yellow acacia. . . The 
squirrel chooses forks from 30 cm to 4 m high for this. Mushrooms 
stored low down are later buried by snow. However, they are not lost and 
the squirrel digs them out from the snow. " 

With the onset of severe frosts the squirrel feeds almost exclusively on 
mushrooms. This gives it sufficient food in a short time so that it does not 
need to climb trees to search for pine cones. Gol'tsmaier observed that the 
Teleut squirrel feeds on the following mushrooms: Boletus scaber, 
B. rufus, Boletus luteus, B. bovinus, B. s ub t o m e nt o s u s 
Boletus c h r y s e nt he r e on , B. piperatus, Pluteus nana, 
Clitocyle sp. , Pholiota sp. , Gomphidius glutionosus, 
Ganode r ma sp. 

Gol'tsmaier writes: "The squirrel prefers different mushrooms in 
different years. In 1930 most of the stored mushrooms consisted of 
Boletus luteus, B. scaber and B . rufus, and other species were 
poorly represented. On the other hand, only a few Boletus scaber and 
B. rufus appeared in stores in 1931. Instead there were Go m p h i d i u s 



314 



glutionosus. Boletus luteus and B . bovinus. The favorite 
mushroom in this year was Boletus luteus. This may be deduced from 
the fact that this was the first mushroom to be eaten from the food stores. " 

The squirrel removes the dried mushroom from the branch, holds it 
between its forefeet and rapidly eats it, turning it between its feet. It often 
eats only the pileus, throwing the stem away. Gol'tsmaier did not observe 
how the squirrel eats fresh mushrooms on the ground. To judge by tooth 
marks it seems that the animal does not pluck the mushroom but begins to 
eat it from the pileus. 

According to Gol'tsmaier, the Teleut squirrel eats large numbers of dry 
mushrooms daily, as many as 35. 

The diet of the Teleut squirrel also includes other vegetable foods. 
Animal foods are less common. Squirrels were found to eat the following: 
sawfly larvae, ant eggs (cocoons), butterflies, flies. Only once did 
Gol'tsmaier find remains ofaLacerta agilis in the stomach of a 
squirrel. 

According to E. V. Kozlova-Pushkareva (1933), squirrels of the Hentey 
Range (Northern Mongolia) stored bones of the skull of roe deer for the 
winter. They put the bones on thick branches near the stem of the trees. 
Bones are rich in ash. These stores of bones were made by squirrels 
in years with rich cedar crops. It should be mentioned in this connection 
that in German forests squirrels were found to gnaw the shed horns 
of ungulates (Rimand, 1928). Rimand reports that squirrels gather in 
permanent resting places of roe deer (cf. V.N. Kaverznev, "Squirrels 
and Squirrel Hunting" (Belka i belichii promysel), Moscow 1931, p. 28). 
363 In captivity the squirrel may attack large birds. Gol'tsmaier writes: 
"On 1 August 1931 we liberated a wounded jay (Garrulus infaustus) 
into a cage of squirrels. The bird was immediately caught by a young 
male squirrel who broke its skull and ate its brain. " 

Gol'tsmaier made the following table of the food of the Teleut squirrel in 
various months. 

Months Type of food 

January Mushrooms, pine nuts 

February Mushrooms, pine nuts 

March Mushrooms, pine nuts 

April Mushrooms, pine nuts 

May Mushrooms, pine pollen, animal food, rarely pine nuts 

June Mushrooms, pine nuts, ant cocoons and other animal food 

July Mushrooms, pine nuts, ant cocoons, sawfly larvae and other animal food 

August Mushrooms, pine nuts, ant cocoons, sawfly larvae and other animal foods tree 

leaves 

September Mushrooms, pine nuts, sawfly larvae and other animal food 

October Mushrooms, pine nuts and other vegetable food, ant cocoons, insect larvae 

November Mushrooms, nuts, aspen buds and other vegetable foods, sawfly larvae and other 

animal foods. 

December Mushrooms, pine nuts, sawfly larvae 



The squirrel needs water. Observations of this animal in captivity show 
that it drinks much. In dry years the squirrel keeps close to water in the 
forest. In winter it may satisfy its thirst with snow. 



315 



365 



Thus to summarize the data on food of the squirrel: In coniferous forest, 
the main food is coniferous seeds. ■■' In the oak and nut tree forests, 
acorns and nuts are the main food. For European and most Siberian 
squirrels mushrooms are a secondary food in years with plenty of cones 
and other basic foods. In years with small crops of tree seeds, the 
squirrel feeds on mushrooms, berries, buds and also animal foods. As 
described above, the Teleut squirrel eats large numbers of mushrooms. 
This is due to the poor flora of the pine forests, which gives only a limited 
choice of food, mainly mushrooms and pine nuts. 

The food of the squirrel may change sharply under certain conditions. 
K. Plyater-Plokhotskii ("Vestnik Dal'ne-vostochnogo filiala Akademii 
Nauk", 1935) reports that large numbers of squirrels migrated from forests 
of the Far Eastern taiga to fields of corn and sunflower because of the 
failure of the crop of pine nuts in 1935, Great damage was caused to the 
plantations. The damage caused to corn by the squirrel is typical. It 
climbs on the plant, tears off the leaves enveloping the cob and plucks the 
seed with its incisors. It opens it, eats the germ and discards the rest. 

Detailed work has recently been published on methods of determination of 
age and length of life of squirrels. We have quoted the work of M. K. 
Serebrennikov (1930) and N. P. Naumov (1934) on changes in skull and teeth 
of squirrels. Naumov also studied the structure of the claws. He studied: 
l) the number of grooves on the surface of the claw; 2) the number of 
tubules in the deep stratum (between the superficial and the sterile stratum) 
from the germinal matrix in the last phalanx to the tip of the claw. These 
tubules (called "Ausfiillungshorn" by German authors) are easily seen with 
a magnifying glass through the surface layer of the claw. 

There is no need for histological sections, like in the study of bone. 
It was found that the number of grooves on the claw** (at its base and on its 
sides) increases regularly with age. One new groove is added each year. 
The older grooves lie near the tip of the claw, the younger grooves near 
its base. Examination of a large amount of material showed that there 
are deviations. There is sometimes a correlation between the number of 
grooves and the age of the animal, but in other cases there is no 
correlation. Claws of old specimens frequently have not enough grooves. 
This may be caused by the disappearance of grooves through abrasion. 

Analysis of the number of tubules of the deep stratum did not give clear 
results. The growth of layers of tubules with age could not be exactly 
determined. These characters should therefore be used with caution and 
only as control of more reliable characters like structure of teeth and skull. 

As shown above the age of a squirrel can be more or less exactly 
determined by the rate of abrasion of the teeth. There occur very old 
specimens (8-9 years). The length of life of the squirrel is determined 
differentlyby different authors. Korscheltt claims that the life of the 



* A list of important trees in Russian forests: Picea excelsa Link, P. obovata Ledebour, P. 
jezoensis Carr. , Abies sibirica Ledeb, A, nephrolepis Maxim., Larix sibirica Led., 
L. gmelini, Pinus, Pinus sibirica Mayr. , Pinus pumila (Pall. ) Rgl. , Pinus koraiensis 
Sieb. et Zuce. , Quercus petraea, Q. robur L. , Q. m ongol ic a Fischer, C or ylus avellana 
L. , Juglans mandshurica Max, and others. The seeds of these trees are eaten by the squirrel. 
** The grooves are studied with a mrgnifying glass with a magnification of 10 to ISX. The claw is 
illuminated from different directions, 
t Korschelt, Lebensdauer, Alter und Tod, Jena, 19?4. 



316 



squirrel may last 10-11 years. S.A. Severtsov='= thinks that its length of 
life is 12 years. 

According to N. P. Naumov (1934), only 20-25% of the new generation 
live to the age of one year. There is thus a very high mortality among the 
young: 10-15% reach the age of 2 years; 3-5% — 3 years; 1-2% — 4 years; 
1-0.5% — 5 years. 

Where squirrels are hunted regularly the percentage of animals reaching 
the age of one year and reproducing is lower than that in regions where the 
animal is hunted irregularly. 

N.P. Naumov states that old, experienced squirrels have many 
advantages over the young in regions with irregular hunting; they are more 
able to hide from their principal enemies. They also stand periods of 
food scarcity better than do the young, because of their experience, their 
stores of food, etc. Where huskies are used for hunting, the squirrels 
are found by traces of their activity and animals of any age or sex may be 
caught. The experience of the old squirrel is of no avail against the dogs. 
An equal number of young and old specimens are killed. This eliminates 
the selection in a natural environment, without the activity of man. The 
chances of a squirrel reaching old age are therefore very small in hunting 
territories, and its life is shorter by 2-3 years. Adults which live through 
the winter are 1 to 6 years old and are in their prime. Where squirrels 
are not hunted, about 25% of the adult population is senile. The females 
of this group are probably sterile or remain unfertilized as they mate with 
very old males. This reduces the number of young (N.P. Naumov). He 
writes: "We assume that the large number of senile, sterile specimens 
causes a reduction of the reproduction rate and decreases the percentage 
of young. However, when natural selection is replaced by hunting, the 
population is rejuvenated and the reproduction rate increases. " 

Where hunting is intensive, each pair has about 9 young, while pairs in 
unexploited territories produce only 4 young (N.P. Naumov). 
366 According to Gol'tsmaier male Teleut squirrels are more numerous than 
the females. Of 249 specimens obtained, 56%, were males and 44% 
females. Of the newborn 62% are males, and only 38% females. This 
difference disappears later and the sex ratio becomes about 1:1. According 
to S. V. Lobachev (1932), 49% of the squirrel population were males and 
51% females. The sex ratio was thus 1:1. N.P. Naumov (1934) analyzed 
squirrel populations and found that the sex ratio is about 1:1, with slight 
prevalence of males. 

Most observations show that there are two periods of rut. This, however, 
varies with the locality. For example, in the southern and central parts 
of Russia there are three periods of reproduction (N.P. Raspopov and 
Yu. A. Isakov, 1935). In the north the squirrels have two litters or even 
only one per year. Females have estrus at different periods, according to 
the locality. According to L. P. Sabaneev, estrus begins in late February 
in the Moscow Region (10-25 January according to more recent observations 
by M.P. Raspopov and Yu. A, Isakov). In the Upper Vychegda area 
estrus begins in early February (S. V. Lobachev, 1932). Inthe northern part of 
theTobolarea it begins in the middle of March (Dunin-Gorkavich, 1908). In 
the Ket area of the Narym Territory it begins in late March (Afanas'ev, 



* S. A. Severtsov, "The Relation between Length of Life and Fertility in Various Mammals" (O 

vzaimootnoshenii mezhdu prodolzhitel'nost'yu zhizni i plodovitost'yu raznykh vidov mlekopitayushchikh). 
Izvestiya Akademii Nauk SSSR, 1930. 

317 



1928) and in the Tara District in early March (V.E. Ushakov, 1927). In 
the Turukhansk Territory it begins in late February or March (M. F. 
Krivoshapkin, 1865; P. Tret'yakov, 1869); in the Angara area in the 
middle of February (V.N. Troitskii, 1930); in the Barguzin Territory in 
late January or February (K. A. Zabelin, 1926); in the Anadyr Territory in 
the middle of February (N. Sokol'nikov, 1927). 




ncuRE 178. 



A squirrel nest 

Drawing by A. N. Formozov 



367 Weather certainly affects the onset of estrus. Late spring and frosts 
delay the rut, and good weather hastens it. The physical condition of the 
animal and its food also affect the rut. In emaciated animals, estrus is 
delayed and prolonged. This results in many barren females. In years 
with much food in autumn the animals are well nourished, and the spring 
rut appears early. 

In Siberia strong snowfall and snow on the trees inhibit the activity of 
the squirrels. N.P. Naumov writes: "The squirrel shows little activity in 
winter. It spends the severe frosts in semihibernation. This is followed 



318 



by a period of increased activity caused by strong winds. These winds, 
which come rarely in late January, usually in middle or late February, 
blow the thick snow (which hinders the movements of the squirrel) from 
the branches. The snow may be 5-10 cm thick on the branches, 
especially in the region of the Siberian anticyclone where there is little 
wind in winter. The snow often covers the cones. Observations in the 
Turukhansk Territory (winter 1927- 1928) showed that heavy snow on the trees 
and late appearance of winds forced the squirrels to stay in their nests to 
the middle of February although the temperature was favorable. " 

During the rut the males chase the females in groups of four to six. They 
often fight for a female. The battlefields of males consist of large areas of 
snow covered with tracks. The males are very excited, run about, squeak, 
jerk their tail up and down, chase females and fight. They also joggle their 
jaws, grinding their teeth, and often give high-pitched squeaks like the 
mewing of a cat. They smell and lick the female's urine on the snow. 

The chase takes place on the ground and on the trees. When a group of 
squirrels is chased by a dog, it settles on a tree. The female nearly always 
stays below the males which sit higher up among the branches, move their 
feet up and down and patter. 

In the first estrus in early spring or in late winter the estrus of old 
females appears earlier than that of young females, which breed for the 
first time. That females are chased by several males may be explained 
by the fact that sexual excitement appears earlier in males. This creates 
a temporary surplus of males. The chase itself usually takes place in the 
morning beginning shortly before sunrise and ending towards noon. It often 
begins again towards evening. M. P. Raspopov and Yu.A. Isakov made 
observations on the copulation of squirrels. After an energetic sex play 
the male leaps onto the female, who rises on her legs, folds her ears 
backwards and turns her tail sideways. The male clasps her flanks near 
the hind legs with his forefeet, holds her with his muzzle and copulates. 
Copulation is accompanied by loud grumbles and squeaks of the male and by 
complete silence of the female. Copulation lasts about 15 minutes. At the 
end the male grows weak and hangs helplessly on the female. Blood oozes 
from the genitals of the female after the first copulation. The male covers 
the female about eight times during the day. Fertilization probably takes 
place when the exhausted male hangs on the female. It was observed in the 
Zoological Garden that squirrels rub their genitals against the snow and lick 
the penis after copulation. 

Estrus lasts about 2 weeks. The female lets the males copulate during 
3-4 days, towards the end of the rut. During the other days she runs away 
or drives the males away. 

Recent observations show that the gestation of the squirrel lasts 35-40 
days. 

The second rut begins at different periods after the first. Its appearance 
is influenced by a good food supply. Poor physical condition and little food 
retard the appearance of the second rut. 

V. G. Stakhrovskii (1932) states that the second rut begins 36-37 days 
after the birth of the first litter. The young open their eyes at about this 
period. S. V. Lobachev (1932) observed during the Upper Vychegda 
expedition that the second rut begins when the young of the first litter become 
lndef>endent in the middle of June. The data of Gol'tsmaier (1935) confirms 
Lobac'hev's data. N.P. Naumov (1934) thinks that in years with poor food 



319 



369 




FIGURE 179. Copulation of squirrels 

Drawing by V. A. Vatagin 

supply the birth of the first litter is separated from the second estrus by 
the nursing period of the young, 70-75 days. In years with a good food 
supply this is reduced to 50-55 days. A.N. Formozov made observations 
in the Krasnye Baki District of the Gorki Region, in summer 1932. This 
summer was preceded by a year with a rich yield of spruce nuts. On 3 
July 1932 he caught a gravid female (weight 386 g) with 7 embryos. A 
male (weight 306 g) with very large testes was killed near the female on the 
same tree. Nearby he found two young squirrels, one of which was a 
male (weight 121 g). The female was still feeding the young 
of the first litter, as her teats still contained milk. The embryos 
were 3X2 cm long, and were apparently in the second half of their 
development. This female had certainly copulated while still suckling the 
first litter, probably 20 days after birth. The period between the two 
litters was shortened by 15-20 days in this female. 

Some authors state that squirrels may have a third and even a fourth 
estrus. M.P. Raspopov and Yu.A. Isakov (1935) record that they observed 
periods of rut in the Moscow Zoological Garden: the first in the middle 
of January, the second in the middle and end of March, the third in early 
June and the fourth in early August. The fourth estrus was observed in a 
female Teleut squirrel the third rut of which had taken place in early June. 
As the observations were carried out in a zoological garden, they cannot 
be applied to wild animals. First of all, the squirrels obtained a varied and 
plentiful food supply. This is important, as a good physical condition 
certainly furthers reproduction. Secondly, different geographical forms 
were observed in the zoological garden, central Russian Teleut squirrels 
and Manchurian squirrels. The heterogeneity of the material reduces 
the value of the results. 

Estrus in autumn is rarely observed in squirrels of the same year which 
become gravid for the first time. G. K. Gol'tsmaier (1935) reports that on 
16 November 1931 he obtained a 5-7 -month-old female Teleut squirrel of 
which the teats had been sucked and the niammary glands functioned. If we 
assume that this female gave birth on 15 November, it must have had estrus 



320 



370 



on 10 October. Most females apparently reach sexual maturity only in 
spring of the next year, i.e., at an age of 8-12 months. The estrus of 
these young females lasts t h e whole spring. Yearling females 
usually do not have a second estrus. In Gol'tsmaier 's opinion this 
explains the presence of many sterile females in the second half of summer. 
In the spring of the next year, estrus usually begins and ends earlier in 
2-3-year old females than in one-year old ones. These females also have 
a second, summer estrus in June -July. Gol'tsmaier gives a table 
summarizing these phenomena (see below). 

The table shows that the sexual cycle of the squirrel lasts about 8 
months. During this time one may find females in rut and suckling their 
young. 

Many authors observed that winter nests are often occupied by couples. 
N. P. Naumov found a couple in a winter nest in the Turukhansk Territory. 
Similar observations were made by A.N. Formozov in oak forests in the 
vicinity of Gorki. He found that without exception all squirrels for me d 
couples already in autumn. He also states that the Mar i in the 
central Trans-Volga region make a peculiar squirrel hunt in autumn, 
beating the air with a switch or making noises with a ramrod in the barrel 
of a gun, to lure the animal. These noises resemble the sounds of the 
squirrels during the breeding period. This method resembles the autumn 
hunt of the hazel hen with a whistle. This system uses the fact that the 
hazel hens form couples already in autumn and each is attracted by the 
voice of the other sex (the voices of the male and female hazel-hens differ 
slightly). 

Periodicity of estrus of the Teleut squirrel (Gol'tsmaier) 



^^~->v^_^ Months 


>. 
















E 


<D 


o 

o 






Age ^^"\.,^^ 


B 
a 


1 

(U 


2 


a, 

< 


>• 


_3^ 


3 


3 
< 


a 
00 


> 



2 


a/ 
o 

Q 


Subadultus 
(4.5-12 months old) 
















Estrus rarely 








Adultus 






Estrus 






Estrus possi- 










(1-2 years old) 












ble exception- 
ally 










Adultus 




Estrus 






Estrus 












(2-3 years old) 























Another interesting feature of the sexual cycle of the squirrel is the 
ability of gravid females to hide in the forest. S. V. Lobachev (1932) 
described the second estrus and beginning of gravidity as follows: "it 
seemed as if there were no females at all in the forest. This was clear 
proof of the ability of the gravid females to hide in high and dense spruce 
trees, only rarely descending to the ground. " 

The number of young in each litter varies greatly, five to ten in adult 
females and three to six in young ones. N. P. Naumov ("Material on the 
Reproduction Rate of Squirrels" (Materialy k poznaniyu urozhaya belki), 
Trudy po lesnomu opytnomu delu. Issue VII, 1930, p. 107) did not find more 



321 



371 



than ten young in one nest. According to Gol'tsmaier the number of young in 
each litter is four to six (five on the average), but these data are based on 
little material. The number of young per litter is affected by the age of 
the female and by its nutrition. N. P. Naumov's observations (1934) in the 
Surgut Territory show the connection between the rate of reproduction, i.e., 
the number of embryos, and the nutrition of the female. In this district the 
squirrels suffered severe famine during the winter of 1931-1932 because 
of a very poor pine-nut crop in 1931. Local hunters found squirrels which 
had starved to death in spring. It was found that the stomachs of the dead 
animals were either completely empty or full of conifer buds, Polyporus 
betulinus (a fungus on birch) and lichens (Usnea). The animals were 
emaciated. This poor physical condition delayed the estrus. Resorption of 
embryos also took place. The uterus of a female caught on 24 July 1932 
contained eight embryos. Five of them were of normal size, but the others 
were less than half as large. The resorbed embryos were not in one group 
at the tip of the uterus, but alternated with normal ones. Thus, starvation 
caused a reduction of progeny by 37.5%, by resorption of embryos. According 
to N. P. Naumov, the reduced reproduction of squirrels in summer 1932 
seemed due to E-avitaminosis*. E.M. Vermel' made histological studies of 
organs of squirrels obtained in the Surgut expedition. He found pathological 
degeneration in the liver and adrenals. The changes consisted in an 
increased absolute and relative number of binucleate cells, and in the 
appearance of cells with four and eight nuclei in the liver. This indicates 
a deterioration of the physiological condition of the animal. He also found 
that the adrenal medulla disappeared. 

These histological changes were found only in adult squirrels which had 
suffered from the winter famine. Young squirrels were not affected. 

The development of the young was recently described by V. G. 
Stakhrovskii (1932), M.P. Raspopov and Yu.A. Isakov (1935) and O.K. 
Gol'tsmaier (1935). According to M. P. Raspopov and Yu. A. Isakov, the 
newborn are completely naked, their skin is pink and their eyes show 
in a lilac color through the eyelids. The groups of vibrissae are already 
visible. When the young are taken out they roll into a ball, curl their 
tail between the legs and cuddle up against each other. 

Measurements and weight of the newborn common Moscow squirrel: 



372 



Sex 


Weight in 


grams 


Body 


length 


in 


cm 


Ti 


ail 1 


ength in cm 


Male 


8 

7.5 
8.S 
8 






5,6 
5.6 
5.3 
5.6 










2.6 
2.1 
2.3 
2.4 


" 


7 






5.1 










2.2 



The toes of the newborn are connected. The toes of the forelegs 
separate first, the lateral toes before the inner toes; the inner toe 
separates first. The toes of the hind legs separate in the same order. 
According to M. P. Raspopov and Yu. A. Isakov, the toes of the forelegs 
separate on the 9th day, and those of the hind legs on the 12th day. 



Vitamin E is an antisterility vitamin which stimulates reproduction. It is present in the germs of wheat and 
other seeds, and in vegetable fats. 



322 



The claws of the newborn are white. On the 2nd day pigment spots 
appear on the claws, under the skin and at the base of the claws. 
Translucent blood vessels appear at the tip of the claws. The pigment spot 
at the base of the claws later enlarges towards the tip. The claws attain 
their definite pigmentation towards the 30th day. 

The ears of the newborn are covered with skin and pressed to the head. 
The skin is gradually drawn in, and on the 4th day the ears begin to project 
and the cartilages develop. On the 9th day the invagination of the skin 
continues, and the posterior cartilage projects. On the 18th day the 
anterior cartilage is differentiated, and its margins diverge. On the 20th 
day the ventral cartilage takes shape and the skin becomes so deeply 
invaginated that the auditory meatus is clearly visible on the 2 1st- 22nd 
day. 




FIGURE 180. Eighteen-day- old squirrels 

Drawing by V. A. Vatagin 

Eyelids, eye-slits and eyelashes are well formed on the 27th day. The 
eyes open on the 29 -30th day. * When the eyes open the animals are able 
to climb. 

G. K. Gol'tsmaier (1935) observed that the legs grow irregularly in this 
early stage. They are very short in the newborn. When the animals begin 
to see the legs are very long. This is possibly an adaptation which enables 
them to fall from considerable heights without injury. Gol'tsmaier 
observed such a young animal falling from a height of 5 m onto sand, without 
any harm. 

M. P. Raspopov and Yu. A. Isakov observed that the gums of the mandible 
became swollen on the 13th day. On the 20th day the gums had projecting 
tubercles with well marked blood vessels at the tip. On the 21st day the 
lower incisors broke through in three out of five young. They were 0.5mm 
long. On the 22nd day the lower incisors broke through in the remaining 
two young. 
373 In young Teleut squirrels in the Moscow Zoological Garden the incisors 
broke through on the22-23rd day. Three days later they were twice as 
long (l mm) but were still loose and movable. 



* According to V. G. Stakhrovskii (1932), the eyes of the Moscow squirrels open on the 36-37th day. 
According to Gol'tsmaier, the eyes of the Teleut squirrels open on the 33rd day. 



323 



The upper incisors of Teleut squirrels could be felt through the gums 
on the 30th day. They broke through, however, only on the 37- 39th day. 
In the common Moscow squirrel this was delayed; the incisors broke 
through only on the 41st day. At this stage the lower incisors were already 
6-7 mm long. The molars could be felt through the gums on the 45th day. 
The only hair on the body of the newborn was groups of vibrissae (see above). 
On the 3rd day a few hairs could be seen with a magnifying glass on the spine 
near the base of the tail. On the 4th day, a few pigmented short hairs could 
be seen all over the body with a magnifying glass. These hairs rapidly 
become dense on the occiput. White hairs appear on the belly of 6 -day-old 
animals. From the 8th day the skin takes on a blue -lilac tone in some 
places. The first parts to darken are the tail and the hind soles. The 
pigment in the bulbs of the new hair then spreads to the spine and the whole 
back. At the same time both sides of the feet become dark. On about 
the 20th day the dorsal side of the animal is covered with dense reddish 
hair. On the 30 -32nd day tufts 3mm long begin to grow on the ears. At 
this stage the terminal hair of the tail is 20 mm long. 

It is not known exactly when the animal is weaned. M. P. Raspopov and 
Yu. A. Isakov observed that the young still sucked on the 43rd and even the 
52nd day of life. 

V. G. Stakhrovskii (1932) thinks that the young begin to feed independently 
about the 40th day. This agrees with observations of other authors. The 
animals probably do not stop sucking altogether in the first period of 
independent feeding. This was reported by M. P. Raspopov and Yu. A. 
Isakov. 





FIGURE 181. Forty-day-old squirrels 



Drawing by V. A. Vatagin 



374 G. K. Gol'tsmaier (1935) states that blind newborn tolerate fasting well. 
A one -day fast does not cause any harm, but perhaps retards growth. 

The daily increase in weight of the newborn varies. It is about 1.5 g on 
the average. The curve of growth and increase in weight is irregular. 
According to M. P. Raspopov and Yu.A. Isakov, a 5 l/5-month-old 
Moscow squirrel has the weight and size of an adult. The weight of such a 
specimen was 250 g, and it was slightly heavier than its father. 

G. K. Gol'tsmaier observed that young Teleut squirrels of the spring 
litter reach the size of the adults in 4-7 months. 

M.P. Raspopov and Yu.A. Isakov (1935) observed that 47-day-old 
Moscow squirrels could already climb on trees. After descending to the 

324 



375 



base of a tree, they held on to the trunk with their hind feet and reached for 
food on the ground with their forefeet and teeth. They thus collected pine 
nuts in the open-air cage and bit off shoots ofChenopodium . On the 52nd 
day they fed on pine nuts, strawberries and bilberries. 

Adult females take good care of their young. They sleep with them in the 
nest during the suckling period. In case of danger they transfer them to 
another nest. V. G. Sakhrovskii (1932) saw the following in the 47th section 
of the Pogonno-Losinyi forest. A female, with a litter of open-eyed young, 
clinging to her neck, climbed on a tree and jumped from one tree to 
another. 

Gol'tsmaier states that the female transfers her young from one nest to 
another two or three times. He writes: "Transferring her young, the 
fenaale held its flank between the teeth so that the head of the young was 
under her head and its tail above her neck. The body, head and feet of the 
young were rolled up under the head of the mother, not to hinder her running. 
The female runs on the ground for long distances, when carrying her young, 
climbing trees only in case of danger. The female moves in a short gallop. 
A female who is forced to climb a tree may drop the young. The young 
squirrel then climbs helplessly on the tree, and the female remains 
beneath it, probably to catch it when it falls, " 

Much has been published on the molt of squirrels. The most important 
is an article by Prof. B. A. Kuznetsov, "Fur Structure and Molt in the 
Squirrels" (Stroenie mekha i lin'ka belki.— Izvestiya Assotsiatsii nauchno- 
issledovatel'skikh institutov, I (1-2), Moscow 1928). This work was 
supplemented and corrected in parts in N. P, Naumov's article "Biology of 
the Reproduction of the. Common Squirrel" (Biologiya razmnozheniya 
obyknovennoi belki, in the collection "Ekologiya belki" (The Ecology of 
Squirrels), Moscow 1934), and also by G,K, Gol'tsmaier (loc.cit., 1935), 
M.P. Raspopov and Yu. A. Isakov (loc. cit. , 1935). 

The autumn molt of the squirrel has been studied in detail, but not 
the spring molt, 

N.P. Naumov (1934) came to the following conclusions: 

1. The spring molt starts on the head and proceeds towards the tail. 
The first part to molt is the "spectacles" around the eyes. At the same 
time the molt begins on the occiput along the line connecting the middle 
of the ears. Much earlier [sic], the hair of the tail begins to fall out. 
However, the skin does not yet show any signs of molt. " The molt of the 
head then spreads to the cheeks, forehead and ears. From the head the 
molt spreads to the shoulders and back, along the spine. At this stage, 
the lead-gray or bluish color of the skin of the back becomes rust colored 
due to the appearance of new hair. The molt then spreads to the flanks and 
legs. At this time the molt has not yet begun on the tip of the muzzle and 
the ears. The duration of the spring molt, like that of the estrus, depends 
on the physical condition of the aninaal. The spring molt follows 
immediately after the spring estrus. The males molt first and the molt 
lasts until the middle of the summer. Molt begins later in females, but is 
more rapid, and ends with the birth of the young. 

2. The tail molts once a year. The other fur molts twice a year. The 
molt of the tail is very prolonged. In the first molt period** (at the end of 



* This is generally true of all molting parts of the body, in which the appearance of dark spots and patterns 
on the skin is preceded by loss of hair. 
** Before the beginning of the spring estrus. 

325 



winter) part of the ordinary and guard hairs and nearly the whole underfur 
fall out. The fur of the tail becomes stiff. The molt of the tail begins after 
an interruption (resting stage) in the second half of June. It begins in the 
middle of the tail and continues towards the tip and the base. 



•/•»5-*V!^ 




FIGURE 182. Diagram of sequence of spring molt of the 
squirrel 

After M. P. Raspopov and Yu. A. Isakov 

3. The autumn molt begins at the root of the tail. The first stage is the 
appearance of small, isolated groups of hair forming patches or stripes on 
the buttocks near the tail. The molt intensifies and the area at the base of 
the tail becomes larger, cordiform and of a uniform bluish black color. 
This area is surrounded by a zone of lighter color which gradually becomes 
pale towards the periphery. The spot then widens towards the head in 
two stripes (B. A. Kuznetsov). The area between these two stripes does 
not molt for some time. At the same time the hind legs begin to molt. The 
autumn molt ends on the nose, occiput and forelegs. It is not the same in 
the two sexes. The males molt first, immediately after the heat. Females 
begin molting at the end of the breeding period. They molt, therefore, much 
later than the males. Adults molt in two stages. Females in their second 
pregnancy begin to molt about one month later than those which do not 
reproduce a second time. These differences make it possible to determine 
the number of sterile females which do not become gravid a second time. 
In autumn sterile females molt soon after the males. 

Molt in young squirrels of the same year. After leaving the nest the fur 
of the young is delicate, short and sparse, but it grows gradually and 
becomes denser. The thin and easily torn skin becomes tougher and 
thicker. In summer the color of the skin becomes blue because of the 
appearance of new hair and the growth of the present hair, but there is no 
molt. The fur on the lower part of the back and the tail reaches its full 
length first. This continues towards the head, and ends on the occiput and 
the forehead. While the fur of the body reaches its full length the fur of the 
tail molts. Before molt the fur of the tail is short, luxuriant and soft. 
After molt it becomes dense, long, with well developed guard hairs which 
give thickness and the typical bilateral "combing" of the hair. In the second 
half of August the fur of the young of the first litter reaches its full length, 
and the molt of the tail ends in early September. The molt of the young 



326 



begins in September and follows the same sequence as the autumn molt in 
adults. It seems impossible to determine a difference in the molt periods 
of young males and females. The molt of the young squirrels to some 
extent lags behind that of adult males and considerably precedes that of 
adult females. 




FIGURE 183. Diagram of the sequence of autumn molt in the 
squirrel 

After M. P. Raspopov and Yu. A. Isakov 

G. K. Gol'tsmaier records that on 16 November he found a young female 
of the first litter with complete summer fur and clear skin. Her mammary 
glands secreted milk, as she was gravid in her first autumn. Pregnancy 
apparently retarded the molt in this female. 





Molt in young 


Teleut squirrels 




Stage of molt 


Age in months 


Date of capture 


Sex 




1 ^ 


19 Sept. 


female 


Beginning .... 


5 
1 2.5-3 


24 Sept. 
9 Sept. 


male 
female 




J 3 


13-15 Sept. 


female 


Middle 


1 4.5 
) 3.5 


9 Oct. 
19 Oct. 


male 
male 




] 5.5 


31 Oct. 


male 


End 


4.5 


1-4 Nov. 


female 




3 


18 Nov. 


male 



N. P. Naumov (1934) studied molt periods and molt in young squirrels of 
the second litter. He found that they molt rapidly, as they have little time 
until the beginning of frosts. Unlike the first litter, they do not pass 
through a long period in which they attain full size and grow new hair. 
Squirrels of the second litter thus begin the autumn molt immediately. 
N. P. Naumov observed that body and tail molted at the same time in some 
cases. This never happens in adults or in young of the first litter. 

As stated above, duration of molt depends on the physical condition of the 
animal and on its sex. Males begin to molt earlier. The skin of the 
scrotal region becomes blue even during estrus. However, molt is more 



327 



378 



prolonged in males than in females. In males it often lasts until the second 
estrus, sometimes even including estrus. The females begin molting later, 
but the molt is more rapid and ends with the birth of the young. 

M. P. Raspopov and Yu. A. Isakov observed molt in squirrels of the 
Moscow Zoological Garden. ■•' They found that the spring molt lasts 2 l/2- 
3 months, and the autumn molt 1-3 months. 

G.K. Gol'tsmaier (1935) gave a table (p. 327) which shows that the older 
the animal the sooner molt begins and ends, and the younger the animal 
the later it molts. 

G. K. Gol'tsmaier found that at all ages autumn molt begins in August 
and ends in December. In the following table he arranges the molting 
period in age groups, without distinguishing between sexes: 



Molt in Teleut squirrels 



Age 


Months 


August 


September 


October 


November 


December 


Juvenis 




Molt begins in the 
second half of the 
month 


Molt 


End of molt 


End of molt in 
late litters 


Subadultus 


- 


Molt begins in the 


Molt 


Molt ends in the 


Molt of females 






first half of the 




second half of 


born in autumn 






month 




the month 




Adultus 


Molt begins in the 


Molt 


Molt 


Molt ends at 


Presumably, end 




last days of the 






about the 20th 


of molt in 




month 






of the month 


females bom in 
autumn 



Squirrels migrate in various seasons. The most intensive migrations 
take place in late summer and in autumn. These migrations were recently 
described by N. P. Naumov (1930, 1934), S. V. Lobachev (1932), and 
especially by A.N. Formozov (1934, 1936). 

Fluctuations in numbers of squirrels have been known for a long time by 
hunters and have been described in the zoological literature. The tables 
in N, P. Naumov's work "Periodicity of Fluctuations in Numbers of the 
Common Squirrel" (Periodichnosf v kolebaniyakh chislennosti 
obyknovennoi belki, in: "The Ecology of the Squirrel" (Ekologiya belki), 
Moscow 1935, pp. 27-31) show clearly that these fluctuations may cover 
large areas. On the other hand, migrations consist only in a regrouping 
of populations on relatively small areas of their range, not more than some 
hundreds of kilometers. A.N. Formozov (1935) writes: "Fluctuations in 
numbers and migrations are related, but the latter are of a subordinate 
character. Mass migrations are a result of mass reproduction. An 
increase of the squirrel population may result from immigration of animals 
from other areas, and a decrease of the population may be caused by 
emigration of animals. Reduced numbers of squirrels are a result of 
379 epizootics and a low reproduction rate. These two are the result of famine 
after crop failure. Reduced reproduction is thus the reaction of the 
population to unfavorable conditions. The migrations, which are fatal for 



* Without distinguishing between the sexes. 



328 



most migrating animals, are a similar reaction. They are a rapid and 
immediate response to adverse conditions. . . Epizootics are a delayed 
response. After crop failure, sick squirrels are usually found in late 
autumn. The greatest numbers of dead, frozen and weak animals are found 
in the middle and the end of the winter. Squirrels are completely absent 
or occur in very small numbers a year after a crop failure, i.e. , in the 
next hunting season, which may be a season with plenty of food. " 

Fluctuations in numbers and migrations of squirrels depend largely 
on the food supply. It was found that years with plenty or with very few 
coniferous seeds occur with a certain periodicity. Years of plenty are 
followed by a rest period of the conifers, which may consist of several 
years with poor crops. The squirrel populations show the same periodicity. 
It should be stressed that multiplication of squirrels is preceded by rich 
crops of coniferous seeds. Data on the periodicity of crops of coniferous 
seeds have been published. In the Moscow Region spruce crops recur at an 
interval of 3 years, in the Leningrad District every 4 years (A.N. 
Formozov, "Fluctuations in Numbers of Commercial Animals" (Kolebaniya 
v chislennosti promyslovykh zhivotnykh), Moscow 1935, pp. 39-40). The 
cedar crops have a different periodicity which varies with the locality. 
According to A.N. Formozov (1935), cedars yield crops every 3-4 years in 
the western part of their range and in the central part of the Siberian taiga. 
Rich crops are rare in the Narym Territory and in general in the northern 
taiga. They recur at an interval of 7-8 years, according to some authors 
every 10 years. In the European part of its range, the Siberian larch gives 
crops every 6-7 years, possibly every 7-10 years. Pines give crops 
frequently in the Moscow Region, almost every year. In the central part 
of the U.S.S.R. rich pine crops recur every 3-5 years, in Lapland every 
10 years, at 68°30 N every 10-20 years, and in Prussia every 3 years. 

The periodicity of coniferous crops is connected with climatic and soil 
conditions, insolation rate, density of forests, etc. Regions with a mild 
climate, good soil and adequate insolation, e.g., forest borders, have 
rich crops and smaller intervals between the years of plenty. Different 
regions with the same conifer species have a different crop periodicity. 
Some isolated areas in different parts of the range of a species of tree 
may have similar crops at similar time intervals. A.N. Formozov (1935) 
proposes to call such areas "regions of similar productivity". 

If the crop periodicity of conifers in a district is known, years with 
many squirrels, famine years and migrations can be predicted accurately. 
380 There are also accidental, secondary factors which may affect the 

squirrel population, e.g., climatic phenomena of some years such as late 
or early frosts, rainy spring, summer or autumn, sever winter frosts, 
summer droughts, etc. 

Natural calamities like forest fires, mass infestation of seeds by insects, 
etc., have also to be mentioned. Such events may result in movements and 
migrations of squirrels. * 

A.N. Formozov (1933) observed that low reproduction and local 
migrations of Siberian squirrels (caused by poor pine -nut crops) occur 
together with migrations ofNucifraga caryocatactes 
macrorhynchos to Europe. Migrations of N u c i fr a ga c. 
macrorhynchos to Europe make it possible to predict exactly reduced 



* Summer migrations of squirrels are usually caused by drought or forest fires. 



329 



squirrel populations in the northern Urals and Siberia. In years with poor 
pine -nut crops this bird appears in the southern Urals and in the central 
Volga Region in August. The Siberian squirrel population can thus be 
predicted 2-2 1/2 months before the squirrel hunt. 

Loxia curvirostra, Loxia pytyopsittacus and L o x i a 
leucoptera bifasciata also migrate after the failure of spruce and 
pine crops. Reduced squirrel populations and migrations in regions with 
poor crops can be predicted by the migrations of these birds. 

Squirrels generally migrate in a "broad front". Such a front was 300km 
long in 1928 in the lower reaches of the Amur, 100 km long in 1917 along 
the Northern Dvina and 300 km in 1930 in the Leningrad Region. Migrating 
squirrels never move in compact groups, like reindeer herds. They move 
singly, sometimes out of the sight of their fellows. The squirrels become 
more crowded when the animals cross obstacles, e.g., rivers. The 
size of the flock affects the migration rate. The greater the number of 
animals, the quicker the available food is consumed, and the shorter the 
stay in a locality. The rate of migration is 3-4 km/hour on the average. 
The time in which migrating squirrels pass a given point can be determined 
by observation points which are very few in the Soviet Union. In 1917 the 
squirrels migrated through the Northern Dvina region for more than 45 
days (A.N. Formozov). In 1932 they migrated across the Ob (the Surgut 
Territory) for more than 2 months (N.P. Naumov and I.D. Kiris). 

Migrating squirrels often surmount considerable obstacles. They then 
become less timid and more excited than usual. They swim across rivers 
„ and lakes on their way. Squirrels have been observed to cross the Klyaz'ma, 
Selendzha, Msta, Volkhov, Vyatka, Kama, Northern Dvina, Sos'va, Yenisei, 
Amur rivers, etc. According to B. M. Zhitkov (1935), amass migration to the 
northeast was observed in the Sos'va District (east of northern Urals) on 
4 June 1935. The squirrels swam across 1 km of water. Many of them 
drowned. Squirrels are not stopped even by such wide water bodies as the 
Tatar Strait, Taz Bay, and Chivyrkui Inlet of Lake Baikal, which is about 
12 km wide. 

Migrating squirrels which swim across rivers may climb on rafts, boats, 
barges and steamers for rest. They may enter large towns like Kirensk, 
Tyumen', Petrozavodsk, Krasnoyarsk, Tomsk and even the outskirts of 
Moscow. 

It is usually difficult to determine the distances covered by the migrating 
animals. Squirrels have been observed to migrate 250-300 km beyond their 
usual range. According to S.I Orlov (1930), squirrels never occurred in the 
Taz District (67°20' N and 49°50'E). However, large numbers of squirrels 
migrated along the Taz in autumn 1923. They reached the Taz Bay and 
attempted to swim across. This attempt proved fatal because the animals 
were exhausted from the long journey and starvation in the tundra (the 
dwarf larch trees of the tundra did not supply sufficient food). Orlov adds 
that in spring 1924 he found several squirrel nests in dwarf alder forests. 
However, the squirrels in them were frozen to death. 

In 1932 squirrels appeared on the Kanin Peninsula, about one degree 
north of the forest limit. 

The migrations of squirrels to Kamchatka are noteworthy from the point 
of view of zoogeography. It has been generally accepted that no squirrels 
live or ever lived in Kamchatka, as the way to it is barred by the tundra. 



330 



According to P.T. Novograblenov (1930), a report was received in 
Petropavlovsk in 1920 that squirrels had appeared in northern Kamchatka. 
No such reports were received before that date. The appearance of 
squirrels in Kamchatka was received with great surprise by the hunters, 
and the animal was regarded as a rarity. In winter 1923-1924 it was 
definitely established that squirrels had invaded Kamchatka. This migration 
was preceded by increased numbers of squirrels in northeastern Siberia, 
probably because of rich food crops. The squirrels apparently entered 
Kamchatka from Gizhiga and Penzhina*, through the valleys of the Talovka, 
Pustaya, Lesnaya, Palan and Tigil' rivers. During this migration, the 
squirrels crossed the so-called Parapol'skii Dale, a broad tundra area 
covered with mountain-pine thickets. From the valley of the Tigil' river 
they continued through the valley of the Anaun and Kozyrevka rivers, and 
finally reached the valley of the Kamchatka river. The animals moved up 
and down this valley until they reached the valley of the Avacha river. 
According to unpublished data by S.N. Barmantov (1932), squirrels were 
„„ found along the Dvugortochnaya and Kamchatka rivers after their occupation 
of the peninsula. Many specimens were found in the region of Mil'kovo- 
Nachiki. The squirrels then slowly began to appear in the Petropavlovsk 
District, e.g., in the villages Koryaki, Elizovo, Nalychevo, and Nikolaevka. 

In the final stage of long migrations the squirrels lose their purpose. 
As remarked by A.N. Formozov, they move in a state of abnormal 
excitement and sometimes arrive at very barren places. Such migrations 
may end with the death of the whole population. 

Squirrels do not hibernate and are active even during severe frosts 
(according to G. K. Gol'tsmaier's observations in West Siberia). In East 
Siberia, e.g., in the Turukhansk Territory, the squirrels keep to their nests 
and remain inactive during the second half of winter, when the severest 
frosts occur. The same is sometimes observed in the central and northern 
parts of European U. S. S. R. 

P. A. Manteifel' (1927) states that squirrels are never seen outside their 
nests during severe frost*, they stay in them in a semi-somnolent state. 

The squirrel has many enemies. In eastern Europe it is persecuted 
by the pine marten, which attacks it at night in its nest. In Siberia the 
squirrel is hunted by the Siberian ferret. In northeastern Siberia squirrels 
are possibly suffocated in their nests by ermines (S,I. Ognev, 1926). 
According to V. lokhel'son (1898), hunters in the Kolyma Territory state 
that squirrels are attacked by flying squirrels. lokhel'son writes: "During 
the fight the flying squirrel spreads its parachuting membrane. The 
squirrel bites into this membrane and tears it, but the flying squirrel wins 
just like the ermine." 

This doubtful observation was recently discussed by G.K. Gol'tsmaier 
(1935). He writes: "The hunters of the Srostinskii pine forest consider the 
flying squirrel an enemy of the Teleut squirrel and believe that it eats 
both adult and young squirrels, and then occupies their nest. Hunters of 
the Verkhneobskii pine forest state that the flying squirrel only occupies 
nests of squirrels, but does not cause them any harm. Our observations 
do not confirm that the flying squirrel attacks the squirrel, but the flying 
squirrel certainly occupies its nests. Almost all flying squirrels caught 



I.N. Malyshev (1936) thinks that the squirrels entered Kamchatka mainly through the valley of the 
Penzhina river. 



331 



383 



384 



in the Srostinsk pine forest (in which they are numerous) were found in 
squirrels' nests. The question is whether the flying squirrel drives the 
squirrels out, or settles in abandoned or auxiliary nests. The fact that we 
found a flying squirrel in a freshly built squirrel nest (on 24 August) is 
in favor of the first alternative. " 

These ecological aspects of the biology of squirrels and flying squirrels 
should be verified. 

The most serious enemy of the squirrel is the goshawk. It is also hunted 
by sparrow hawks, kites and various owls, e.g. , tawny owl, Ural owl and 
eagle owl. Crows and magpies are dangerous to young squirrels which have 
left their nest but do not yet climb well. 

The squirrel is of great commercial value. The yearly production of 
squirrel furs constitutes about 30% of the whole Russian production of furs 
(B.A. Kuznetsov, "Science of Staple Commodities" (Tovarovedenie), 
Moscow 1932, p. 335). According to A. Emel'yanov ("Pushnoe Delo", 
No. 1/2, 1927, p. 16), an average of 15 million squirrel pelts were 
processed yearly. In the late 1920's about 10 million pelts were exported. 

The squirrel hunt is called in Russian "belkovan'e" or "belkov'e", and 
squirrel hunters are called "belkovshchik" . Squirrels are usually hunted 
with dogs. Amateur hunters in the central part of European U.S.S.R. 
often hunt with mongrel dogs, claiming that they are better suited for 
hunting mammals and birds. The object of the dogs is to find the squirrels 
and bark when they climb the trees. When the squirrel hunters begin the 
hunt, they send the dogs to look for the animals. Sometimes the squirrel 
does not fear the barking and even looks at the dog with curiosity. However, 
it sometimes becomes frightened and passes to other trees as when the dog 
stands up and scratches the bark of the tree. Experienced dogs usually sit 
calmly some distance from the tree and try not to lose sight of the victim. 
The animal is sometimes so well hidden that it is not visible. The hunter 
then waits until the animal shows itself by a movement, or he beats the 
stem of the tree, forcing the squirrel to move. Huskies are especially 
suitable for this hunt. Squirrels are usually shot with shot guns or small- 
bore rifles. 

Squirrels are rarely caught with traps. The only type of trap suitable 
for squirrels is the "plashka". It consists of 2 boards one above the other. 
These boards are fixed so that they form an acute angle. The lower board 
is fixed in notches between the trunks of two trees. The movable heavy upper 
board is raised and held by catches and a horizontal piece with a bait. The 
"plashka" is usually fixed 1-1.5 m above the ground. When the squirrel 
touches the bait the upper board drops and crushes it. Squirrels are also 
trapped by crossbars. Branches are split and the two halves are pulled 
apart and held by catches. When the animal touches the bait, it operates 
the catch and is caught by the closing halves of the branch. 

Fleas (Ctenonotus octodecimdentatus) and ticks are 
ectoparasites of the squirrel. Its intestinal parasites include tapeworms and 
the following nematodes: Syphadia obvelata, Viammia scrjabini, 
Strongyloides and some species of Heligmosominae (Family Trichostrongy- 
lidae), etc. 



332 



Subspecies of the squirrel Sciurus vulgarus 

359. Sciurus vulgaris ognevi Migulin (1928). Central Russian 
squirrel 

1895. Sciurus vulgaris Satunin K. A. , Pozvonochnye Moskovskoi gubernii, Mlekopitayushchie 

(Vertebrates of the Moscow Province. Mammals), p. 10. 
1913. Sciurus vulgaris subsp. Ognev S. I., Mlekopitayushchie Moskovskoi gubernii (Mammals of the 

Moscow Province), pp. 265-281. 
1925. Sciurus vulgaris v a rius Turov S. S., Mlekopitayushchie Ryazanskoi gubernii (Mammals of the 

Ryazan Province), p. 59. 
1927-1928. Sciurus vulgaris ogne vi Migulin A. A., Obzor gryzunov Ukrainy (A Survey of Ukrainian 

Rodents), "Zakhyst roslyn", No. 3-4, pp. 12-13. 
1929. Sciurus vulgaris Kuznetsov B. A. , Zametki o faune mlekopitayushchikh Pogonno-Losinoostrovskogo 

lesnichestva (Notes on the Mammal Fauna of the Pogonno-Losinoostrovskii Forestry), Trudy po lesnomu 

opytnomu delu, Tsentral'naya lesnaya opytnaya stantsiya, Issue VI, p. 107. 

TYPE LOCALITY AND DEPOSITION. Former Borovsk County of the Kaluga Province. Type in Kharkov 
Central Station for Plant Protection. 

MATERIAL EXAMINED: 155 specimens. 

DIAGNOSIS. Winter fur of two types: 1) spruce -inhabiting squirrels: 
gray with a buff tinge. Spine and lower part of back often with brownish 
rust tones ("bald-spine"). Tail of the brown or dark brown type (see above); 
2) pine -inhabiting squirrels: back light ash-gray with a buff tinge. Bald- 
spine usually well marked, rust-red. Tail of the red type. Red-tailed 
squirrels constitute 25-30% of all squirrels. 

The color of summer fur on the back varies from intense red-chestnut- 
brown (in dark brown-tailed specimens) to light ocher-reddish rust (in 
light red- tailed specimens). 

MEASUREMENTS small: length of skull (Figures 165, 168, 184) 47.2- 
53.2 mm (M. 51.2 mm), condylobasal length 42.8-48 mm (M. 46.4 mm); zygomatic 
width 29. 8-33. 4mm (M. 31.5 mm); length of upper tooth row 8.3-9.9 mm 
(M. 9.2mm); hind feet 54-61 mm (M. 57.8mm); tail 160-190 mm 
(M. 173.1mm). 

DESCRIPTION. Color of winter fur of the two types as described. 
Many transitional colors occur. 

1. Spruce -inhabiting squirrel, dark type (see Plate VI, lower figure). 
Back gray with a buffy tinge (mouse -gray). This tone sometimes 
approaches ash color. Dark gray mottling always well marked. Spine and 
lower part of back often with brown-rust -gray tones (bald-spine) on tips 
of underfur and broad subapical bands of guard hair. 

Buff tones always well marked on head, nose and cheeks. Lower parts 
of cheeks and chin pale buff-gray color passing gradually into the white 
„_j. color of the throat. Base of ears usually brown- gray-buff. Two thirds of 
the ears, including the tip are black-brown. Feet and part of antebrachium 
ocher-rust (between orange -cinnamon and sayal-brown). Belly and groin 
white. Squirrels of this type belong to the chestnut-tailed and dark-chestnut- 
tailed type. Dorsal side of tail black-gray with a slight rust tinge, which is 
well marked on the terminal hair. Hair roots dark ash-gray followed by 
buff- rust colored bands and black tips. Terminal hair of tail with buff -rust 
colored tips. Middle of ventral side of tail ash-gray, darkened by blackish 
hair tips. Sides of ventral side with buff-rust colored tones caused by the 
subapical bands. 



333 




s > 

'S <• 



334 



386 




nCURE 184. Skull of 
Sciurus vulgaris 
o g n e V i Migulin, No. M. 
1300, Moscow Region 
(former Moscow Province) 



2, Pine -inhabiting squirrel, light type. Back light ash-gray with a 
buff tinge (light mouse-gray); spine and lower part of back with "bald spine" 

consisting of reddish rust tones. Back with dense 
mottling of a darker gray. Rust tones present on 
head, cheeks and forehead. Ears light ocher-red 
(between orange -cinnamon, mikado -brown and 
tawny). Feet and antebrachium of forelegs reddish 
rust colored. Tail of the red-tailed type. Dorsal 
side of tail reddish brown-ocher (between orange - 
cinnamon and tawny). Hair roots on dorsal side 
light ash-gray, with brownish ocher-red apex. 
Some hairs have a black apex. The middle of the 
ventral side of the tail has well-marked ash-gray 
tones. Sides of ventral area with a broad stripe 
caused by rust-red hair tips. 

The summer fur (Plate VI, upper figure) also 
shows two types connected by transitions. 

In squirrels of the dark type, back red-brown- 
chestnut (between kaiser -brown and tawny). Nose 
and cheeks reddish rust-buff of varying intensity. 
Occiput with darker brownish tones like in the 
middle of the back. Flanks and legs more pure 
reddish rust (between ferruginous and tawny). 
Throat, chest, belly and groin white. Sides of 
throat pale buff-rust, little differing from white middle part of throat. 
The white throat area of the summer fur of most squirrels is wedge- 
shaped anteriorly. Tail of the dark chestnut type. Roots of hair on tail 
light, buff- gray, followed by narrow blackish or black-brown bands, 
broader rust-buff subapical bands and still broader black or black-chestnut 
tips givingablack-chestnut tone tothe dorsal side of the tail. Middle of ventral 
side of tail dirty rust-gray. Sides of ventral area with a fringe of reddish- 
rust and black- chestnut tones. Margins of tail with a black- chestnut tone. 

In specimens of the light type, the whole back (from the occiput) lighter, 
ocher-red-rust (between orange -cinnamon and tawny). Forehead and 
cheeks with buff -yellow shades near ochraceous -tawny. Flanks rust- 
orange (near zinc-orange). Tail of red-tailed type. Dorsal side generally 
red-ocher-rust (between orange -cinnamon and tawny). Hair roots light gray- 
buff, followed by narrow black- gray bands, and rust -red tips which are 
yellower proximally and redder and more rust colored distally. Middle 
of ventral side of tail light ash-gray, flanked by ocher-yellow tones. Sides 
of ventral area with a broad ocher-red-rust fringe. 

GEOGRAPHICAL DISTRIBUTION. Throughout the central part of 
European U.S.S.R. The northern boundary apparently follows the line 
Luga - Novgorod - Kostroma - Perm. The western boundary follows the 
line Pskov - Velikie Luki - Torzhok - Vyaz'ma - Kaluga. The southern 
boundary begins in the southern part of the former Kaluga Province, 
continuing to Tula - Ryazhsk - Penza - Syzran', and turns north along the 
right bank of the Volga Belebei - Yelabuga. Near Birsk (on the Belaya river) 
the boundary meets that ofS. v. bashkiricus. The eastern boundary 
turns north along the Kama to Perm. 



335 



360. Sciurus vulgaris fedjushini Ogn, (1935). Belorussian 
squirrel 

1929. Sciurus vulgaris Fedyushin, A. V. , Dinamika i geograficheskoe rasprostranenie okhotnich'ei 

fauny BSSR (Population Dynamics and Geographical Distribution of Game Animals in the Belorussian 

S. S. R. ), Minsk, pp. 55-58 (partim.'). 
1935. Sciurus vulgaris fedjushini Ognev S. I. Sistematicheskii obszor belok, vodyashchikhsya v 

SSSR (A Systematic Survey of the Squirrels of the USSR), Byulleten' nauchno issledovatel'skogo instituta 

zoologii Moskovskogo universiteta, No. 2 p. 43. 

TYPE LOCAUTY AND DEPOSITION. No. M.3657, 22 October 1929. Pleshchenitsy Forestry of the 
Minsk District, A. V. Fedyushin. S.I. Ognev's collection. 

MATERIAL EXAMINED: holotype and 68 paratypes. 

DIAGNOSIS. Winter fur of two types. Squirrels with dark fur 
predominate. Back gray with chestnut and buff shades which are darker 
than in dark Moscow winter specimens. Grayish brown bald-spine usually 
well developed. Tail of the chestnut or dark-chestnut type. Winter fur 
of the light type usually darker than the light type of the Moscow squirrel. 
Gray tone with a dirty-buff tinge. Tail of the light chestnut or red-tailed 
type. 

Summer fur of a saturated reddish brown, much darker than in 
S. V. ognevi. 
387 MEASUREMENTS: Length of skull 51 .6-53 mm; condylobasal length 
48-49 mm; zygomatic width 31.7-32.5 mm; length of upper tooth row 9.6- 
10.5 mm; length of hind foot 57 mm. 

DESCRIPTION. Squirrels of the dark type predominate. Back dirty- 
gray (between mouse -gray and deep mouse -gray, occasionally with a neutral 
gray shade), darker than in dark S . v . ognevi . Well marked gray-brown 
mottling on back. Many specimens show bald spine, i.e., well marked 
grayish brown tones on spine and lower part of back (between olive -brown 
and natal-brown). Flanks frequently with a buff tinge (buff-brown) which is 
also well marked on nose and cheeks, mixed with wood-brown and buff- 
brown tones. Base of ears brown, and the other parts chestnut-black. 
Soles chestnut -gray (between hair-brown and benzo-brown) with a rust 
tinge. This color is also marked on antebrachia and shoulders. Tail of the 
dark-chestnut or dark type. In tails of the dark type, the brown bands of 
the hair are almost absent and the tail is generally grayish black, with 
broad gray roots of the long guard hairs. Fur coarser and less silky than 
that ofS. V. ognevi. 

Squirrels of the light type are red tailed or light -chestnut tailed, 
differing from the Moscow pine -inhabiting squirrels in the darker gray 
color of the fur. Bald spine well marked. 

Color of summer fur usually more intense than in most specimens of 
S . v . ognevi . Back saturated reddish chestnut (dark kaiser-brown). 
Nose, forehead and cheeks buff- gray (hair -brown) with a rust tinge and 
dark gray mottling. Flanks and legs richer red-rust colored. Chin and 
lower part of cheeks gray-buff (light drab) gradually passing into white in 
middle of throat. In red-tailed specimens dorsal part of tail of one color. 
In chestnut -tailed specimens tail with a marked black tinge, making the 
tail appear nearly black. 

SYSTEMATIC NOTES. Although there are no skull measurements, this 
subspecies seems well characterized. Series differ from S. v. ognevi 
as follows: 1) darker tone of winter fur; 2) color of summer fur more 
intense; 3) lower percentage of red-tailed specimens; 4) fur coarser and 



336 



less silky. Measurements larger than those ofS. v. ognevi. This, 
however, should be confirmed. 

GEOGRAPHICAL DISTRIBUTION. Not well known. The subspecies 
apparently occurs in the forests of Minsk, Polotsk and Mogilev. In the 
north it occurs to about the Velikie Luki, in the east to the line Smolensk - 
Vyaz'ma - Roslavl'. These boundaries are not certain and should be 
verified. 

361. Sciurus vulgaris formosovi Ogn. (1935). Northern squirrel, 
or Formozov's squirrel 

1925. Sciurus vulgaris Shchegolev V. N. , Fauna, biologiya i ekonomicheskoe znachenie 

mlekopitayushchikh Cherepovetskoi gubernii (The Fauna, Biology and Economic Value of Mammals in 

the Cherepovets Province), p. 18. 
1925. "Belka" ("Squirrel"). Milovanovich D, , Okhotnichii promysel v Perchorskom krae (Commercial 

Hunting in the Pechora Territory), "Okhota i okhotnik", p. 43. 
1930. Sciurus vulgaris Lobachev S. V. , Obzor okhotnich'ikh promystov Vyatskogo kraya ^A Survey of 

Commercial Hunting in the Vyatka Territory), Trudy po lesnomy opytnomu delu, Tsentral'naya lesnaya 

opytnaya stantsiya, Issue VII, p. 72. 
1932. Sciurus vulgaris Lobachev S. V. , Verkhnevychegodskaya ekspeditsiya (The Upper Vychegda 

Expedition), pp. 97-147. 
1935. Sciurus vulgaris formosovi Ognev S. I. , Sistematicheskii obzor belok, vodyashchikhsya 

v SSSR (A Systematic Survey of the Squirrels of the U. S. S. R. ), Byulleten' nauchno-issledovatel'skogo 

instituta zoologii Moskovskogo universiteta, No. 2, p. 44. 

TYPE LOCALITY AND DEPOSITION. No. 7, female, 11 November 1930, Yakovlevo Village of the 
Sharya District of the Gorki Region (former Nizhnii Novgorod Region), A. N. ForMozov. Type in S. I. 
Ognev 's collection. 

MATERIAL EXAMINED: holotype and 109 paratypes from adjacent districts. 

DIAGNOSIS. Winter fur of one type. Back pure gray with well marked 
dark gray, dense mottling. Bald-spine rare. Tail of the chestnut tailed 
or dark chestnut -tailed type. Transitional specimens between chestnut- 
tailed and dark-tailed type occur rarely. Red-tailed specimens very rare, 
only 2-3% of the population. Summer fur of fully grown specimens dark, 
intense chestnut -reddish, darker than that ofS. v. ognevi. 

MEASUREMENTS. Length of skull 49-52.3 mm (M. 51.2 mm); condylo- 
basal length 43-48.7 mm (M. 46.3 mm); zygomatic width 28.3-32.2 mm 
(M. 31.2mm); length of upper tooth row 9-10 mm (M. 9.2 mm); hind feet 
54-61mm (M. 57.6mm); tail 140-185mm (M. 167,3mm). 

DESCRIPTION. Back pure gray (neutral-gray) with a light buff tinge 
(mouse-gray) and dense dark gray mottling. Series of this subspecies 
differ sharply from series of Moscow squirrels in their purer gray color. 
Bald-spine specimens are rarer than among S. v. ognevi. Forehead 
sometimes with a buff- gray tinge (drab). Specimens without buff tones on the 
head are common, their color is gray with darker mottling. Lower parts 
of cheeks and chin light gray or buff-gray, gradually passing into white in 
the middle of the throat. Color of flanks like that of back. Ears black, 
sometimes with a chestnut tinge at the base. Tail of the chestnut -tailed 
type. Dark-tailed specimens also occur. Gray roots of hair on tatil well 
developed in some dark-tailed specimens from Arkhangelsk. Rust-brown 
hair on soles of winter fur sometimes retained. 

There are rarely specimens with a color resembling that of the central 
Siberian squirrel, i,e, , with a reduced white area on the belly. To this type 
belongs specimen No. M,4242, male, of my collection (caught by 



337 



390 



V. Parovshchikov on 29 September 1934 in the vicinity of village Tundra in 
the Kholmogory District, former Arkhangelsk Province). The back of this 
specimen is gray with dark mottling and a well marked bald- spine. Flanks 
of a purer gray slightly different from that of back. White area of belly 
reduced. Throat with a white wedge, which is surrounded by a dark gray 
fringe. Ears black. Tail of the chestnut -tailed type, with well developed 
black hair tips. V. Parovshchikov informed me that he found a specimen 
of similar color among many hundreds of squirrels from Arkhangelsk. 

Summer fur of old specimens darker than that of old Moscow S . v. 
ognevi. Back between chestnut -brown and kaiser -brown, i.e., intense 
chestnut-red. Nose and cheeks with buff-gray-rust tones. Flanks in many 
specimens saturated red-ocher-chestnut (between orange-rufous and tawny). 
Feet of same color as flanks. A reddish orange tone in lower part of neck 
and on chin, surrounding a white wedge in the middle of the throat. Tail 
of the chestnut or dark chestnut type. Young much lighter and yellower. 

SYSTEMATIC NOTES. Differs from S , v. ognevi as follows: l) purer 
gray color of winter fur; 2) fur denser and more luxuriant; 3) lower 
percentage of bald-spine specimens*; 4) much lower percentage of red- 
tailed specimens (2-3 % as against 25-30 % in S . v . ognevi). 

GEOGRAPHICAL DISTRIBUTION. S, v. formosovi occurs in 
northeastern Europe. The southern boundary of the range apparently 
follows the line Perm - Kostroma - Novgorod. It is also distributed in the 
basins of the Pechora and Northern Dvina. Squirrels of the Kholmogory 
District belong to this form, and not to the closely related form S . v . 
varius , but they are perhaps transitional to this. In Finland and the 
Kola Peninsula S. v. varius is replaced byS. v. formosovi. The 
squirrels at the coast of the Gulf of Finland are of an intermediate type, 
more closely related to V ar i u s than to o gn e v i . Along its southern 
boundary the range ofS. v. formosovi passes gradually into that of 
S.v. ognevi. 

362. Sciurus vulgaris varius Barrett -Hamilton (1899). ex Kerr 
(1792). Kola squirrel 

1762. Sciurus varius Brisson A. D. , Regnum Animale in classes IX, Lugduni Batavorum, p. 106. 

1789. Sciurus vulgaris varius Gmelin in Systema Naturae Linnaei I, ed.XIII, p. 146. 

1792. Sciurus vulgaris v arius Kerr K. , Animal Kingdom, p. 255. 

1868. Sciurus vulgaris Kessler K. , Materialy dlya poznaniya Onezhskogo ozera i Obonezhskogo kraya 

(Materials on the Onega Lake and the Onega Territory), Sankt Petersburg, pp. 21-22. 
1884. Sciurus vulgaris Pleske Th. , Uebersicht der Saugethiere und Vogel der Kola-Halbinsel, S. 105-108. 
1899. Sciurus vulgaris varius Barrett-Hamilton G. , On European Squirrels, Proceedings of the 

Zoological Society of London, p. 6. 
1910. Sciurus vulgaris varius Trouessart E. L, Faune des Mammiferes d'Europe, pp. 117-118; Miller 

G. S. Catalogue of the Mammals of Western Europe. London, 1912, pp. 906-907. 
1928. Sciurus vulgaris v ari us SerebrennikovM., Synopsis of Russian Squirrels, Doklady Akaemii Nauk 

SSSR, p. 421. 

TYPE LOCALITY AND DEPOSITION. Brisson, who described S c i u r u s v a r i u s records its distribution as 
follows "Habitat Europa". According to Gmelin and Kerr, its range is northern Europe. The first exact 
indication of the type locality was given by Barrett-Hamilton, i. e. , northern Scandinavia and Lapland. 
Holotype unknown. 

MATERIAL EXAMINED: 20 specimens. 



The rust tone of bald-spine S. v. formosovi (on spine and lower part of back) is superficial, not deeply 
colored hair. 



338 



91 



DIAGNOSIS. Winter specimens are mostly light colored and bald- spine. 
Specimens of a more intense gray color are rare. Bald- spine squirrels 
occur frequently. Tail usually of the chestnut-tailed type. Red-tailed 
specimens constitute 30% of the population. Dark-tailed specimens are 
rare. Winter fur more luxuriant and silky than in Moscow squirrels. 
Sunamer fur of adult specimens saturated, red-chestnut-rust colored. 

Length of skull 49-53.6 mm; condylobasal length 44-48.6 mm; zygomatic 
width 30.2-31.6 mm; length of upper tooth row 9-9.8 mm. 

DESCRIPTION. Winter fur. Back very light bluish gray in most Kola 
squirrels (between gull-gray and pale neutral-gray, Ridgway, 1912), with 
fine mottling. Nose and cheeks with a light gray -buff- rust tinge. Flanks 
lighter than back, of a pure gray-blue (gull- gray), sharply contrasting with 
the white belly. The color of the ears in such specimens varies between 
rust-red (tawny, mikado-brown) and black-chestnut or even black. Feet 
more or less rust colored. Tail rust colored as in chestnut -tailed 
specimens. Red-tailed specimens occur frequently, constituting 30% of 
the squirrel population of the Kola Peninsula. There are also many 
transitions between chestnut -tailed and red-tailed squirrels. The basal 
third of the tail in such specimens is of a rich reddish rust color (orange - 
cinnamon). Roots of hairs in this area bluish gray, followed by narrow 
blackish bands and broad reddish rust bands. Some hairs with small black 
tips. In the middle of the tail, particularly near the tip, the black hair 
tips become wider, so that the tail appears blackish. Tips of terminal 
hairs rust colored. Medium line of ventral side and base pale bluish gray. 
Sides of ventral area rust-red followed by a broad black-chestnut fringe. 
Fur luxuriant, soft and silky, similar to that ofS.v.formosovi in full 
pelage. 

Dark gray, dark-tailed squirrels are rarer on the Kola Peninsula. Back 
ash-gray (between light neutral-gray and neutral-gray) in such specimens, 
with well marked dark gray mottling. Head dark ash-gray with fine dark 
mottling, and rust-buff tones on tip of nose and lips. Gray color lighter on 
flanks, giving a bluish shade. Feet, especially soles, with gray hair. Tail 
gray-black, without any rust shades. Hair roots of tail dark ash-gray; tips 
of hair of tail black. 

Summer fur red-chestnut-rust colored (between kaiser-brown. Hay's 
russet with chestnut-brown shades). General color very similar to that of 
summer fur ofS. v. formosovi. Forehead and cheeks with buff -rust 
shades. Rust -red tones on lower parts of cheeks and chin sharply set off 
by white wedge in middle of throat. 

Flanks and feet rich rust-red color. Tail of the chestnut and dark- 
chestnut type. Red-tailed specimens are common (see above). 

SYSTEMATIC NOTES. Differs from S . v. ognevi as follows: l) 
usually purer light bluish gray winter fur; 2) denser, more silky and 
luxuriant fur; 3) bald- spine specimens rare as compared with the Moscow 
squirrel; 4) darker color of summer fur. 

Differs from S. v. formosovi as follows: l) lighter bluish gray color 
of winter fur; 2) large percentage of red-tailed specimens (30% as 
against 2-3 % in S . v . f o r mo s o vi ); 3) large number of red-eared 
specimens which are very rare in winter specimens ofS. v. formosovi. 

I compared Swedish specimens ofS. v. varius (from near Upsala) in 
the ZMAN collection with S. v. vulgaris and found that its summer fur 
is very dark, the tail chestnut colored. The back of Swedish specimens is 



339 



dark chestnut, warm-sepia (pi. XXIX, Ridgway, 1912). The tail is still 
darker and the flanks more rust colored (with mikado-brown tinge, pi. XXIX). 

The chestnut color is retained to a large degree in winter fur. A light 
silver-gray tinge at the hair tips. Skull small, length about 52mm (cf. 
G.S. Miller's description in: Catalogue of the Mammals of Western Europe, 
1912, p. 905). 

GEOGRAPHICAL DISTRIBUTION. Kola Peninsula (Lapland and Finland). 
Squirrels from the Onega Territory apparently belong to this subspecies. 

363, Sciurus vulgaris ukrainicus Migulin. Ukrainian rust- 
colored squirrel 

1855. Sciurus vulgaris Severtsov N. A. , Periodicheskie yavleniya v zhizni zverei, ptits i gad 

Voronezhskoi gubemii (Periodic Phenomena in the Life of Mammals, Birds and Reptiles of the Voronezh 

Province), p. 379i 
1894. Sciurus vulgaris Silant'ev A. A. , Fauna gadov (The Reptilian Fauna), Sankt Petersburg, p. 35; by 

the same author: Zoologicheskie issledovaniya (Zoological Investigations), Trudy ekspeditsii Lesnogo 

departamenta, IV (2) : 11- 12. 1898. 
1915. Sciurus vulgaris Gorbachev S. N. , Mlekopitayushchie Orlovskoi gubemii (Mammals of the Orel 

Province) [according to recent data], p. 6. 

1923. Sciurus vulgaris fuscoater Ognev S. I. , Fauna nazemnykh pozvonochnykh Voronezhskoi gubemii 
(The Fauna of Land Vertebrates of the Voronezh Province), pp. 153-154. 

1924, Sciurus vulgaris fuscoater Migulin A. A. , Mlekopitayushchie Khar'kovskoi gubemii (Mammals 
of the Kharkov Province), "Priroda i okhota na Ukraine", No. 1-2, pp. 119-128. 

392 1927-1928. Sciurus vulgaris ukrainicus Migulin A. A. , Obzor gryzunov Ukrainy (A Survey of the 

Ukrainian Rodents), "Zakhist roslyn", No. 3-4, 1927-1928, pp. 11-12. 
1928. Sciurus vulgaris fuscoater Gavrilenko N. , Opyt sistematicheskogo kataloga zverei 

Poltavshchiny (A Systematic Catalogue of the Mammals of the Poltava District), p. 10. 
1928. Sciurus vulgaris fuscoater Serebrennikov M. , Synopsis of Russian Squirrels, Doklady 

Akademii Nauk, p. 425. 

TYPE LOCAUTY AND DEPOSITION. No. 621, female, May 1915, Sumy County of the Kharkov 
Province, Kiyanitsa Townlet. Type in the collection of the Kharkov Station for Plant Protection. 

MATERIAL EXAMINED; 9 specimens. 



DIAGNOSIS. Winter fur retaining marked brownish rust tones along the 
spine and in lower part of back. Tail brown-rust colored. Ears lighter, 
also brown-rust. Fur coarse, not luxuriant. Color of summer fur on back 
and tail rich ocher-red-rust. 

MEASUREMENTS: Large; length of skull 53-53.4 mm; condylobasal length 
48.2-50 mm; zygomatic width 29.2-32.6 mm; length of upper tooth row 
10.1mm; hind feet 59-61 mm; tail 160-200 mm. 

DESCRIPTION. Winter fur (Plate VII) distinguished by a different 
intensity of gray. In the grayest specimens, back (except for the spine) 
dirty ash-gray (between mouse-gray and light neutral-gray) with fine 
chestnut-gray mottling. Brown-rust tones similar to cinnamon-pecan- 
brown or rufous-tawny well marked along spine, and in lower part of back. 
Spine also with well marked fine brownish rust mottling. Cheeks, forehead 
and occiput light gray- rust with dark gray mottling. Ears light brownish 
rust (cinnamon-rufous to pecan-brown). Brown-rust color also on feet. 
Dorsal side of tail brown-rust (between cinnamon-rufous -tawny and hazel). 
Medium line of ventral side with ash-gray hair roots. 



340 



PLATE VII 




Ukrainian rust-colored squirrel (Sciurus vulgaris ukrainicus Migulin). Winter fur 

Drawing by V. A. Vatagin 



341 



Specimens with rust -brown tones occur together with gray specimens. 
Rust-brown tones well marked on back, especially along spine and on flanks. 
Gray color of flanks with a rust-buff tinge. 

Winter fur differs from summer fur in well marked gray color of flanks. 
Ears with long hair tufts of pale brownish rust tone (cinnamon-rufous, 
pecan-brown). 

Color of summer fur on back, legs and tail, saturated ocher-red-rust, 
between cinnamon -rufous, hazel and kaiser -brown. Color of flanks 
richer, lighter and more yellow. Nose and cheeks, sides of neck and chin, 
with grayish tones. Chest with white wedge, apex pointing anteriorly. 
393 SYSTEMATIC NOTES. Differs from S. v. ognevi by its large size 
and brownish rust color of winter fur. 

GEOGRAPHICAL DISTRIBUTION. Bryansk, Trubchevsk and Dmitrievsk 
forests, in the Poltava District, from Romny in the south to Krasnograd 
(apparently only rarely). It is also found in the former Izyum, Sumy and 
Zmiev counties of Kharkov Province. In the east this animal occurs in 
isolated localities, e.g., the Grafskaya forestry (north of Voronezh), the 
Khrenovsk forest. It was rare in the Shipov forest (former Pavlovsk 
County), and has now disappeared from there. According to D. Kapranov 
("Hunter's Gazette" (Okhotonich'ya gazeta). No. 13, 1928, p. 5) several 
families were found in ravines in Ternovaya and Dashkina of the former 
Boguchar County of the Voronezh Province. This is the most south- 
eastern record ofS. v. ukrainicus. 

364. Sciurus vulgaris kessleri Migulin (1928). Kessler's squirrel 

1840. Sciurus vulgaris Nordmaim A. , Observations sur la Faune Pontique; in: A. Demidoff, Voyage 

dans la Russie meridionale, III, pp. 54-55. 
1850. Sciurus vulgaris Kessler K. F. , Zhivotaiye gubemii Kievskogo uchebnogo okmga (Animals of 

the Provinces of the Kiev District), pp. 38-40. 
1858. Sciurus vulgaris Belke G. , Esquisse de I'Histoire Naturelle de Kamienietz-Podolski. Bulletin 

Naturel de Moscou, p. 26. 
1877. Sciurus vulgaris Tacznowski L. , Liste des Vert^bres de Pologne, Bulletin Zoologique de France, 

p. 129. 
1915. Sciurus v u 1 g a r i s Sharleman E. V. , Mlekopitayushchie okrestnostei Kieva (Mammals of the 

Vicinity of Kiev), Izdatel'stvo Kievskogo omitologicheskogo obshchestva, pp. 50-51. 
1925-1926. Sciurus vulgaris f usco ate r KhranevichV,, Narys faunyPodillya(ASurveyof the Podolian 

Fauna), pp. 21-22. 
1927-1928. Sciurus vulgaris kessleri Migiilin A. A. , Obzor gryzunov Ukrainy (A Survey of 

Ukrainian Rodents), "Zakhist roslyn", Nos. 3-4, p. 12. 
1930.-S (ciurus) v (ulgaris) kessleri and S . v. ukrainicus Danilovich A. P. , O belkakh 

okrestnosteiKieva (Squirrels of the Vicinity of Kiev), "Okhotnik", March, Moscow, p. 20. 

TYPE LOCALITY AND DEPOSITION. Vicinity of Zhitomir. Holotype in A. A. Migulin's collection. 

MATERIAL EXAMINED: 40 specimens. 

DIAGNOSIS. Winter fur of light specimens (spine and lower part of 
back) with rust-brown tones. Dark specimens with dark brownish rust tones. 
Tail rust-brown. Ears deep gray-chestnut with a light reddish tinge. The 
subspecies differs, in this respect, from S . v. ukrainicus, the ears of 
which are light (see above). 

Fur coarse, not luxuriant. Summer fur on back and tail of two types: 
a) rich ocher-red-rust; b) saturated red-chestnut 

MEASUREMENTS larger than in S. v. ukrainicus: Length of skull 51 - 
55.2 mm (M. 53.7 mm); condylobasal length 48-52.8 mm (M. 49.5 mm); 



342 



395 



zygomatic width 31-34,3 mm (M. 32.5 mm); length of upper tooth row 9.4- 
10.2 mm (M. 9.8 mm); hind feet 53-65 mm {M.58mm); tail 180-205 mm 
(M. 188mm). 

DESCRIPTION. Winter fur of two types: l) light type. These 
specimens closely resemble the rust-brown type ofS. v. ukrainicus 
Migul. (see above). S.v.kessleri, however, differs from S . v . 
ukrainicus in the darker color of the ear s . The ears ofS. v. kessleri 
are dark chestnut-gray with a light reddish tinge (natal-brown or benzo- 
brown, occasionally near fuscous) in darker specimens. 

2) Dark type. Top of head, lower part of back and spine dark rust-brown 
(between russet andProut's brown). Neck, shoulders and flanks with a 
marked gray tinge and brownish-chestnut mottling. Chin and sides of neck 
gray-buff (light-drab), differing from rust color of cheeks. Ear tufts 
fuscous, i.e. , deep gray-chestnut with a light red tinge. 

Feet rich rust-brown (of a purer shade than russet). Tail rust-brown 
(between russet andProut's brown). Hair on tail with ash-gray-chestnut roots. 
Ventral side of tail like dorsal side. However, middle of ventral side with 
more marked gray shades. 

Summer fur. In specimens of the light type, color like that of S . v . 
ukrainicus except for the ears, which are chestnut-gray. Top of head and 

backof dark squirrels saturated red -chest nut, be- 
tween bay and chestnut. Flanks and legs lighter and 
redder, between ferruginous and kaiser-brown. 
Chin and sides of neck lighter, buff, clearly 
different from dull reddish chestnut-gray color 
of cheeks. Color of tail resembling that of back, 
a little redder and more rust colored. Ash- 
chestnut- gray hair roots well marked in median line 
of ventral side. 

SYSTEMATIC NOTES. S . v . ukr a in i c u s 
and S. V. kessleri differ in series in a number 
of'characters (see diagnosis). 

The most constant character is the color of the 
ears . Those ofS. v. kessleri are usually 
much darker than those ofS. v. ukrainicus. 
A. P. Danilovich (1930) recorded both S . v . 
kessleri (5 specimens) and S. v. ukrainicus 
(2 specimens) near Kiev. This author probably 
considered light specimens ofS. v. kessleri as 
ukrainicus. I examined a large series of S . 
V. kessleri and did not find a single 
ukrainicus among them. All specimens from 
the left-bank Ukraine proved to belong to typical 
ukrainicus . 
Series of S . V . f us c o at e r Altum (type from Harz Mts. , Germany) 
should be compared with S. v. kessleri to determine their relationship. 
G. S, Miller (1912) gives a good description ofS, v. fuscoater which 
shows that the winter fur ofS, v. fuscoater differs from that of S . v . 
kessleri as follows: l) light type — purer and richer reddish tone 
(rufous) with light buff-gray shades on the sides of the back (pale smoke- 
gray); 2) dark type — difference even more marked. Back of S , v . 
fuscoater chestnut, without any reddish brown tinge, with a gray tinge 




FIGURE 185. Skull of 
Sciurus vulgaris 
kessleri Migulin, No. M. 
3704, from the former Kiev 
Province 



343 



396 



and a black tail. I never found black-tailed specinaens ofS. v. kessleri. 
S. V. kessleri seems therefore to be a well characterized form of S . v . 
fuscoater . 

GEOGRAPHICAL DISTRIBUTION. Forests of the right bank of the 
Dnieper. It is found in the north in the Minsk and Gomel districts. It 
occurs throughout Podolia and the northern, forested part of Bessarabia. 
The southern boundary of the range is not known. It is found in isolated 
forests from Balta and Uman' up to Zvenigorodka and Cherkassy. 

365. Sciurus vulgaris bashkiricus Ogn, (1935), Bashkirian 
sqtiirrel 

1928. Sciurus vulgaris exalbidus (partim.') Serebrennikov M. , Synopsis of Russian Squirrels. 

Doklady Akademii Nauk SSSR, p. 424; Serebrennikov M. K. , Materialy po sistematike i ekologii 

gryzunov yuzhnogo Zaural'ya (Materials on the Systematics and Ecology of the Rodents of East of the 

Southern Urals), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, 1929, pp. 280-281. 
1935. Sciurus vulgaris bashkiricus Ognev S. I. Sistematicheskii obzor belok, vodyashchikhsya v 

SSSR (A Systematic Survey of Squirrels of the U. S. S. R. ), Byulleten' Nauchno-issledovatel'skogo 

instituta zoologii Moskovskogo universiteta, No. 2, p. 46. 

TYPE LOCAUTY AND DEPOSITION. No. M. 3352, female, 25 November 1929, Buzuluk pine forest, 
Shkotovskoe forestry (former Buzuluk County of the Samara Province). Holotype inS. I. Ognev's collection. 

MATERIAL EXAMINED: holotype and 14 paratypes. 

DIAGNOSIS. Winter fur light, between pale whitish buff-gray to bluish 
gray. Winter fur always with gray mottling. Many specimens bald-spine, 
with buff-ocher tones along spine and on lower part of back. Ears between 
ocher-red and blackish or black-chestnut. Chestnut-tailed specimens 
common. Dark-tailed specimens rarer. Fur soft but not luxuriant. 
Summer fur without the intense color of northern or Moscow squirrels. 
Summer fur ocher-red-chestnut. 

Skull larger than that ofS. v. ognevi: Length of skull 51-54.7 mm 
(M. 53.2mm); condylobasal length 47,3-48.7 mm (M. 48.2 mm); zygomatic 
width 31. 2-34. 2mm (M. 33.2mm); length of upper tooth row 9. 2-10, 2mm 
(M. 9.7 mm); hind feet 51 .3-57 mm (M. 52.8 mm); tail 164-208 mm 
(M. 185.6mm). 

DESCRIPTION. Winter fur uniformly light gray. Back of light 
specimens pale whitish gray with buff shade and gray mottling. Back 
between light gull-gray and pale smoke -gray. Lower part of back and 
occasionally spine with pale buff-ocher tones (bald-spine). In more 
intensely colored specimens, back more bluish-gray (between gull-gray 
and smoke -gray) with gray mottling. 

In light-colored specimens, sides of neck white. White area of belly 
unusually large, high on the flanks, to the shoulders, haunches and tibiae; 
top of head and occiput, forehead and cheeks with red-ocher tones. Nose 
buff-white with a weak ocher tinge. Ears red-ocher (orange-cinnamon). 
Tips of ears with black-chestnut tufts. Chestnut-tailed specimens common. 
Roots of hair on tail broad, pale whitish gray (pale smoke -gray), followed 
by pale straw-ocher subapical bands and black tips. Middle of ventral side 
of tail whitish. Ocher subapical bands of hair well marked, rich ocher-red 
(between orange -cinnamon and sayal-brown), the tips of hair black or black- 
chestnut. The tip of black or black- chestnut hair is yellowish buff, so that 
it appears mottled. 

In light specimens, feet in winter fur rich, light reddish ocher. 



344 



Dark-colored squirrels (see above). Cheeks, forehead and occiput with 
a light yellowish buff tinge. Base of ears dull rust-buff, long hair on ears 
black or black- chestnut. Soles, forelegs and antebrachia in part with a dull 
rust -buff tinge. 

Tail of the dark-tailed type. Hair of dorsal side with broad ash-gray 
roots (between gull-gray and deep gull-gray); hair tips black, so that the 
dorsal side appears dark. Middle of ventral side ash-gray, sides with a 
gray-buff tone in front of the zone with black hair tips. 

Summer fur. Back ocher-red-chestnut (between tawny and cinnamon- 
rufous), generally purer and more reddish than inS.v.ognevi and 
S. V. varius. 

Flanks and cheeks with reddish buff tones. Feet reddish ocher. Most 
squirrels examined in summer belonged to the chestnut-tailed type. Red- 
tailed specimens were rare. 

SYSTEMATIC NOTES. The Bashkirian squirrel differs from the Moscow 
squirrel in its paler and lighter color. These differences are especially 
marked in winter specimens. S.v.bashkiricus is closely related to 
the Ural mountain squirrel (see below), but differs from it in the less pure 
gray color of the winter fur, the buff tinge on the back and ocher tinge on 
occiput. S.v.bashkiricus resembles the Teleut squirrel. M.K. 
Serebrennikov (1928, 1929) considers the Bashkirian squirrel as identical 
with the Teleut squirrel. However, S.v.bashkiricus is much smaller 
and more short-tailed than the Teleut squirrel. Furthermore, Bashkirian 
squirrels never have gray tails, which are typical for the Teleut squirrel. 

GEOGRAPHICAL DISTRIBUTION. Buzuluk pine forest and forests of 
Bashkiria, to the middle reaches of the Belaya river in the north. This 
region is inhabited by a type intermediate between S. v. bashkiricus 
and S. V. ognevi. 

365'. Sciurus vulgaris bashkiricus natio ur ale ns is Ogn. (1935). 
Ural mountadn squirrel 

1874, Sciurus vulgaris Sabaneev L. P. , Pozvonochnye srednego Urala (Vertebrates of the Central Urals), 

p. 9. 
1897, Sciurus vulgaris Zarudnyi N, , Zametki po faune mlekopitayushchikh Orenburgskogo kraya 

(Notes on the Mammal Fauna of the Orenburg Territory), p. 2. 
1928. Sciurus vulgaris varius Dulkel 'skaya N. M. , Opyt obzora fauny mlekopitayushchikh 11' menskogo 

zapovednika (A Survey of the Mammalian Fauna of the Il'menskii Reserve), pp. 26-28. 
1935, Sciurus vulgaris bashkiricus natio uralensis Ognev S.I., Sistematicheskii obzor belok, 

vodyashchikhsya v SSSR (A Systematic Survey of the Squirrels of the U. S. S. R, ), Byulleten' Nauchno- 

Issledovatel'skogo instituta zoologii Moskovskogo urdversiteta, No. 2, p. 46. 

TYPE LOCALITY AND DEPOSITION. No. M. 1650, sex unknown, December 1926, Miass of the 
Zlatoust District (former Coimty of the Orenburg Province), Il'menskie Mountains, S. I. Snigirevskii. 

MATERIAL EXAMINED: holotype and 13 paratypes, 

DIAGNOSIS. Closely related to S. v. bashkiricus, but differing from 
it in the purer gray color of the back. In winter fur, back bluish smoke - 
gray (deep gull- gray) with a light buff- gray tinge (smoke -gray). Color of 
head and occiput like that of back. Base of ears smoke -gray. Tufts on 
ears black or black- chestnut. Tail of the chestnut or dark-chestnut type. 

Summer fur resembling that ofS. v. bashkiricus, but more chestnut- 
yellow, without the rust-red tones characteristic ofS. v. bashkiricus. 

Measurements like those of the Bashkirian squirrel. 

SYSTEMATIC NOTES. Closely related to S. v. bashkiricus, 
transitional to West Siberian squirrels. The winter fur differs from that 

345 



ofS.v.martensi in being less luxuriant and in having a more intense 
gray color of the back. S.v.martensi is larger than the Ural squirrel 
(see below). 

GEOGRAPHICAL DISTRIBUTION. Central Urals, in the northern parts 
and on its eastern slopes. I have specimens from the Nyays' river (eastern 
slopes of Northern Urals). Their summer fur resembles that of the central 
Ural squirrels. 

366, Sciurus vulgaris martensi Matschie (1901). West Siberian 
squirrel 

71792. Sciurus vulgaris arge nteus Kerr R. , Animal Kingdom, p. 256 (terra typica not indicated). 
1804. Sciurus vulgaris albus Dvigubski J. , Prodromus Faunae Rossicae, p. 86 (partim.') ("In 

deserto Barabensi ad fl. Samaram Obum, ad Baikalum. . . "). 
398 1904. Sciurus martensi Matschie P. , Zoographische Betrachtungen iiber die Saugetiere der 

nordlichen Alten Welt. Arch, f, Naturgesch. , Beiheft zu Bd. 67, S. 313. 
1915. Sciurus vulgaris Chugunov S. M., Mlekopitayushchie i ptitsy Surgutskogo uezda (Mammals and 

Birds of the SurgutCounty), Ezhegodnik Tobol'skogo gubemskogo muzeya, XXIV, pp. 32-33. 
1928. Sciurus vulgaris nadymensis Serebrennikov M. , Synopsis of Russian Squirrels, Akademiya 

NaukSSSR, p. 422. 

TYPE LOCALITY. Sciurus m arte ns i was described by P. Matschie from the left bank of the lower 
Yenisei. 

This so-called "species", which is in fact a subspecies ofS. vulgaris, was described on the basis of 
A. Middendorf 's work "Sibirische Reise", Saugethiere, VJJgel und Amphibien, St. Petersb. , 1867, S. 80. 
Middendorf gives a description of the light form, indicating that it occurs along the left bank of the 
Yenisei and along its right bank between the Stony Tunguska and the Polar Circle. 

Middendorf writes: "Die gewohnliche Farbung der Eichhomchen am unteren Jenisej, zwischen der 
Stein-Tunguska und Polarkreise, zumal auf dem linken Ufer des Stromes, ist ein helles Crau, welches um 
eine, nur bei dem geuauesten Vergleiche in die Augen fallende, Tinte heller ist als das Grau der baltischen 
Eichhomchen. Bei nUherer Untersuchung findet sich dann, dass das Grau der Jenisej Eichhomchen deswegen 
heller erscheint, weil das Weiss, welches jederseits die schwarze Binde jedes Deck-haares begrenzt, ein reines 
Weiss ist, und keine Spur einer gelblichen Tinte verrSth. " 

The author stressed that this squirrel is chestnut-tailed and that its ear tufts are chestnut-black. The 
above quotation shows that it is based on light specimens which inhabit the left bank of the Yenisei. On 
p. 81 Middendorf says that squirrels from the right bank of the Yenisei are darker: "Auf dem rechten Ufer 
des Jenisej sind, wie oben angefUhrt, die Balge dunkler. . . " 

M, K. Serebrennikov 's assumption that the description of m arte nsi applies to dark specimens from the 
right bank of the Yenisei is therefore erroneous (see below) 

Middendorf's type of martensi is apparently lost. 

MATERIAL EXAMINED: 1 19 specimens. 

DIAGNOSIS. Winter fur on back very light buff -gray with light grayish 
mottling. Specimens of a more intense gray color are rare. Bald -spine 
specimens are also rare. If a bald-spine is present, it is little marked 
on the lower part of back near the tail. Tail of chestnut type. Dark- 
tailed specimens are rare, red-tailed specimens very rare (see below). 
Winter fur luxuriant and silky. Summer fur of most specimens light, 
ocher-yellow-red, sometimes buff-yellow-gray, rarely darker, red-brown- 
ocher. Fur usually tricolored because of the presence of the light buff- 
ocher color of the flanks. White area of belly large: inner sides of hind 
legs and antebrachia, sides of neck and groin white. 

Length of skull 51-55.8 mm (M. 52.4 mm); condylobasal length 46-51.2 mm 
(M. 47.2 mm); zygomatic width 29.5-32.8 mm (M. 30.4 mm); length of upper 
tooth row 9-lOmm (M. 9.3 mm); hind feet 52. 1-6 1,1 mm (M. 57.9 mm); tail 
145-192mm (M. 166.1mm). 



346 



399 



DESCRIPTION. Color of winter fur more or less constant. Back 
very light buff-gray (light mouse-gray) with dense but weak gray mottling. 
Some deviations from this typical color type of the Ob squirrels occur, 
such as more intense gray tones (towards light neutral-gray and neutral- 
gray). Color of flanks even lighter, pale gray with a light buff tinge. In 
full winter fur, top of head light buff-gray, cheeks pale buff- gray and 
ears black. Tail generally of chestnut type. The hairs have ash-gray 
roots, rust-brown subapical bands and broad black or black- chestnut tips. 
Some hairs on the tail, especially near its end, have brownish rust tips. 
The rust tones on the tail disappear in dark-tailed specimens, and only 
ash-gray roots and black-chestnut tips remain. 

Color of summer fur very variable, but most specimens have a rather 
light color. Back of adult and old squirrels generally light ocher-yellow- 
red (tawny with a light reddish and yellow tinge). Nose and anterior part 
of head buff -gray- white. Cheeks also with a light buff tinge. Reddish 
ocher tones predominating on occiput. The subspecies is characterized 
by the very light, well marked buff-ocher (ochraceous-buff) 
color of the flanks sharply contrasting with the white underparts. The 
animal is therefore tricolored. Legs rich ocher-red-yellow. Tail 
generally chestnut (see above). 

The light-colored type includes specimens with buff- ye How -gray back 
with a light red tinge in summer (ochraceous -tawny with a tawny shade). 
Anterior part of head and nose with many whitish hairs. Feet ocher- 
yellow. Flanks with a stripe of pale yellow-ocher fur (light ochraceous- 
buff) causing it to appear tricolored. 

The dark-colored type includes a smaller number of intense chestnut- 
ochraceous colored squirrels, the backs of which are between kaiser- 
brown and hazel. Color of head more intense than in preceding forms, 
without a buff tinge. Flanks with well marked yellowish ocher fringes. 

Color of young generally more yellow and lighter than of adults. Back 
of very young squirrels often dull buff-gray with a light ocher tinge. 

SYSTEMATIC NOTES. Kerr (1792) mentioned S c iu r u s vulgaris 
argenteus ("silvery common squirrel") with the following description: 
"is larger than the ordinary varietas and of a glossy silver-gray color". 
He did not indicate the type locality. It may be assumed that his material 
consisted of West Siberian squirrels, the winter fur of which has these 
characters. I. Dwigubski (1804) mentioned S c i u r u s vulgaris albus, 
which is in part a synonym ofmartensi, as the Ob is mentioned as one 
of its habitats. 

The confused synonymy ofS.v. martensi with S.v. nadymensis 
is discussed above. 

The West Siberian squirrel is closely related to S. v. bashkiricus 
natio uralensis, and differs from it as follows: l) lighter buff tone of 
back in winter specimens; 2) more luxuriant fur; 3) large size. 
400 It was stated above that chestnut -tailed squirrels predominate among 
West Siberian squirrels. I.D. Kiris collected a large series of squirrels 
from Surgut. He estimated the percentage of color types and found the 
following: 

I. Dark chestnut-tailed 3.7% (4) 

II. Chestnut-tailed 93.5% (102) 

typical chestnut-tailed 62% (68) 

light chestnut-tailed 31.1% (34) 

III. Red-tailed 2.8% (3) 

347 



401 



The tails of some very young squirrels resemble those of chipmunks, as 
the hair is not fully grown. Such young specimens show a greater 
percentage of red -tailed specimens. The following figures were obtained: 

I. Chestnut-tailed 72.2% (13) 

typical chestnut-tailed 5.5% (1) 

light chestnut- tailed 66.7% (12) 

II. Red-tailed 27.8% (5) 

These data show that the age of the squirrel is an important factor in 
descriptions. 

S. V. martensi apparently forms a separate natio in the southern part of 
of its range, which is transitional to the Teleut squirrel. Squirrels 
between Novosibirsk and Narym have gray tails, but there are also 
chestnut-tailed specimens with a gray tone on the tail. 

These squirrels are intermediate between the Teleut squirrel and S . v . 
martensi in some respects. In the Tomsk-Novosibirsk region, and also 
north and southeast of this region, there are many light chestnut-tailed 
squirrels with marked rust tones on the tail and with red-rust-coiored ears. 
The winter fur of these squirrels is very light, pale buff-gray with weak 
mottling. I cannot determine the systematic position of these specimens at 
present, for lack of material. 

The furriers consider these squirrels as a separate Tomsk strain. 
According to B. A. Kuznetsov (in litt. , 1935), there are no black-tailed or 
dark-tailed specimens in this strain, which consists mainly of chestnut- 
tailed specimens, of both dark and light types. A few specimens have red 
tails. B. A. Kuznetsov examined 96,581 pelts and found the following: 

Dark-tailed 49% 

Chestnut-tailed 44% 

Red-tailed 7% 

The Tomsk strain is apparently an intermediate form (natio) between 
martensi andaltaicus (?). 

Further study will show whether these squirrels deserve a separate 
name. 

GEOGRAPHICAL DISTRIBUTION. A large part of West Siberia in the 
middle and lower reaches of the Ob and along the Narym. The subspecies 
reaches the Yenisei in the east. The light -colored squirrels gradually 
disappear on the right bank of this river, and are replaced by the darker 
subspecies . The southern limit of the range of S. v. martensi is not 
exactly known. It is also not known which form occurs in the Tara region, 
between Narym and Tomsk and near Novosibirsk. 

367. Sciurus vulgaris exalbidus Pallas (1778). Teleut 
squirrel 

1788. Sciurus vulgaris varietas e x a 1 bi da Pallas P. , Novae Species Quadrupedum e Glirium ordine, 

p. 374, 
1792. Teleutskaja belka or Squirrel of the Teleutian tartars. Kerr R. , Animal Kingdom, p. 256 

(Upper parts of the Ob). 
1811-1831. Sciurus varius (partim'. ) Pallas P. , Zoographia Rosso-Asiatica, pp. 183-185. 
1911. S (ciurus) Talahutky Brass E. , Aus dam Reiche der Pelze, S. 586 (South Siberia). 



348 



1923. Sciurus vulgaris Martino V. and E. , Materialy po sistematike i geograficheskomu 

rasprostraneniyu mlekopitayushchikh Kirgizskoi stepi (Materials on the Systematics and Geographical 
Distribution of Mammals of the Kirghiz Steppe), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, 
Vol. XXIV, pp. 21-22. 

1928. Sciurus vulgaris exalbidus Serebrennikov M. , Synopsis of Russian squirrels, Doklady 
Akademii Nauk. 
TYPE LOCALITY AND DEPOSITION. Described by Pallas from a specimen from the region of isolated 

pine forests along the Irtysh and Ob. Type lost. 
MATERIAL EXAMINED: 210 specimens. 

DIAGNOSIS. Winter fur light buff- gray with grayish mottling. The 
following variations occur: l) darker color of back. The dark specimens 
of the Teleut squirrel are, however, much lighter than dark S . v . 
martensi. 2) albinism. These whitish squirrels usually show 
yellowish buff bald-spine. Tail of most specimens pale gray with a 
blackish gray tinge and usually very pale yellowish rust shades. Chestnut- 
and dark-tailed specimens never occur. Hair of winter fur very long and 
dense. 




FIGURES 186, 187. Left, skull of S ciurus vulgaris exalbidus Pallas, 
No. 333 female, Korosten forestry. West Siberia, G. K. Gol'tsmaier. Right, 
skull of S ciurus vulgaris rupestris Thos. , No. M.3776, Sakhalin 

Summer fur usually light, between pale ocher-yellow and more saturated 
orange-ocher. White area of belly large. Inner sides of hind legs and 
antebrachia white. The white color reaches far along the sides of the neck. 
Groin white . 
402 Size very large: length of skull 54.6-59.2 mm (M. 56.9 mm); condylobasal 
length 50-53.2 mm (M,52mm); zygomatic width 33.1-36 mm (M. 34.9 mm); 
length of upper tooth row 10-11.1 mm (M. 10.4 mm); hind feet 56.1-65.5 mm 
(M. 62. 9mm); tail 153.2-222mm (M. 196.2mm). 

DESCRIPTION. Winter fur. Back light buff-gray with dense, well 
marked mottling which is not dark gray. Back pale mouse -gray. Color of 
flanks lighter, palid mouse -gray. Mottling practically absent on flanks. 
Hair of belly white, without gray roots. Outer parts of shoulders and 
haunches lighter than back (between pale mouse -gray and pallid mouse - 
gray). Forehead and occiput gray with a slight rust tinge. Anterior part 



349 



403 



of forehead, cheeks and region around the eyes pale yellowish buff- rust 
(light pinkish cinnamon) with a light cinnamon tinge which is especially 
marked in the middle of the muzzle. Tufts of ears luxuriant, ocher -red- 
rust (orange -cinnamon). Feet more rust colored and richer than the ears 
(pinkish cinnamon). Tail very long and luxuriant, gray, generally pale 
mouse -gray with a blackish gray tinge and yellowish buff hair tips. Color 
of hair on tail of typical specimens as follows: roots broadly whitish gray 
(pallid mouse -gray), followed by black bands which are well marked on the 
sides, and pale ye How -buff (pinkish buff) hair tips. Whitish roots of hair 
clearly marked in the middle of the ventral side of the tail. 

The above is a description of typical Teleut squirrels. The color of 
the winter fur is very constant, with only the following small variations. 

Dark-colored type. General color of back more intense, not so buff, 
more bluish (between pale mouse-gray and light mouse -gray with a pallid 
neutral-gray tinge) and with a fine, more yellow-gray mottling. Flanks 
lighter than back. Belly pure white. Color of head like that of preceding 
type, but the gray tones are generally more intense. Tufts of ears 
luxuriant, dark chestnut -brown (between mikado-brown and verona-brown). 
Soles rust -yellow (cinnamon). Outer parts of hind soles with broad black 
fringe. Tail gray with well marked black hair tips. Color of hair on tail 
as follows: roots broadly light gray (pale mouse -gray); hair tips long, 
black-chestnut or black. Terminal hair of tail with light gray roots, dull 
yellowish buff bands, blackish subapical bands and yellowish buff tips. 

The light-colored type shows a tendency towards albinism. Back whitish 
gray with a buff tinge, which is especially marked along the middle of the 
spine (pale drab-gray or even more whitish). . Muzzle, region around the 
eyes and forehead very light, yellowish buff (light pinkish cinnamon); 
flanks whitish, belly pure white, feet light, yellowish rust colored (between 
light pinkish cinnamon and pinkish cinnamon). Ears light ocher-rust 
colored (between pinkish cinnamon and vinaceous -cinnamon). Roots of hair 
on tail mainly whitish gray, followed by narrow pale dull yellowish buff bands, 
gray-black subapical bands and pale rust -buff tips. Terminal hair of 
tail without gray-black bands, only with broad dull rust-buff tips. This 
combination results in a pale gray-buff color. 

Summer fur. Back of full grown specimens between pale ocher-yellow- 
rust (pinkish cinnamon to cinnamon) and darker orange -ocher (orange - 
cinnamon), sometimes chestnut-ocher (mikado-brown). Head usually lighter 
and more yellow than back. Color of ears like that of back, sometimes 
darker. Flanks light yellowish with a rust tinge (cinnamon-buff to pinkish- 
cinnamon). Color of feet similar to that of back. Color of tail variable, 
generally whitish gray in lighter specimens, with yellowish buff and dark 
gray shades caused by bands of these colors. Tip of tail pale yellowish 
buff. In darker specimens tail more ash-gray with well marked black-gray 
hair tips. There are also squirrels with pale gray tails darkened on the 
upper part by blackish hair and having almost pure white tips. 

Color of young squirrels duller, lighter and yellower than that of adults. 
Color of back of young between cinnamon and pinkish cinnamon. Tail of 
young usually pale gray with a well marked yellowish rust tone, especially 
near the tip. 

SYSTEMATIC NOTES. This subspecies is well characterized. It is the 
largest subspecies. The tail is pale gray. The Teleut squirrel is a link in 
the chain of West Siberian squirrels. Its color generally closely resembles 



350 



that ofS. V. bashkiricus, but the two are not identical. It does not 
reach the Volga, as M.K. Serebrennikov (1928) thought. The Teleut 
squirrel is also closely related toS.v.martensi. It is easily 
distinguished from these subspecies by its large size and long tail. We 
stated above that a transitional type between martensi and exalbidus 
occurs between Novosibirsk, Tomsk and Narym. Specimens from 
Novosibirsk in my collection resemble Teleut squirrels, but their ear 
tufts are black. This was never seen in true S . v . e xalb id us . More 
material on the Tomsk squirrels should be collected, so as to be able to 
determine their relationship to squirrels in the Kalba Range, which 
were recently described by V. A. Selevin (1934) as a separate race. 

GEOGRAPHICAL DISTRIBUTION. The Teleut squirrel mainly occurs in 
pine forests along the right bank of the Ob, between Biisk and Novosibirsk. 
It is also found in narrow strips of pine forests in the Kulundin Steppe. 
Some of these pine forests, e.g., the Burlin forest, are north of the 
Kamen' and the Teleut - Gorkoe lakes. Farther southwest the Teleut 
squirrel occurs in "ribbon" pine forests along the Irtysh (near Semipalatinsk 
and east and west of this town). The ribbon pine forests along the Irtysh 
include the Loktevskii, Shul'binskii, Korostelevskii, Srostinskii and 
Severnyi pine forests. 

The northern boundary of the range of the Teleut squirrel is not known 
exactly. G.K. Gol'tsmaier (1935) assumes that the northwestern boundary 
follows the line Sverdlovsk - Chelyabinsk - Kurgan. In my opinion, the 
Teleut squirrel does not occur there, but is replaced by the Ural squirrel 
on the Urals as far as Sverdlovsk and apparently by the Bashkirian squirrel 
in the Chelyabinsk District. Farther east, near Novosibirsk, the range of 
the Teleut squirrel joins that of the West Siberian squirrel (S. v. martensi 
or a related form) which occurs north of the town, while the Teleut squirrel 
is found in adjacent pine forests, south of the town. From Novosibirsk 
the boundary turns southeast and follows the forest steppe which separates 
the pine forests along the Ob from the forests of the Salair Range. The 
boundary then crosses the Biya river above Biisk, alongthe line connectingthe 
village of Karabinskoe with the estuary of the Kazh river (a left tributary of the 
Biya). Accordingto S. N. Korchev, the Teleut squirrel occurs in smallnumbers 
further along the river Biya, along its tributary Lebed', in small pine 
forests along these rivers, in the lower reaches of the Baigol (a tributary 
of the Lebed') and its tributary, the Klyk. The occurrence of the Teleut 
squirrel along the Baigol and Klyk rivers is of special interest. It inhabits 
isolated pine forests in the Altai taiga among cedar and spruce forests, in 
which the Altai squirrel occurs; their ranges are adjacent and partly 
overlapping. 

Pallas was thus apparently wrong in stating that the Teleut squirrel 
occurs along the Chulym to the Yenisei. 

We have no exact data on the distribution of the Teleut squirrel in the 
pine forests west of the Irtysh, towards Pavlodar and Kokchetav. G. K. 
Gol'tsmaier (1935) writes: "At present, the Teleut squirrel does not occur 
farther east in the pine forest near Karkaralinsk, on the Kokche-Tau or 
along the upper reaches of the Tobol river. " 

N.M. Mikhel' ("Trudy Kazakhskoi bazy Akademii Nauk SSSR", 1934, 
pp. 35-51) does not record the Teleut squirrel from the Borovskii forest. 
A.M. Belyaev ("A List of the Mammals of Kazakhstan" (Spisok gryzunov 
Kazakhstana), Trudy Zoologicheskogo instituta Akademii Nauk, II, 1934, 



351 



p. 30) states that the Teleut squirrel is a "foreigner" in the pine forests of 
the former Kokchetav County. 




FIGURES 188, 189. Skull of S c i urus vulgaris exalbidus Pall., No. M.3906, female, 
25 Nov. 1931, Srostinskii pine forest, former Zmeinogorsk County of the Tomsk Province. 

Drawing by Yu. A. Kostylev 



352 



406 



In September 1933 N. A. Bobrinskii and S. A. Severtsov caught two 
Teleut squirrels* in an oak-birch forest, 50 km southwest of Kokchetav. 
It seems that the animal occurs rarely in this region. 

368. Sciurus vulgaris kalbinensis Selevin (1934). "Borovushka" 
or the Kalba squirrel 

1934. Sciurus vulgaris kalbinensis Selevin V. A. , Predvaritel 'noe opisanie novykh form gryzunov 

iz Kazakhstana (A Preliminary Description of New Forms of Rodents from Kazakhstan), Byulleten' 

Sredneaziatskogo gosudarstvennogo universiteta, 19(13) : 75. 

TYPE LOCAUTY AND DEPOSITION. Collected 30 Dec. 1931, Ayudin pine forest on the southern slope 
of the Kalba Altai. Type (No. 3, adult) in V. A. Selevin's collection. 

MATERIAL EXAMINED." I have, unfortunately, not seen any material. Selevin does not state the 
number of specimens examined. 

DIAGNOSIS (after V. A. Selevin): "Color similar to that of S c iu r us 
vulgaris exalbidus Pall., but slightly darker and grayer in winter 
fur. Distribution of color areas as in the Teleut squirrel. Winter fur of 
adult specimens often with traces of bald-spine, i.e., rust-colored spots 
on posterior part of back. This is a juvenile character in S. v. exalbidus. 
Color of tail differs sharply from that of the Teleut squirrel, it is rich 
rust, rarely chestnut, while the tail of the Teleut squirrel is always gray**. 
When the tail of the Kalba squirrel is chestnut colored, the whole animal 
is darker. S. v. kalbinensis, therefore, apparently has different color 
types, including light rust-tailed and dark chestnut -tailed forms. Its 
skull measurements are intermediate between those of the Altai and the 
Teleut squirrel. Condylobasal length 49.5-50 mm; zygomatic width 32.0- 
34.5 mm. " 

SYSTEMATIC NOTES. It is difficult to determine the differences between 
S. V. kalbinensis and its geographical neighbours from V.A. Selevin's 
short description. It certainly differs from the Teleut squirrel to judge 
from the description. 

The differences between the Kalba squirrel and the Teleut squirrel 
were mentioned before the publication of V.A. Selevin's description (B.A. 
Kuznetsov, "Rodents of the Semipalatinsk District" (Gryzuny 
Semipalatinskogo okruga), Byulleten' Moskovskogo obshchestva ispytatelei 
prirody, XLI (l-2):80-81. 1932). Kuznetsov writes: "The second form 
found in pine forests of the Kalba Range differs from the Altai 
specimens as follows: l) larger size; 2) back light silver-gray in winter 
fur and rich rust in summer fur; 3) tail of an intense red color. To judge 
from pelts, the Kalba squirrel is a separate, so far undescribed 
subspecies. " 

A comparison between S. v. kalbinensis and squirrels from the 
Tomsk Territory (see above) would be interesting. 

GEOGRAPHICAL DISTRIBUTION. Pine forests on the southern slopes 
of the Kalba Altai. 



* It is difficult to decide whether squirrels from forests near Kokchetav are identical with the true Teleut 
squirrel (the latter is probably larger). 
** This is not correct (see above). 



353 



369. Sciurus vulgaris jeni s s e jens i s Ogn. (1935). Yenisei 
squirrel 

1867. Sciurus vulgaris Middendorf A. Th. , Sibirische Reise, S'iugethiere, S. 78-83. 

1880. Sciurus vulgaris Tel (in NordenskjiSld's book), Ekspeditsiya k ust'yu Eniseya (An Expedition to 

the Estuary of the Yenisei), p. 178. 
1920. Sciurus vul garis Solov'ev D. K. , Sayanskii promyslovo-okhotnichii raion (The Sayan 

Commercial-Huiiting District), pp. 255-257. 

1924. Sciurus vul garis Kozhanchikov L. and I. , Promyslovaya okhota i rybolovstvo v Minusinskoi 
taige (Commercial Hunting and Fishing in the Minusinsk Taiga), Ezhegodnik Gosudarstvennogo muzeya 
imeni Mart'yanova, II (2) : 131-136. 

1925. Sciurus vulgaris andSciurus vulgaris calotus Shukhov I. I. , Obzor rasprostraneniya 
okhotnich'ikh i promyslovykh mlekopitayushchikh v Eniseiskoi gubemii, Turkestanskom krae i v 
Uryankhaiskoi zemle (A Survey of Game and Commercial Mammals of the Yeniseisk Province, Turkestan 
Territory and the Urianghai Land), p. 15. 

1930. Sciurus vulgaris martensis (sic'. ) Zverev M. D. , Obzor gryzunov yuzhnykh chastei 

Minusinsko-Khakasskogo okruga (A Survey of the Rodents of the Southern Part of the Minusinsk-Khakassian 
District), Sbomik Srednesibirskogo geograiicheskogo obshchestva, p. 83. 

1930. Sciurus vulgaris subsp. Yakovlev E. O. , K granitsam rasprostraneniya promyslovykh zverei i 
ptits V Turukhanskom krae (Ranges of Commercial Mammals and Birds in the Turukhansk Territory), 
Trudy Zoologicheskoi sektsii Srednesibirskogo geograficheskogo obshchestva. Issue 1, p. 57; Pogudin A. A. , 
Ocherk okhotnich'ego promysla Kirenskogo okruga (A Survey of Commercial Hunting in the Kirensk 
District), Trudy Lesnoi opytnoi stantsii. Issue VII, 1930, p. 95. 

1935. Sciurus vulgaris j eniss e j e nsis Ognev S. I. , Sistematicheskii obzor belok (A Systematic 
Survey of Squirrels), Byulleten' Nauchno-issledovatel'skogo instituta zoologii Moskovskogo 
univereiteta, No. 2, p. 47. 
TYPE LOCAUTY AND DEPOSITION. No. 75 male, 23 Oct. 1926. Lower Tunguska, Turukhansk 

Territory, N. P. Naumov. Type in the MSZM collection. 
407 MATERIAL EXAMINED: Holotype and 340 skulls and pelts. Forty of the pelts examined were with skulls. 

DIAGNOSIS. Color very variable. The most typical color of the winter 
fur is an intense bluish ash-gray with slight dark gray mottling. Tail 
generally reddish rust-chestnut with a marked black tinge. There are also 
a few red-tailed, dark-tailed and chestnut -tailed specimens. Winter fur 
on back very light ash-gray in red-tailed specimens. Summer fur of three 
types: l) Back black-gray-chestnut; 2) Back dark chestnut; 3) rich rust- 
red-ocher specimens. White area of belly much reduced. Groin and inner 
sides of legs of dark color, not white. 

Length of skull 51.3-53.7 mm (M. 51.9 mm); condylobasal length 45. 1- 
49,2 mm (M. 47.2 mm); zygomatic width 29.2-32.2 mm (M. 30.6 mm); length 
of upper tooth row 9.1-10 mm (M. 9,5 mm); hind feet 56-62.5 mm 
(M. 58.8 mm); tail 165-191 mm (M. 172.2 mm). 

DESCRIPTION. Color of winter fur very variable. Most specimens of 
this subspecies have a characteristic color of the tail which is transitional 
between the red and chestnut types (see below). In typical specimens the 
color of the winter fur on the back is an intense bluish ash-gray (light 
neutral-gray and neutral-gray) with fine dark gray mottling. Color of 
hair roots on back deep slate -gray (deep neutral-slate -gray), more intense 
than inS. v. martensi. Gray color of anterior part of head, nose, 
forehead and cheeks with a buff-rust tinge (between wood-brown, buff-brown 
and drab). Buff-gray shades at the sides of throat. Middle of throat with 
a white wedge. Color of ears between intense ocher-brown (kaiser-brown) 
and black. Gray color of back sharply contrasting with white color of belly, 
which is generally much less than in S. v. martensi, as the inner parts 
of the haunches and antebrachia are gray. Soles dark blackish gray. 



354 



Tail generally reddish rust- chestnut with marked black or deep black- 
chestnut tones. Color of hair on tail as follows: roots smoky blue -gray 
(light neutral-gray), followed by broad rust-brown bands (kaiser-brown, 
hazel) and by broad and well marked black or black -chestnut bands and 
rust colored tips which are especially marked towards the tip. Tail of 
specimens with rust-colored ears usually more rust colored than in dark- 
and black-eared specimens. There are all possible transitions to dark- 
chestnut-tailed and dark-tailed specimens. In these squirrels the color 
of the back is usually more intensely gray than in red-tailed and chestnut- 
tailed specimens; color of back intermediate between mouse -gray and 
neutral- gray. 

408 The color of the tail varies between dark red and dark brown. Together 
with these squirrels occur red-tailed specimens, with a light ash-buff-gray 
back. Ears and tail rich rust-yellow-ocher (between tawny, sayal-brown 
and mikado -brown). Roots of hair on tail pale ash-gray (light mouse -gray), 
followed by broad rust-colored bands. Tips buff-yellow with a light ocher 
tinge (cinnamon -buff). These specimens closely resemble S . v . 
martensi in their light color, but differ from it in the smaller white 
area on the belly. Inner sides of hind legs and antebrachia light gray with 

a light buff -rust tinge. 

Summer fur of three types: l) darkest specimens. Color of back 
intense black- gray (between fuscous -black and fuscous), flanks a little 
grayer than back (between fuscous and chaetura-drab). Anterior part of 
head and cheeks with a slight gray tinge. Feet black- chestnut. Tail of 
chestnut or dark type. I did not see any black-tailed specimens in summer. 
2) dark-chestnut specimens (apparently the most common type of summer 
fur). Back generally dull dark chestnut (slightly more brown than natal- 
brown). Head and cheeks with a buff-gray tinge. Feet deep chestnut- 
black-brown. Flanks with a dark gray-chestnut transition zo