Division of “Gumads if Bite A Ut
S.1. OGNEV swim
*, ides rTUTION
iM SBRR SE RRS
MAMMALS OF THE US.S.R.
inp 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
f
t
, f
aL]
f
a
S.I. Ognev
MAMMALS of the U.S.S.R.
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
y Russian English
page page
BORenerd 25.06 RA «gi OG SATs AMON RIRe, « qth Lie gMeEE Ee ike bY Schiele L. 8 ix
Order Five
GLIRES LINNAEUS. RODENTS
Fart Suborder. DUPLICIDENINAMA 4 diasp . emo lets Bae 12 2
marmuey Vacomyidae, Pik aS) oi. iss «+05 cn qm enee SMEAR Nee - 13 3
Genus Ochoton'a Links jis 6 @ ado chets Oh). ae Bode es: 14 4
Subgenus Pik a dbaceé pede is. ce tlevawnhds oP SOS Y Qi ererwet 18 8
149. Ochotona (Pika) alpina Pallas (1773).
Altai or Alpine spill es. 6, apees & me slersitedo ope et ancters hey 19 9
309. Ochotona (Pika) alpina alpina
Pall, (i773). Central Altai (pileany, .6. ts eleven 31 20
310. Ochotona (Pika) alpina nitida
Hollist. (1902). «Bast Altad joie. wi «mcr « 32 21
311, Ochotona (Pika) alpinachangaica
SHDS MOV Tana y OTE ooo isles esc. gage yeyelle « 33 22
312, Ochotona (Pika) alpina scorodu-
movi Scalon (1935). Skorodumov's
Alipine foukar . pon tt use Po eke ogo 88S we cyene oye orone 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) hyperborea Pallas (1811).
Wortheria polka is alas,» 4 evade cueterirey wil aie crenatarekes «take 38 26
315. Ochotona (Pika) hyperborea
hyperborea Pall, (1811), Northern pika 48 a5
316. Ochotona (Pika) hyperborea
ferruginea Schrenck (1858). Russet
Norgiein, piliay io cyep tet aye oliisiou ys ¢ ayatelo. cers yeh oye 49 36
317. Ochotona (Pika) hyperborea
cinereoflava Schrenck (1858), Sl 38
Okhotsk Northerm pias. dwajseeyel. cial: ober os ape
318, Ochotona (Pika) hyperborea
uralensis Flerov (1927). Northern
Wralipiies o 18 ca aeece ne BEACH Orer sone ceca 51 38
ili
319, Ochotona (Pika) hyperborea
mantchurica Thos, (1909). Central
Siberian Northern pika) (7% s:.)a theta «siete 52 39
Subgenus Octhotona Link: occ ei sanes con See eee 55 42
151. Ochotona (Ochotona) daurica Pall. (1776).
Dawisan PIR «os sna Pies bie wel oie dengan eee 56 43
320, Ochotona (Ochotona) daurica
daurica Pall, (1776), Eastern Daurian pika 62 48
321, Ochotona (Ochotona) daurica
altaina Thos, (1911), Western Daurianpika 63 49
152. Ochotona (Ochotona) rufescens Gray
(1842). Afghan pikarey Pres eee es ans ener so 65 52
322, Ochotona (Ochotona) rufescens
regina Thos, (1911), Kopet-Dagh pika 76 61
153. Ochotona (Ochotona) pricei Thomas
(1911). . Price's: pika.-. ja72cevine cadens s naceie 77 62
323. Ochotona S, (Ochotona) pricei
Thomas (1911), Mongolian pika or Price's
pik /sAted toate ass see ete eietmate at Mets 82 67
324, Ochotona (Ochotona) pricei
subsp. nov. Price's Semipalatinsk pika.... 84 69
Subgenus C onothoa Lyont (5: 43 5ae-. 2 os Stes sede athe 85 69
154, Ochotona ({Conothoa) macrotis Giinther
(1875). Large-eared pika or Indian pika ....... 85 70
155. Ochotona (Conothoa) rutila Severtzov
(1873), Red piltarces.e1 Pate Meee ements. ats 90 74
Species of the Ochotona Sula ade O. Sh imerand:
GLOUP: —cetosie s aielevesvererecete DEO ae ee eters Sele ite 97 80
156. Ochotona pusilla Pallas (1768), Small pika
or chehuistica 7— 0 sates B/ NGls. he atahs aioe Rha tetame re ate 98 82
325. Ochotona pusilla pusilla Pallas
(1768). European chekushka ............ 104 89
326. Ochotona pusilla angustifrons
Argyropulo ii Siberian chekushka ... 105 90
Family Leporidae, Hares ........ SPINOR rT e a aed dee itn. 106 91
Genus Lepus Linnaeus (1758) 4378.40.28 0I.. SOP GES, 108 93
Subgenus Eulagus Gray J. E. (1867). Desert and
Folad hare ...0..0ade ee Poet tees oe Oks the Me: BAe 109 93
157, Lepus (Eulagus) europaeus Pall, (1778) 112 96
327. Lepus (Eulagus) europaeus euro-
paeus Pall, (1778). European common
hare... «meee eae TE or woe Bae as 140 t2t
328, Lepus (Eulagus) europaeus
hybridus Pallas (1811), Central Russian
common Rate sche aah ener re & phn Beret 142 123
iv
Seo:
330.
Sail
332,
333.
Lepus (Eulagus) europaeus
tesquorum Ogn. (1923). Steppe common
Lepus (Eulagus) europaeus trans-
sylvanicus Matchie (1901), Roumanian
or Crimean common hare ..........-. Sod
Lepus (Eulagus) europaeus
caucasicus Ognev (1929), Caucasian
common hare....... Be eo UA CFO, OF 2 ao) Ones ON
Lepus (Eulagus) europaeus cyrensis
Satun, (1905). Transcaucasian common hare
Lepus (Eulagus) europaeus caspicus
Ehrenberg (1830), Caspian common hare...
158. Lepus (Eulagus) tolai tolai Pall. (1778).
Molar Ware) eeerertelo-
159. Lepus (Eulagus) t
e ecoeeeve so
ibetanus Waterhouse (1841).
DeSertwhare ccc cis sce sce se ele) siie seers cal ste Bete rei cee
334,
334', Lepus (Eulagus) tibetanus lehmanni
natio zaisanicus Satun, (1907). Zaisan
desert hare ......cccceseseseeees Sood
335. Lepus (Eulagus) tibetanus quercerus
Hollister (1912). Chuya hare .............
336. Lepus (Eulagus) tibetanus subsp.
Kopet-Dagh desert hare ...... SUMNER Ree ON
337. Lepus (Eulagus) tibetanus desertor-
um Ogn, et Heptner (1928). Desert sand hare
338. Lepus (Eulagus) tibetanus buchari-
ensis Ogn. (1922), Buchara desert hare....
339, Lepus (Eulagus) tibetanus pamiren-
sis Blanford (1875), Pamir desert hare .....
Subgenus Lepus s, str, Linnaeus (1758) ..........6.---
160. Lepus timidus Linnaeus (1758). Blue hare .
340. Lepus timidus timidus Linnaeus (1758).
Northern blue hare ..........-.++ss:00- ao08
341. Lepus timidus kozhevnikovi Ogn.
(1929), Central Russian blue hare..........
342, Lepus timidus sibiricorum Johans.
(1923). West Siberian blue hare ...........
343, Lepus timidus transbaikalicus
Ognev (1929), Transbaikal blue hare .......
344, Lepus timidus gichiganus J. Allen
Lepus (Eulagus) tibetanus lehmanni
Severtzov (1873). Syr-Darya desert hare....
(1903). Gizhiga, orOkhotsk blue hare .......
147
150
153
155
158
161
172
186
190
191
193
195
tSi7
199
201
201
237
240
242
244
247
Bae.
128
131
i] 333}
345.
346,
Lepus timidus mordeni Goodwin
(1933). Maritime territory blue hare .....
Lepus timidus orii Kuroda (1928),
Sakhalin bitie have Po, Vette at tie he eee
347, Lepus timidus kolymensis Ogn.
(4922), : -Kolyma bime*haret i027 fr. is 22
348. Lepus timidus begitschevi
Koljuschev (1936). Taimyr blue hare......
349, Lepus timidus tschuktschorum
Nordquist (1883), Chukchi blue hare .....
Genus ATtolayas Ognewr. Sekt s a tere ce cote tekstas ets
161, Allolagus mandshuricus Radde (1861),
Manchurian ‘hare ite. > Stee e'e © sialenies tis mane ahane
Genus Oryctolagus Lilljeborg (1873) ...... at ele an Rie ere
162, Oryctolagus cuniculus Linn, (1758).
Theta @Rt Meee crus 5 fed een ase he SGN Pare cain eat
Second Suborder. SIMPLICIDENTATA .........0c00- yh aA ch
Family Pteremyidae, Flying Squirrels. os scies scorers wasidad ale
Genus Pteromiys.@) Cavier (1617) cece cai oe ch leon ik vau a
163, Pteromys volans L. (1758). Common flying
Squadra 95.8 ase ds ake Se 0 04-0 ere enatnste areata
350,
350,
352,
353;
354.
355.
356.
357.
358,
Family Sciuridae ,.
Pteromys volans volans Linn. (1758).
Common flying squire) di aanes ona hamects
Pteromys volans ognevi Srogan.
(1936). Western flying squizrel or Ognev's
flying squirrel) 1f.-.),\. p « bite ate oroppiew't wa eres a
Pteromys volans gubari Ogn, (1934),
Western Siberian flying squirrel ..........
Pteromys volans betulinus Serebr,
(1929), Semipalatinsk flying squirrel .....
Pteromys volans turovi Ogn, (1929),
Central Siberian flying squirrel ...........
Pteromys volans athene Thos, (1907).
Sakhalin flying squisrel | .cccacwgurwe sypicap
Pteromys volans incanus Miller
(1918). Yakut flying squirrel ............
Pteromys volans anadyrensis
subsp. nov. Anadyr flying squirrel ........
Pteromys volans arsenjevi Ogn.
(1934), Ussuri flying squirrel ............
Subfamily Sciurinae, Squirgeis:.)./.4 Fs, 2. ss aden Bose ede:
Genus Sciurus Linnaeus (1758) ....... sous wintalee te = oie bate %
Subgenus Sciurws "Linnaeus (1758) 0:2: 6:00 0 sis'sts's a'efea's
vi
306
308
310
311
314
316
319
321
323
325
B27
327
329
EF
217
219
221
222
225
226
226
236
239
246
248
249
252
268
270
271
272
274
276
278
281
282
284
285
286
287
164. Sciurus vulgaris Linn, (1758).
AlN Seema Sooacoabcooau0o0K By OSORNO ION
359. Sciurus vulgaris ognevi Migulin
(1928). Central Russian squirrel........
360. Sciurus vulgaris fedjushini Ogn.
(1935). Belorussian squirrel ....... cess
361. Sciurus vulgaris formosovi Ogn.
(1935). Northern squirrel, or Formozov's
Snare sa ropes aucel sy aaenelaWeNe ial adi ieiie) aaaiian oe ast 500
362, Sciurus vulgaris varius Barrett
Hamilton (1899) ex Kerr (1792). Koka
Reeth) ooo ddooacodoWoOOdooednROOUIDO
363. Sciurus vulgaris ukrainicus
Migulin, Ukrainian rust-colored squirrel
364. Sciurus vulgaris kessleri Migulin
(1928) oisessler's Squiirelie re ettaeueie iy i-l
365. Sciurus vulgaris bashkiricus
Ogn, (1935). Bashkirian squirrel .......
365'. Sciurus vulgaris bashkiricus natio
uralensis Ogn, (1935). Ural Mountain
SQUITTOU i cienctareiete noose a oud tertoninatss Ryo Sane
366. Sciurus vulgaris martensi Matschie
(1901), West Siberian squirrel .........
367, Sciurus vulgaris exalbidus Pallas
(2778); slieteut squirrel iy. vals chai: cieclersie: «lc
368. Sciurus vulgaris kalbinensis
Selevin (1934). 'Borovushka" or the Kalbin
SQUATTCU SY Macs wate a olor ate cNeHehees ener oneucksseeholte
369. Sciurus vulgaris jenissejensis
Ogn, (1935). Yenisei squirrel ..........
370. Sciurus vulgaris altaicus Serebren.
(1928). Altarsquimrel’ 2.55 0.5. a oe se
371. Sciurus vulgaris jacutensis Ogn.
(1929) )i Yakut squirrel ))53. 0.2% Sratetetet
371', Sciurus vulgaris jacutensis natio
anadyrensis Ognev (1929). Anadyr
Squirrel varie sie Merstoneistotes slisttene ieodetauteuetts :
372, Sciurus vulgaris fusconigricans
Dwigubski (1804), Transbaikal squirrel
373. Sciurus vulgaris mantchuricus
Thos, (1909). Manchurian squirrel......
374, Sciurus vulgaris rupestris Thos.
(1907). Sakhalin-Amur squirrel .........
Subgenus Mrenme Ss! HOSS fics sve ceieie «a e'ale.s scbaodoc
165. Sciurus (Tenes) persicus anomalus
vii
388
389
391
393
395
397
397
401
405
406
409
410
415
416
417
419
421
Bee
287
333
336
337
338
340
342
344
345
346
348
353
354
356
357
361
362
364
366
367
Gmel, .(1778). . Caucasian squimvel iis cawais «os 2»
Subfamily Spermephilonsinae sink five sae es oie eater 6 9 ee, wn
Genus Spermophilopsis W. Blasius (1884) .........
166.Spermophilopsis leptodactylus Licht,
(1923). Long-clawed Ground SQuirrel cui. ee Ss
375. Spermophilopsis leptodactylus
leptodactylus Licht. (1823) ........
376. Spermophilopsis leptodactylus
schumakovi Satunin (1908), Kushka
or Shumakov's ground squirrel ......... ar
Subfamily Marmotinae. Chipmunks, Ground Squirrels and
IMEDMNEH Camo Oud mono uOU ODED Moo oounbanc6é6déanc
Genus Eutamias@@roucssart (1S80)e eae. wns ce wins «
167, Eutamias sibiricus Laxm. (1769).
The Ciipmuinks | ars':e.0.a ale = eaetaters (tele ans a: a elas o's
3//, Eutamnrais sibiric us saba tres
Laxmann (1769), Central Siberian
CHIPURAGI ‘ateras sualersietanel cece ol tarrette wer aie oa
378. Eutamias sibiricus striatus Pallas
(1778). West Siberian or European
GHIPMAGTE fasiece- 1 eunera ake uctacoren eore he as eis
379, Eutamias sibiricas jacutensis
Ognev (1936). Yakut chipmunk.........
380, Eutamias sibiricus orientalis
Bonhote (1898), Ussuri chipmunk ........
381, Eutamias sibiricus lineatus
Siebold (1826). Amur chipmunk ........
SUPPLEMENT ‘TO VOLUME IEE, cc. sous diste ciaielc (or ie masteretane > aus
Tables 1-61
Maps I-IX
viii
Re PB:
422
432
433
435
452
454
456
458
483
485
486
487
488
490
a
368
S77
378
380
394
395
397
398
421
422
423
424
425
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., 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 areference 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 spermophiles
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
158
10
arey uouruIOD pammyded yyIM xoJ
r )
11
12
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 processes 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.
Testes lying either permanently in the abdominal cavity or in an inguinal
sac, rarely inthe scrotum. Uterus double (uterus dupiex). 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 adaptec
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:
1 (2). Incisors i 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,
1 :
2 (1). Incisors T 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: oe number of molars varying between ; in Leporidae to : in
'3Ps
Ochotona. In some extinct Ochotonidae, e.g., Prolagus Pom. (lower
European Miocene), the number of molars was reduced to - 3
Molars rootless, hypselodont with deep enamel folds on crown; cement
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. Asa 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 ofthe 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 méxillary 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
14
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 FAMILIES OF DUPLICIDENTATA
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 aa
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
poe Leporidae.
Family LAGOMYIDAE, PIKAS
1866. Lagomyidae, Lilljeborg, W., Systematisk 6fversigt af de gnagande daggdjuren, Glires,
Nova Acta Soc, Scienc,, Upsala, Vol. VI.
1897, Ochotonidae, Thomas O,, 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 Mammiféres 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: iz; cS; Ps: 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
Lacépéde and Conothoa Lyon*.
* Pikas of the pusilla-tibetana 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.1, 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.
15
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 Ochotona Link
1773, Lepus Pallas P., Reise durch verschiedene Provinzen des Russischen Reichs, Bd. II, Anhang, S.
701-702 (partim!), Le pus Linnaeus, 1758.
1795, Ochotona Link H,F., Beitrage zur Naturgeschichte, Bd.I, Theil 2, S, 74 (type Ochotona minor
Link,=Lepus ogotona Pall. =Lepus dauuricus Pallas),
1799, Pika Lacépéde Bern. G,E., Tableau des divisions des Mammiféres, Paris, p. 9 (type Pika
alpinus =Lepus alpinus Pallas),
1800, Lagomys Cuvier G., Legons d'anatomie comparée, Vol.I, tab. I (Lagomys Storr, Prodromus
Methodi Mamm,, 1780, pp. 34-40; this description was apparently made according to an Arctomys
specimen).
1867. Ogotoma 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 PallasiizLagomys ogotona
Waterhouse, Lepus ogotona Pallas),
1904, Conothoa (subgenus!) Lyon M.W., Classification of the Hares and Their Allies, Smiths,
Miscellan, Collections, Vol. XLV (1), parts 3-4, p. 438 (type Ochotona roylei 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); i1-1*.
Rostrum of flattened skull narrow, this being a reliable difference
between the genus Ochotona andthe 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, witha
* [p=pectoral, i=inguinal. }
16
Single enamel cap. In Leporidae it has a double enamel cap. The structure
of the second upper molar (m2) 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 macrotis Giinther. Large-eared pika.
3 (2). Interorbital region of adult skull relatively narrow and convex in profile,
4 (5). Length of skull not over47mm, Vibrissae 40-55mm long; most vibrissae white, few very dark-
brown. Ears with well marked pale margins, Color of summer fur on back pale yellowish straw-gray to
duller and darkerstraw-gray. Belly pure white. Neck without a broad pale collar,
Ochotona daurica Pall. Daurian pika.
5 (4). Length of skull in adults usually 46mm to 53mm, Most vibrissae dark brown; longest
vibrissae with long white tips, 45 to 55mm 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 43 mm 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 22mm; diastema longer than 9mm, Hind foot (without claws) more than
30mm,
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,. cinereofusca. 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,
PIATE L
Red pika (Ochotona rutila Sev.)
Drawing by V. A, Vatagin
Altai pika (Ochotona alpina Pall.)
Drawing by V. A, Vatagin
17
11 (10). Skull much smaller, less than 43mm 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 22mm; upper diastema less than 9mm long, Hind foot (without claws)
less than 30mm long. Soles grayish brown or gray; whitish gray in winter.
Ochotona hyperborea Pall. Northem 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 45mm.
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.
=} cwccce co weg
see a-p---n- Ee
oe
FIGURE 1. Skull measurements in pikas FIGURE 2. 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
frontals,
(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,
18
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 (=<Conothoa Lyon).
Type species: O. daurica Pall. Palatine and incisor foramina fused
and exposed.
2) Pika Lacépede, 1799 (FLagomys C. Cuvier).
Type species: O. alpina Pall. nec Storr., 1780. Palatine and
incisor foramina separate. Skull flattened in profile.
3) Ogotoma Gray, 1867. Type speciesO. 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
Giinth. andO. rutila Severtz. which were considered as one species by
many authors would have been ascribed to different subgenera according to
the structure ofthe 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 Bichn. 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 toO. 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 includingO. daurica, curzoniae,
ladacensis,srufescens, prices.
2) Roylei group including O. roylei, macrotis, rutila,
erythrotis, gloveri, wollastoni, nubrica.
3) Alpina group including O. alpina, hyperborea, princeps,
schisticeps pucollanis’:
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 Lacépede
1799, Pika Lacépéde B,G,E, Tableau des divisions des Mammiféres, Paris, p. 9.
1S
Skull strongly flattened in the interorbital and frontal region; frontal
fenestrae aS arule 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 70mm.
Scree-inhabiting, semisylvan and sylvan species distributed in the tundra
to the shores of the Arctic Ocean.
Type species: Pika alpinus =*Lepus alpinu’s Pall.
The following species also belong to this subgenus: Ochotona
hyperborea.Pall., O. princeps Richardson, @.\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 (wild 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 Reichs, II,
1773, S. 701, Taf. II, B, (half-tone plate ofentire 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) Schreber I, Ch. , DieSdugethiere, 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., Russkaya fauna (Russian Fauna), II, 1851, p. 791-793, plate 63, fig. 2 (entire animal,
color plate); 6) Brandt, I.F., Untersuch, tiber die craniolog. Entwick. und Classification der Nager, 1855,
Pl, XI, Fig. 11-20 (skull and teeth); 7) Schrenck L,, Reisen und Forschungen im Amur Lande, I, 1858,
S. 150, Pl. VIII, Fig. 2 (Lagomys hyperboreus var, cinereofusca=Ochotona alpina cinereofusca (entire
animal, colorplate); 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.)}, 1, 1933, p. 81, fig. 74 (skull, dorsal), fig. 75 (D)
{incisor foramina of skull); 14) Sbornik: '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 45mm in fully grown and old
Specimens, 57mm in very large specimens. Interorbital space flattened
and broad in adult and old specimens, 5to 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 subspecies are numbered separately in this volume.
20
fenestrae. Vomer ventrally covered by margins of premaxillaries,
palatine and incisor foramina separate. Most vibrissae black or dark
brown, 60 to 70mm 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 inits gray or brownish gray color, streaked with
blackish hair. Completely melanistic individuals occur frequently.
Body length may reach 250mm (usually about 200 mm); hind foot may
be 35mm long (usually about 30mm).
FIGURE 3. Skull of Ochotona alpina alpina Pall., FIGURE 4, Skull of Ochotona alpina
dorsal, Specimen No, M. 327018 Dec, 18, 1929. Southern alpina Pall,, ventral (same specimen as
Altai, near Argat, Katon-Karagai. S.I1,Ognev'scollection in Figure 3)
Drawing by Yu. A. Kostylev
FIGURE 5. Skull of Ochotona 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 Ochotona alpina alpina Pall.,
(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
23
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
FIGURE 8. Skull of Ochotona alpina
alpina Pall., No, 149, 1897. Central Altai,
A, A, Silant'ev, KZMAN*. Very large
specimen
Drawing by Yu. A. Kostylev
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, pl. XXIX), belly
pale, yellowish ocher, intermediate
between pinkish cinnamon and
cinnamon-buff (Ridgway, pl. XXIX)
although lighter.
The color of pikas from the Chuya
Alps (Chegan-Burgazy -O. alpina
nitida Hollister) is more intense, of
a dark brown shade (intermediate
between mikado-brown and snuff-brown,
Ridgway, pl. XXIX). Flanks colored
a brighter rust-brown, close to Sayal-
brown and cinnamon (Ridgway, pl. 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,
pl. 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,
* [KZMAN — Collectionof the Zoological Museum of the Academy of Sciences of the U.S.S. R.]
12
24
i) 2h pl. XXIX). Flanks and belly in winter fur much browner with a clear
ocher tinge (cinnamon and clay color, Ridgway, 1912, pl. 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, pl. 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, larrivedatthe
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 Ochotona 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 collectionof 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 of O. 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. Sscorodumovi 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 andO. 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 andO. hyperborea are certainly very closely related.
A taxonomist could, however, identifyO. 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
25
Hangay Mountains. The range of O. alpina reaches from the Altai
Mountains to Transbaikalia in the east and the Hangay Mountains in the
south. The Altai pika lives indifferent habitats, namely screes androcks. 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 Hollister
andO. h. mantchurica Thos. occur together in the Sayan Mountains.
O. a. cinereofusca Schrenck andO.h. mantchurica Thos.
occur together in the Gazimur Mountains; O. a. changaica subsp.
nov. andO. 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 Sorex araneus L. andS. minutus L., in
spite of the fact that both have large and small subspecies. Nor has anyone
united Pipistrellus nathusii and Pipistrellus 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
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. Hollister (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 inthe 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 innorthwestern 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 inthe 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 inthe 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 of O. 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 asO. hyperborea var. cinereofuSsca.
B.A. Kuznetsov (1929) found this species, which he considered to belong to
species alpina (subsp. cinereofusca) in the Udinskii Range. V.N.
Skalon (1936, loc. cit.) thinks thatO. 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. Itis
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 of O. alpina extends. O.
Thomas (1909) identified the Ussurian pikasasO. cinereofusca.
27
This identificationis, 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 of O. alpina inthe 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 of O. 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 - 1m 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.
"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.
Varvinskiiand Makar'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, withacentral 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 inhigh-mountainscrees inthe 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
* [Verst: 1.067km, ]
28
far below us. The last cedars had disappeared. Inthe 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 pika!''*
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 Spirea overgrowth (at 1500 to
1800m altitude). On another occasion pikas were found above the
forest zone (at about 2000m)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. Inthe 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 (Ribes nigrum, R. atro-
purpureum), gooseberry (Ribes grossularia), very fragrant and
Sweet raspberry (Rubus melanolasius) etc.
Later, screes become densely overgrown with Caragana
arbomes censindMonicerajicioe rulea,; (Lb. ‘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'chariver (atributary of the Chulyshman river), and also in the upper
* B.A. Kuznetsov, Gryzuny Semipalatinskogo okruga (Rodents of the Semipalatinsk District), — Byulleten'
Moskovskogo obshchestva ispytatelei prirody, 51 (1—2):115-116, 1932,
17
29
30
reaches ofthe Bidui river (a right-hand tributary of the Abakan river). The
screes have here completely lost their primitive character: alluvialsoilis
never found, and stones androcks are coveredwithmoss. 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 onthe 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 Calamagrostis
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 cinereofuSsca 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. cinereofusca 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 sawO. 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 materialonthe snow-covered Narym Mountains, I observed that the
pikas were busy preparing hay for the winter. In the morning they could be
observed scurrying amongthe 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,
31
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-hoarding 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, Pteridium
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 thatO. a. cinereofusca 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 (16-17cm 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. 2409, 27 Jul. 1936 Kuraiskaya
Steppe, Altai, A.M. Kolosov) or inearly August (No. 23589, 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
32
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 ona
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. alpina.
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 thatthe skinofO. 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 Eversmann E,, Addenda ad celeberrimi Pallasii Zoograph. Rosso-Asiatica, p. 3
(type from vicinity of Uimon),
1899, Lagomys alpinus 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 gryzumami 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 47-57mm (M. 50.2);
condylobasal length 44-53mm (M. 46.8); zygomatic width 22-26mm
(M. 23.6); width of skull at level of auditory meatus 22-25.7mm (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 Ochotona alpina, 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 C. 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), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk,
VII, p. 300,
1912, Ochotona nitida Hollister N., New Mammals from the Highlands of Siberia, Smithson.
Miscellan. Collect., Vol. 60, 14, p.4.
1913. Ochotona nitida Hollister 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 15-51mm (M. 47.6); condylobasal length 41-48 mm (M. 43.7);
zygomatic width 21.2-24mm (M. 22.5); width of skull at level of auditory
meatus 21.2-23mm (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. Hollister (1912) described this form from a
specimen** from the Chegan-Burgazy mountain pass. He stresses the dark
color of the summer fur of O. nitida onthe 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.
** Hollister had a series from the Chegan-Burgazy mountain pass and from Tapuchi (31 pelts) in addition to
this type.
21
34
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 oahangaica 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) & northern Hangay, basin of the Orhon, 1925, P,K. Kozlov; 3) 20 220 9, 8 Jul. 1929, southwestern
Hangay, screes in Ohton-Tengri; 4) No,59 9, 19Jul. 1929Hangay, rocks along river Bombotu (an
eastern tributary of the Buyanta Gol river; 5) No. 26, 1925 (23), 15 Aug. 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
of O. a. changaica in summer fur paler than typical form; upper
surface of light yellowish collar duller thaninO. a. alpina. Color
of winter fur more grayish white than in nominal race.
Dimensions smaller than inO. a. alpina. Skull length 46-48.9mm
(M.48); condylobasal length 42.7-45.2mm (44.3); zygomatic width 22.5-
23mm (M. 22.7); width of skull at level of auditory meatus 22.7-23.2 mm
(M. 23).
SYSTEMATIC NOTES. This subspecies is closely relatedtoO. a.
nitida in its dimensions; it resembles the southern AltaiO. 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), Sbornik Rabot
Protivochumnoi organizatsii Vostochnosibirskogo kraya, Vol.1, 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, inthe collection of the Zoological Museum of the Academy of Sciences.
MATERIAL EXAMINED, I 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 182.8mm; 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 5.4mm; 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 Ochotona alpina alpina Pall. Color of flanks usually
different from that of belly and resembling that of back (Figure 9).
1254
22
35
ADDITIONAL DESCRIPTION. According to V.N. Skalon, the top of the
head and spine of specimens of O. 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 mouSse-gray.
Melanistic individuals never occur.
\
ale i
—ses5
FIGURE 9, Skorodumov's alpine pika, FIGURE 10, Barguzinpika, Ochotona alpina
Ochotona alpina scorodumovi svatoshi Turov, No. M. 39899 19 Aug. 1923,
Scalon, type No. 25,009, ZMAN Shumilikha river, Sosnovka, Baikal, S.S. Turov;
collection collection ofS. I. Ognev
Drawing by Yu. A. Kostylev
The following is a description ofthe color ofatypeofO. 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
SYSTEMATIC NOTES. Ochotona a. scorodumovi is most
closely related toO.a. cinereofusca; it differs from it by its
deeper color with black tones on the back. According to V.N. Skalon,
O. a. scorodumovi isalso somewhat largerthanO. 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 mantchurica 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 witha
skull length of 39.2-42mm, which appear in the table (No. 10-11),
undoubtedly belong to hyperborea.
The only certain difference between O.a.scorodumovi andO.a.
cinereofusca is thus the color. It may be more correct to regard
O.a.scorodumovi as anatio of cinereofusca.
GEOGRAPHICAL DISTRIBUTION. Borzya District of Transbaikalia.
313. Ochotona (Pika) alpina svatoshi Turov. (1924). Barguzin
pika
1923, Lagomys alpinus 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, © faune pozvonochnykh zhivotnykh severo-vostochnogo
poberezh'ya oz. Baikala (The Vertebrate Fauna of the Northeastern Coast of Lake Baikal), Doklady
Rossiiskoi Akademii Nauk, p, 110 (Barguzin 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 svatoschi Vinogradov B.S,, Mlekopitayushchie SSSR (Mammals
of the USSR), Gryzuny, p. 83.
TYPE LOCALITY AND DEPOSITION. M 3985, summer 1922, Shumilikha ravine, Sosnovka,
Barguzin Reserve. Type in S,1I, Ognev's collection.
MATERIAL EXAMINED: 15 pelts and skulls,
DIAGNOSIS. Skull small, length in adult specimens 44-47.2mm
(M. 15.6) [sic]; condylobasal length 40.7-45 mm (M. 42.8); zygomatic width
21.1-22.2mm (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 asa subspecies of hyperborea. This form occupies in fact
an intermediate position between O. alpina andO. 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. svatoshi 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
37 quite possible. It is also unknown how far south this form occurs. An
alcohol specimen of Ochotona alpina caught 327km 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, 1,S. 150, Taf. VIII, Fig. 2.
1923. Ochotona hyperborea cinereofusca Soweby, A. de C., The Naturalist in Manchuria,
II-Ill, p. 174.
1929, Ochotona (Pika) cinereofusca Kuznetsov, B, A. , Gryzuny vostochnogo Zabaikal'ya
(Rodents of Eastern Transbaikalia), Izvestiya Assotsiatsii nauchno-issledovatel'skikh institutov.,
2 (1):87-89,
1933. Ochotona hyperborea cinereofusca 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 40-44.9mm (M. 42.6); zygomatic width 21-23mm
(M. 21.7); width of skull at level of auditory meatus 21-23 mm (M. 22.2). See
mieure Il, p. a7.
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, pl.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, pl. 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.
cinereofuscaandO.a.svatoshi differ from eachother asa 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
38
39
cinereofusca is "richer" than that of svatoshi is incorrect. The
color of O.a. cinereofusca is duller, grayer and yellower than that
of svatoshi, and lacks the intense ocher, reddish, and orange tones
characteristic of the summer fur of fully grown svatoshi. These
differences can be clearly seen in the series.
V.N. Skalon (''Nekotorye zoologicheskie nakhodki v yugovostochnom
Zabaikal'e'' (Some Zoological Discoveries in Southeastern Transbaikalia),
1935, p.85) described a peculiar subspecies, O.a. Sscorodumovi from
near the village of Kailastui (Borzya District). The skull of this form is
larger than that of Och.a.cinereofusca according to him (vide supra)
the mean skull length of O.a. scorodumovi was 46.45mm; the skull
of specimens from the Baiolga Valley (north of the Gazimur Range)
identified by V.N. Skalonas O.a.cinereofusca,is much smaller, its
mean lengthbeing42.75mm. TheskullofspecimensofO. a.cinereofusca
collected by B.A. Kuznetsov on the Udinskii Range is not smaller than that
of O.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 andO.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 recordedcinereofusca 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
itacineneofusca.
O. Thomas (1909) found a specimen of this subspecies in the Ussuri
Valley. His observations should, however, be verified.
150. Ochotona (Pika) hyperborea Pallas (1811),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 nérdliche
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,
Tungus); chipa (Orochi on the Amur); teperkan (Orochi on the Gilyui); piski [squeaks] (Orochi in the
Far East).
PRINCIPAL FIGURES: 1) Schrenck L, V., Reisen und Forschungen im Amur Lande, I, 1858, S. 150,
Taf, VII, Fig. 1(Lagomys hyperboreus var, normalis), Taf, VII, Fig.2(L.h. var. ferruginea),
Taf. VIII, Fig, i (L.h. var, cinereoflava); 2) Flerov K.K. Pishchukha severnogo Urala (The North
Ural pika), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, Vol. XXVIII, 1927, pp. 138-144, figs, 1-3,
pp. 141-142 (entire animal); pl.1V, 1-4, (photograph of skulls).
MATERIAL EXAMINED: 206 pelts (most of them with skulls).
DIAGNOSIS. Skull small, length in old animals usually less than 45mm,
42.5mm in very large and old individuals. Interorbital space of adult and
old specimens flattened and 4.3-5.7mm wide. Anterior third of nasals
without Spoon-shaped widening. Lateralcontours of posterior part ofnasals
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.
x A
FIGURE 11, Amur pika, Ochotona FIGURE 12-13, Skull of Northem pika, Ochotona hyperborea
alpina cinereofusca Schrenk, cinereoflava Schrenck, (dorsal and ventral), No. M. 3989 7/III
No. 588 23/VI 1928. Nerchinsk 1928, Ol'skiy Island, Rassvet Bay, G.D. Dul'keit, S.I, Ognev's
District, B. Kuznetsov collection
Drawing by Yu. A. Kostylev 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 inthe nominate form, grayish brown witha
light straw rusty tinge. Reddish tones (sayal-brown, Ridgway, 1912 pl. XXIX)
well marked on lower part ofback, upper part of back more straw colored.
Flanks darker than back; belly whitish with a distinct straw-yellow tinge.
O.a.cinereoflava has anevenpaler fur. Its summer fur is distinguished
by arich gray-yellow color witha weak rusty tinge. Flanks rich rust-yellow.
40| O.h.ferruginea hasa saturated rust-red-ocher fur, which becomes intense
red-rust-brown inthe 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 inits 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 210mm, usually about 150-160 mm; hind foot 30mm
long, usually 26-27mm or 22-30mm.
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
41
The skull of O.hyperborea 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 alpina the diastema is much longer than the
distance between the alveoli of the molars.
2) The shape of the nasals of hyperborea differs from that of
alpina. This can be seen ina series. The width of the nasals is almost
equal anteriorly and posteriorly inhyperborea while the nasals of
alpina are markedly widened anteriorly.
3) The brain case of O. hyperborea is shorter and more rounded
than in alpina.
4) The orbits of O. hyperborea are more rounded, not as elongate
as inalpina.
All these differences can be clearly seenina series. The differentiation
between the two species is difficult, because the subspecies of alpina
contain some rather small forms, e.g.,O.a.svatoshi, O.a.
cinereofusca etc. However, the above-mentioned characters,
particularly the rostralstructure, always determine the species ofa specimen.
The following table of measurements may prove useful for general
orientation.
O, alpina O. hyperborea
Lenett ofekull .% . B..) item peek esas more than 43mm less than 43 mm
Condylobasal Jength®» 5.) 01%. «el ve fom o ek 5 Cees ny 8 ay! Mt? tt Ate
Zygomatic width... . ere at 2D) tt now 22 "
Width of skull at level of eee meatus \ S. >» ney PAV a PAT
Diastema # | f «Non 2 a «fo « Seite... Satay fat oi" aren oF
Row of upper molars “,-0°. « « « - 7 « Me fe mn" reall da Se"
Body length? s) ts. ah ere tte . oe. ee ss) ~)6 USUallymorethan 190mm _ usually less than 190 "
Length of hind foot without cacti wie iamcine 3o J = i! " ume 30> gf if Mary gl 307 4
Lengthi ofvibmssae iit).\tiie tia oes " 4 Leg6O isi y MS Pac ld 60.
There are five subspecies of Ochotona hyperborea intheU.S.S.R.,
which are described below.
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 inthe 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
42
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.hyperborea. 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 voicesonCape Epra-Ukun. He also found pikas
in the tundra near the village of Pokhodskii (85km from Nizhne Kolymsk
along the Kolyma river). According toA. Bunge (1887) this species reaches the
Yanaand Adycharivers 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 ''Yakutskaya 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
specimens 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
Adycha river); Tumarariver (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 along the Yuzhno-Muiskii Range (I. Polyakov, 1873). To the east it
is found along the Yablonovoi Range, where B.A. Kuznetsov (1929) caught
an excellent series near Kyker. The expedition of B.S. Vinogradov and
S.1. 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 asOchotona sp. is alsoa
Norther pika, (Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, vol. 17, 1912 (1913), p, 397-398.)
29
43
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 mantchurica 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
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
alpina 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 the 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
44
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.
FIGURE 14. Northern pika, Ochotona 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 7-8km from the rock outcrops on
the bank of the Yenisei river (along its lower reaches, 2km 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
45
46
by them, as it consists entirely of gabbro. Pikas are the most widely
distributed animals in the area; no other animal is as common. i
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, andthe 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 10m 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 ownterritory, usually one of the small
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 tracks, sometimes they are concealed under stones
and covered with earth, like tunnels. Thenest is usually constructed among
stones, inacrevice or ahollow. Sometimes one animalhas severalburrows.
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, 9cm wide and 6cm high. Burrows are usually small,
FIGURE 15, Postures of the Ural Northern pika, Ochotona hyperborea uralensis Flerov
Drawing by K.K. Flerov
1-1.5m long, without many branches but with several entrances, connected
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 9-10cm 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 ofa
pika. The preferred diet of pikas in the northern Urals is Veratrum
lobelianum. They usually cut off the upper part of the stalk. The
stalk is cut into pieces 5-10cm 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 Vicia cracca. According to 1. 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 of Ochotona, 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 45-48mm long were found
in the uterus of a female killed on 23 July. At the same time young
specimens 14-15cm 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 inO.hyperborea occurs very late. A specimen in
molt was caught on Jun. 8, 1930. There isaspecimen inthe 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. 2.515.
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 inthe
middle of the molt. Pikas caught in mid-September bore analmost 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 Amur territory that
pikas do not hibernate, coming to the surface of the snew 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 SchrenckL,, Reisen und Forschungen 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
Daggdjursfauna, 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 hyperborea Ognev, S.I., Mlekopitayushchie Severovostochnoi Sibiri (Mammals of
Northeastern Siberia), pp. 70-71.
1926, Ochotona kolymensis Ognev, S.I., 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 specimina 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 known 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, gravish
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 35-40.7mm (M.
37.3); condylobasal length 32.8-38.3mm (M. 34.4); zygomatic width 17.3-
20.7mm (M. 18.5): width of skull at level of auditory meatus 17.2-21.3mm
(M. 18.5); length of row of upper molars 6.1-8mm (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, pl. 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. hyperborea 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. ferruginea Schrenck 1. Reisen und Forschungen im Amur Lande,
I,S, 148-149 (South Kamchatka, Khalzan Mountains).
1887, Lagomys alpinus Bunge A,, Berichte iiber die Expedition nach Neusibirisch, Ins,, S. 40 (nec
Lepus alpinus Pall’),
1922, Lagomys kamtschatica Dybowski B,, Arch, Tow. Nauk, Lwowie, III, p. 10 (nomen nudum! )
1934, Ochotona (Pika) hyperborea turuchanensis, Naumov, N.P,, Mlekopitayushchie
Tungusskogo okruga (Mammals of the Tungus District), Trudy Polyarnoi Komissii Akademii Nauk, Issue
17, pp. 38-42(Lower Tunguska, Uchami trading station).
TYPE LOCALITY AND DEPOSITION, Kamchatka, Khalzan Mountains. Type at the ZMAN collection,
MATERIAL EXAMINED: 57 pelts (most of them with skulls),
36
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,
pl. XXIX). Belly dull whitish yellow. |
This subspecies is larger than the typical form. Skull length 38-41mm
(M.40). According to N.P. Naumov (1934), the skull length of O.h.
turuchanensis reaches 43.2mm; condylobasal length 37-38.6 mm (M. 37)
(according to Naumov to 40.2mm); zygomatic width 19.03-21mm (M. 19.9);
width of skull at level of auditory meatus 19-21.2mm (M.19.8); length of
row of upper molars 7.2-7.3mm (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'' of ferruginea (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 turuchanensis with
O. svatoshi andO.h. uralensis, but he omits the Kamchatka
ferruginea. This form differs from the typical and other forms in the
same characters asturuchanensis.
We regardturuchanensis as synonymous withferruginea; 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 Tumarariver); 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
(100km 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
rangesofO. h. ferruginea andO. h. cinereoflava (vide infra) lie very close together here, but
the boundaries have not been exactly determined,
Bi
51317. Ochotona (Pika) hyperborea cinereoflava Schrenck
(1858). Okhotsk Northern pika
1858. Lagomys hyperboreus var. cinereoflava Schrenck L., Reisen und Forschungen im
Amur Lande, I,S. 150, Pl. VII, fig. 1.
TYPE LOCALITY AND DEPOSITION. No, 689, 18Feb. 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
yeliowish tinge.
Skull length 40-42 mm (M. 40.8); condylobasal length 37-39.21mm
(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 of O. 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 Severnogo Urala (The Northern
Ural pika), - Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, Vol. XXVIII, pp. 138-144, fig. 1-3;
pp. 141-142, pl.IV, fig. 1-4.
TYPE LOCALITY AND DEPOSITION. No. 16107 910 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 35.3-36.6mm (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 7.1-7.5mm (M.
Weak
Body length 167-183mm (M.175.5); hind feet 22-27mm (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
relatedtoO.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., Puteshestviena 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.
1909, Ochotona hyperborea mantchurica 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.1,, Materialy po faune gryzunov
yuzhnoi chasti Eniseiskoi i Irkutskoi Gubernii i Zabaikal'ya (Material on the Fauna of Rodents of the
Southern Part of the Yeniseisk and Irkutsk Provinces and Transbaikalia), Izvestiya Sibirskoi stantsii zashchity
rastenii, No, 2, p. 15 (taiga regions of Transbaikalia, nec. cinereofusca Schrenck:).
1927, Ochotona hyperborea mantchurica AllenG,, Lagomorphs Collect, by the Asiatic Exped. ,
Amer. Mus. Novitates, No. 284, pp. 1-2(aseriesof 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 (1):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), Sbornik
Srednesibirskogo geograficheskogo obshchestva, p. 85 (River Ana, a tributary of the Abakan, western
Sayans),.
TYPE LOCALITY AND DEPOSITION: No. 23 o, 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
54
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, pl. 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 9 16 July 1928, Vicinity of Kyker, Sretensk District, Transbaikalia, B. A. Kuznetsov
Drawing by Yu. A. Kostylev
Skull length in fully grown specimens 39.3-43.3mm (M.41); condylo-
basal length 36.7-40.2mm (M. 38); width of skull at level of auditory
meatus 19.2-21.1mm (M. 20.3); length of upper tooth row 7-8mm (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, pl. 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 witha
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, Ochotona hyperborea
mantchurica Thos,, lateral (same specimen as in Figures 16, 17)
SYSTEMATIC NOTES. O. Thomas (1909) describedO.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 43.4mm; condylobasal length 41mm,
maximum width 22mm; nasals 14.2mm 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.
vy
FIGURE 19, Mandibula of Central Siberian Northern pika, Ochotona
hyperborea mantchurica Thos., (samespecimen 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 to O. 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
4]
55
56
Territory, rivers Lena, Yana, Olenek, etc.) southwards. The differences
between typicalO.h. hyperborea from the eastern parts of the Chukot
Peninsula andO.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 inthe Yeniseisk
Territory, in the taiga near Krasnoyarsk, western Sayans (river Ana, a
tributary of the Abakan), near Lake Hébs6gol 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 asO.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.mantchurica intheir 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 inO.daurica,
curzoniae, rufescens fused and vomer ventrally free; inO.
ladacensis and O. pricei 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, brownishinsummer 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.
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
57
Ochotona curzoniae Hodgson, O.ladacensis Giinther, O.
rufescens Gray, O. pricei Thos. also belong to this subgenus.
151. Ochotona (Ochotona) daurica Pall. (1776).Daurian pika
1776. Lepus dauuricus Pallas P., Reise durch verschiedene Provinzen des Russischen Reichs, III, S. 692.
NAMES: Daurian pika (English); Otogono or Ogotona (German),
LOCAL NAMES: pishchukha-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. III (whole animal), tab. IV A, fig. 16 A and 16 B (skull front and lateral); 2) Schreber1. Ch, , Die
Sdugethiere, Erlangen, IV, 1792, S. 915-918, Tab. CCXXXIX (in color); 3) Simashko Yu. , Russkaya fauna
(Russian Fauna), Vol. II, Mlekopitayushchie, 1851, pp. 793-794, pl. 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.I, Mlekopitayushchie, Issue 5, 1894, pl. XXV, fig. 1-5 (photograph of skull),
pl. 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,
Il, 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), pl. VIII,
fig. XV (photograph of entire animal).
MATERIAL EXAMINED: 47 pelts, mostly with skulls.
DIAGNOSIS. Skull rather large, its length exceeds 40mm in fully grown
specimens. Interorbital space narrow, 3.2-4.2mm; 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, 40-55mm 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.
Winter fur differs markedly from summer fur in its long and lighter
hair, and gray color with a straw-yellow tinge.
Body length to 195mm; hind feet 31mm.
ADDITIONAL DESCRIPTION. In addition to the above-mentioned
characters, the orbits are rounded (not ovalasinO. rufescens, theskull
of which resembles that of daurica); base of triangular fenestrae
relatively low, elongate longitudinally.
The base of the triangular fenestrae of O. 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 of O. 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 tothat of O. 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.
58 FIGURE 20, Skull of Daurian pika, from Shansi (Ningwufu). O.huangensis
Ochotona daurica daurica Pall. from Kansu is apparently closely related
No, M3997 2 june D922. Wear Bosra: to O.daurica, and is possibly only a
ae eae subspecies of this species. The close
Vinogradov and S.1, Obolenskii, S.I, q 4
Oiev's coliveties relation between huangensis ani
Drawing by Yu. A. Kostylev daurica was pointed out by O. Thomas
(1908). The description of O.
huangensis byP. 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, pl. 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 inWestSiberia 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 inthe 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]
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59
in the ZMAN collection. A.N. Formozov (1929) recorded the Daurian pika
from the vicinity of Lake Orog Nur (45° N) on the eastern slopes ofthe Great
Altai. This species also occurs near Uliassutai [Jibhalanta] on Lake Hébségél,
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
tome by A.S. Lukashkin. G. Radde (1862) caught a large series near Tarey Nor.
R. Maak (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".
IfO.bedfordiae Thomas is consideredas identicalwithO.daurica,
the eastern range of O. daurica extends to the Shansi Province in
China, but this is still not certain.
N.M. Przheval'skii (E.A. Biichner, 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 Hébs6gél).
Colonies of O. daurica occur sporadically along river valleys inthe semi-
steppe and desert. Their favorite habitats are thickets oflris biglumis
and Stipa splendens near water. Small plots with fresh grass are
usually found between the scattered hillocks in which the pika builds its
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60
burrows. 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. Biichner, 1890) the Daurian
pika occurs inthe 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 foundin large numbers in
thickets of shrub along rivers."
GENERAL DESCRIPTION AND HABITS. M.N. Przheval'skii (E.A.
Biichner, 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'-pit'- pit', pi-pi-pi-pi"’.
This resembles the warble of some birds, e.g., that of Anthus
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 duringthe 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 inthe 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
numerors exits. The area of the burrow of one rodent is large, it may
reach 25-35sq.m. The number of exits is also very large, 15-20. The
entrances are about 5cm wide. The burrows of pikas apparently were
inhabited previously by voles because a) old stores of voles could be found in
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61
excavated burrows; b) the galleries were narrow (3cm 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 tanacetifolia, Artemisia SiverSsiana,
Potentilla sibirica and Medicago falcata (the order of the plants
in this list is according to their decreasing frequency).
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 inthe heap. In addition to iris, the haystacks contained some
grasses and artemisia. The stacks were about 50cm wide and 35-40cm
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,
Compositae, 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, itusesdew, although this is
rather rare here.
"It is a question what 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 summer. 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 2Sept. had5 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 105mm
long, 20 miles south of Ulan-Bator (Urga) on 18-19 May 1922.
A.M. Kolosov caught a young pika only 75mm long on 8 Jul. 1936 inthe
Chuya Alps along the river Chegan-Burgazy (2000m).
There is a young specimen, 130mm long, in my collection, which was
caught on 25 Jul. 1934 in the floodland of river Toshtugovu, a tributary of
the Shibita, 10km from Toshakty. B.A. Kuznetsov caught a young specimen,
134mm long, 0n10 Aug. 1928 inthe Aginskoe District near Lake B. Tsaga-
Nor. These data confirm the assumptions of A.N. Formozov mentioned
above.
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62
The knowledge about molts in this species is fragmentary. A.M.
Kolosov caught adults on the Chuya Alps on 5Jul. 1936 and9 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 10Jul. 1936 inthe 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 sawthe 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 (Archibuteo 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 Pall. (1776),
Eastern Daurian pika
1776. Lepus dauuricus Pallas P., Reise durch verschiedene 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 Siiden 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. andS.I, Obolenskii, Materialy po faune gryzunov yuzhnoi
chasti Eniseiskoi i Irkutskoi gubernii 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 LOCALITY AND DEPOSITION: In the famous work of Pallas "Reise" etc,, BD. III, 1776, p, 692, we
read: "Vivit in campis, montiumque declivibus arenosis apricis, per totam Dauriam.,.."'.
* [Sajene: 2,134m,]
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 EXAMINED: 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 38.5-44mm (M. 42.4); condylobasal length 36.3-41 mm
(M. 39.6); zygomatic width 20.4-21mm (M. 20.8); width of skull at level of
auditory meatus 19.2-20.2mm (M.30); length of upper tooth row 7.7-8.6mm
(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, pl.XXIX). Neck paler, witha
more yellowish tinge(pinkish buff, Ridgway, pl. 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 44-47mm (M.45.9); condylobasal length 40.7-45.2 mm
(M.43.3); zygomatic width 21.7-23mm (M. 22.2); width of skull at level of
auditory meatus 21-21.5mm (M. 21.2); length of upper tooth row 8.3-8.8mm
(M.8.6). See Figures 21-25.
Wp UG
Parte argh
FIGURES 21, 22, Skull of Daurian (western) pika, Ochotona daurica altaina Thos, 9 July 1936,
Collected by A. M, Kolosov from the Chuya Alps, Chegan-Burgazy
Drawing by Yu. A. Kostylev
65 Body length to 195mm (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 Ureg Nur (to the north of Lake Achita Nur), in Hobdo
(specimens in the ZMAN collection).
50
FIGURE 23. Skull of Daurian (western) pika, Ochotona daurica altaina
Thos, , (lateral) and mandible of same specimen as in Figures 21, 22
FIGURES 24, 25. Skull of young Daurian (western) |pika, Ochotona daurica altaina
Thos, , Altai, Chuya Alps, A, M. Kolosov Drawing by Yu. A. Kostylev
51
152. Ochotona (Ochotona) rufescens Gray (1842), Afghan pika
1842, Lagomys rufescens Gray J., Annals Magazine of Natural History, p. 266.
NAMES: The Afghan pika (English),
LOCAL NAMES: Gurukses-sichan (Turkmenian),
PRINCIPAL FIGURES: 1) Blanford W.F., Eastern Persia, Il, Zoology, 1876, pp. 83-84, pl. 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, pl. 38, fig. 14, 29; pl, 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, I, 1932, p.46, fig. 5 (dorsal, ventral and lateral views of skull); 5) Flerov K.K. andI.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
over 46mm, reaching 53mm. 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 inthe 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, 45-55mm 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, pl. XVI); tips of hair on back rust-brown; general
color rust-brown (sayal-brown, Ridgway, 1912, pl. 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 witha
yellowish tinge.
Winter fur duller, grayer and more brown than summer fur.
Body length up to 207mm (M. 205); hind feet to 35.2mm (M. 34.5); ear
to 27mm (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).
SZ
67
FIGURES 26, 27. Skull of Afghan pika, Ochotona rufescens regina Thos., dorsal and ventral,
No. M.4020 o 23 June 1925. Germab, Kopet-Dagh, S.1. Ognev's collection
Drawing by Yu. A. Kostylev
FIGURES 28, 29, Skull and mandible of
Ochotona rufescens regina
Thos, o 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 thanback, 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, pl. 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 anyon 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 inthis
region: 1) inthe estuary of river Sakis-Yab (Germab); 2)a large colony ina
ravine tothe west of the village, near the so-called ''Cossack camps''. We also
found it along the precipitous walls ofthe 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.1I. Argyropulo (1932) recorded the following localities for O.
rufescens from material inthe 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 andI.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 15km 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).
1254 54
FIGURE 30. Ochotona rufescens regina Thos,, 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. Varentsovy (''Fauna of the Transcaspian Region'' (Fauna Zakaspiiskoi
oblasti) Ashkhabad, 1894, p.14) found a colony of pikas ina 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
tl
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 Mitzel 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.
FIGURE 31, Afghan pika Ochotona rufescens regina Thos,, 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
he
73
FIGURE 32. Afghan pikaOchotona rufescens regina Thos., in the Kopet-Dagh Mountains
Photograph by S.I. Ognev
K.K. Flerov andI.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 of O. 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 sitson 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.1I, 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
the burrow. Pellets accumulate in rather large quantities, sometimes
large handfuls. They look like pellets of hares, but are much smaller
(3-4mm). 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 7cm. The exits are sometimes widened because the soil crumbles.
There is rarely onlya 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
a 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 1/2 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 pikais 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,1. Ognev
59
75
We found a burrow near the Dmitrievskii village in a crackina monolithic
rock. The animal had, eachtime, 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 7cm 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 Capparis shrub ona
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. Inspite 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 ina male (M.4016) caught on 30 June 1925 (Germab
Ravine) persisted in a few traces on the lower part of back and on the flanks.
Based on the examination of material from the Sumbar and Chandyr Valleys,
K.K. Flerov andI.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
7
6 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. Onthe 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. , DieSaugethiere Transcasp, , 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 regina Thomas O., On New Mammals from Central and Western Asia.
Annals and Magaz, of Natur. History, p. 762.
1929, Ochotona rufescens regina OgnevS.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.1,, Materialy po faune gryzunov Srednei Azii, Pishchukhi
Srednei Azii i Kazakhstana (Material on the Rodent Fauna 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.
MATERIAL 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 44-53.2mm (M.50.3); condylobasal length 44.5-
48.7mm (M. 46.4); zygomatic width 23-25mm (M. 23.8): interorbital
space 3-3.8mm (M.3.4); length of upper row of molars 9.4-10.2mm
(M.9.5); Length of body and head 150-219mm (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 typicalO. 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 ofO. 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,
6]
7 Il, 1876, pl. Vi, Figure 2). This specimen was caught in Korud, north of
78
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 TibetanO. curzoniae Hodgson
as definitely identical with rufescens.
This seems doubtful. W.T. Blanford (Fauna of British India, 1888,
p. 457) definitely states thatO. curzoniae Hodgson and O. roylei
Ogilby are closely related. According to O. Thomas (OnSome New Forms
ofOchotona, Annals and Magaz. ofNatur. History, ser.9, Vol. IX, pp.189-
190, 1922), curzoniae differs sharply from rufescens ina number of
characters. These questions can be solved only by examination of more
material.
I have discussed the geographical distribution of O. r. regina Thos.
in detail above.
153. Ochotona (Ochotona) pricei Thomas (1911). Price's pika
1911, Ochotona (Ogotoma) 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 severnoi Mongolii po sboram ekspeditsii
1925 goda (The Mammals of North Mongolia in Collections of the 1925 Expedition), 1929, p.91, fig. 15
(skull dorsal 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 of O. daurica, of average
size. Length 44-49mm. 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 foramina separate (Figures 36-38). Vibrissae dark
brown; longest vibrissae with long, white tips, 60mm 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, pl. XXIX) with some
grayer and rust tones (cinnamon and clay color, pl. 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 andI. M, Gromov (1934) consider the Asterabad pikas as belonging to this form.
79
FIGURE 36. Skull of Price's pika, Ochotona pricei Thos., No. M.3974 10September 1926,
Central Mongolia, Tsagan-Adzir Mountain, 100km south of Uliassutai [Jibhalanta], A.N.
Formozov, S.1I. Ognev's collection
Drawing by Yu. A. Kostylev
Winter fur on back beautiful straw-gray with a pinkish rust tinge
(avellaneous).
Body length to 200mm (usually about 180 mm); hind feet to 33mm
(usually about 30mm).
GEOGRAPHICAL DISTRIBUTION. This species was described by O.
Thomas (1911) 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 40km from Tsetsen Han [Ondér Han]*
A.1. 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 werk, 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.
aay
é
FIGURES 37, 38. Skull and mandible of Ochotona pricei Thos, (lateral)
No, M.3974
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 by O. pusilla south of Karaganda, northwest of
Balkhash in the direction of Berty-Su. V.A. Selevin recorded O. pricei
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), anda
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), inhisdescriptionof Ochotona 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.
80 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
81
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 inthe 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 Capra
sibirica graze, and near marmot colonies. The pikas take refuge in their
burrows in case of need.
"The range ofthis rodent thus reaches from the valleys to altitudes of
about 8000 ft.
"Burrows of Price's pika are usually under rocks or bushes. The widih
and shape of the entrances vary considerably. The entrance may be
either horizontally or vertically elongate, its width 50-120mm. There are
65
82
nearly always deep paths through the mound of earth excavated from the
burrow. O.pricei collects large heaps of rubble (to 1m long) consisting
of small stones (5X5cm), 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: 1) store
in a niche among rocks on 30 Jul. 1926 inthe lower reaches of river Tuin-
Gol (g 84) —Artemisia pectinata, Euratia coratoides,
Carex stenophylla, Tanacetum achillaeoides; 2) store under
stones on 1 Sept. 1946 between rivers Baydarag Gol and Dzabhan Gol
Tanacetum achillaeoides, Carex stenophylla, Artemisia
sp.; inathird 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.
WeneverheardO. 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* 9 (1 Aug. )
was suckling young (milk in the mammary glands) ; 191* 9 (1Sept.)hada
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 1Jun. 40 miles south of Tsetsen Han [Ondér Han]. Other specimens
of the same size were caught on 21 Jun. near Gun Burte; 25Jun. near
Ussuk and 11-24 July near the Artsa-Bogdo Range.
Subspecies of Ochotona 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. hamica Thos. **which was described from Hami[Quomul1] south of the
eagtern spurs of the Tien Shan) is also closely related to O. 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 GrayI,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, pl. 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 41-43.5mm
(M.43); zygomatic width 21-24mm (M. 23.1); interorbital distance 3.2-
4.7mm (M.4.2); width of skull at level of auditory meatus 22-24mm
(M. 23.2); length of upper tooth row 8.5-9.5mm (M.9).
Body length 180-215mm; hind feet 29-33mm; ear 21-26mm.
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 undisStinct.
Legs with whitish hair with a dirtier tinge on the soles. Claws horny
brown.
83 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 pallasii. He wrote as follows 'Ogotoma. Skull:
the orbits very large; space between the orbits narrow; nose narrow,
bent down".
"Ogotoma pallasii (Lagomys ogotoma Cuvier, Waterh.,
Chir iepus ogotoma Pallas, “Glires, 80, t. 3,°4a; f 16 a.)
BM."
Gray did not indicate the terra typica for his Ogotoma pallasii in
this fragmentary description. After examining the holotype of O.
pallasii atthe 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 (3mm). 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 ''Ogotoma pallasii", as there is
no description of this species, and as Gray partly confused his new species
with O. daurica Pall. (=FOgotona Pall.), we think that "Ogotoma
pallasii' is best considered as a doubtful synonym of O. pricei.
The Mongolian pika is closely relatedtoO. daurica, O.
curzoniae, O. ladacensis andO. 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) describedanew species from Taldura
in the southeastern* Altai, west of the Chuya Steppe. He considers O.
sushkini as closely relatedtoO. 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 onthe 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 200mm; _ hind feet (measured on pelt) 28 mm;
ear 18mm.
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 22.4mm; length of bullae osseae 11.7mm; length of
upper row of molars 9.2mm,
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 Roinanov 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 of O. pricei have a richer color than the Mongolian
ones. Inthis 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 of O. pricei except
in the western parts of this range, i.e., the Karkaralinsk Region and
Sary-Su (vide supra).
~ ©, 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
85
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):
116 (nearriver 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.
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 Conothoa Lyon
1904, Conothoa LyonM.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 inO. erythrotis andO. gloveri. Color of summer
fur, and that of winter fur in some specimens, often with well marked
rust-red tones. Pads ofdigits welldeveloped. Vibrissae long, to95mm.
* Described from specimen No, 17422 in the ZMAN.
69
86
The species inhabits high mountains.
Type of Subgenus: Ochotona roylei Ogilby.
This subgenus contains also the following species: Ochotona
macrotis Ginth., O. rutila Severtz., O. erythrotis Buchn.,
O: glover? Thos., O.. “wollastoni. Thos., O.snubrica hos.
GEOGRAPHICAL DISTRIBUTION. Pamir Mountains, Altai, Ferghana,
Gissar Range, Dzhety-Su Mountains, Ladakh, Himalayas (to 20,000 ft), Tibet
and Kansu.
154. Ochotona (Conothoa) macrotis Ginther (1875). Large-eared
pika or Indian pika.
1875, Lagomys macrotis 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 (partim!), Shnitnikov, V.N., Dzhety-Su (Dzhety-Su), p, 129.
1928, Ochotona rutila Vinogradov B,S., Mlekopitayushchie (Mammals), Sbornik: ''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, Giinther (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 Doba (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. Giinther records
it from Doba. 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 47mm.
Interorbital space flat and broad (5.2-6mm). 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 55-70mm. Ears rather long,
to 31mm.
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 200mm; posterior feet 32-34mm.
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 ofbuff-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 macrotis Giinther, dorsal and ventral.
No, 17950 9, 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-bufftinge, 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
sacana 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
87
88
FIGURES 42,43, Lateral view of skull and mandible of large-eared pika, Ochotona
macrotis Ginther, No. 17950,
O. Thomas remarks that O. sacana is very closely related to O.
macrotis. Series of O. sacana can be easily distinguished, according
tohim, fromtrue 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 of O. sacana according to
Thomas are: length of body and head 200mm; anterior feet 33mm; ear
28mm.
Maximum skull length 47mm; condylo-incisive length 44mm;
zygomatic width 22.5mm; nasal bones 16X5.5; interorbital space 5.3mm;
width of skull at level of auditory meatus 18mm; palatine foramina
13.5X4.7; width of palatine bridge 1.8mm; length of upper tooth row
(alveoli) 9mm (type caught in December 1913).
According to Thomas's description and his remark that O. sacana
andO. 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 of O. 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
Ofimacrietis; VO.) nut ilasg andOw enythrotis, forma, 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 of O. erythrotis is morphologically intermediate between
that of O. rutila andmacrotis; there are frontal fenestrae in
erythrotis which are absent inrutila. The structure of the palate
inerythrotis 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 itis
little visible ventrally (see Figures 44, 45).
89 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 macrotis (ona small scale);
the structure of the palate is the same as inmacrotis.
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 Biichner (dorsal and ventral), No. 1500 co‘, 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 -
Kuland Kara-Kulinthe ZMAN collection. O. macrotis also occurs in
mountains near Karakol (Przhevalsk), onthe Terskei Ala Tau, inthe 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 ofO. macrotis (like
O. rutila) inthis region is rather strange. It lives north of Issyk-Kul,
then it is found along the Alexander [Kirghiz] Range, then westward and
_turns 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 inthe southwestern corner of Dzhety-Su, is wedged into the
range of O. rutila.
"The ranges ofthe 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 (Doba).
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 Astragalus frigidus. V.N. Shnitnikov did not find
"haystacks", and he thinks thatO. 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 inthe 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
zhivotnykh (Vertical and Horizontal Distribution of Turkestan Animals): Izvestiya Obshchestva lyubitelei
estestvoznaniya, VIII (2):83.
74
oil
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 rutila Kashkarov D,N., Rezul'taty ekspeditsii v raion ozera Sary-Chilek (Results 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), Sbornik: "Zveri
Tadzhikistana'" (Animals of Tadzhikistan), Izdatel'stvo Akademii Nauk, pp. 250-251,
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: Typecollectedin 1867 by N. A. Severtsov in mountains near
Vernyi — now Alma-Ata (at an altitude of 7000-8000ft). Type number S.11,074, kept in the Moscow ~~y
Zoological Museum,
PRINCIPAL FIGURES: 1) Schaff E., Ueber Lagomys rutilus, Zoologische Jahrbiich, , 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, 1, 1932, pp. 37-42, fig. 2
(foramina incisiva); p. 39, Fig, 3 (skull dorsal, ventral and lateral); 3) Vinogradov B.S., Gryzuny
(Rodents), Sbornik: ''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 48mm, sometimes 52mm. Interorbital space
broad, 5.7-6.9mm, 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, 80-94mm; 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 pl.I, upper illustration) rich rust-
reddish (intermediate between cinnamon and sayal-brown, Ridgway, 1912,
pl. 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 onthe 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 25-28mm.
ADDITIONAL DESCRIPTION. The skull of the red pika differs sharply
from that of O. (Pika) alpina Pall. in the following characters: 1)
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 skullofO. erythrotis Bichn. differs markedly from that of
O. rutila inthe following characters: 1) presence of frontal fenestrae;
75
92
FIGURES 46,47. Skull of red pika, Ochotona 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 of Ochotona
rutila Severtz, (Same specimen
as in Figures 42, 43)
76
93
94
2) different form of nasals; 3) different structure of palate. The
external appearance of O. erythrotis resembles that of rutila
(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.1I. 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, 64km 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 onIskander-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 1925m 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,
} In the Gissar Range.
U7
95
(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 of O. rutila from the Terskei Ala Tau
seemserroneous, asthis region is inhabited by O. macrotis. The
range of O. 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, inthe
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 quite 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 observed 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 45km 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 insmall numbers in screes along
the road to the Talgar Mountain Pass at 2200 minsubalpine 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 ina
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. Theanimalis verytrusting, and lets
itself be approached to 3-4 steps (pl.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.1. 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 onthe 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. Severtsov that he 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.
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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 onthe 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 (1932) 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 (Alticola worthingtoni andEvotomys
frater).
Species of the Ochotona pusilla and
O. thibetana 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 slitinO. pusilla andinO. 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 45mm 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-Ili Ala Tau, near the Talgar
Mountain Pass, 29 July 1938
Photograph by S.I. Ognev
81
98
Species: O. thibetana Milne-Edw. (with subspecies); O. cansa
Lyon (with subspecies); O. sorella 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 relationship between these 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.1 (drawing of entire animal); tab, IV, fig. 3(ear), fig. 4 (skull frontal), fig. 5 (skeleton);
3) Schreber I, Ch. , Die Sdugethiere, Erlangen, B. IV, 1792, S. 906-910, tab. CXXXVII (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, pl. 63, fig. 1 (color
plate of entire animal); 6) Argyropulo A.I., Opredelitel' gryzunov 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, 1, 1932, p.55, fig. 7 (skull of Ochotona pusilla 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 43mm, usually 40mm. Interorbital space of moderate breadth
in comparison with size of skull, 3.6-5mm. 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 200mm, usually about 170mm; hind feet at most
30mm (usually about 27mm).
82
ulseqeA “vy ‘A Aq 2utmeiq
(‘ll@d @I1ltsnd eu0j,0y4d0) exYysnyeyo io extd [[eurs
Il ALVId
83
99 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, forexampleO.hyperborea, the skull
of which is of about the same size as that of O. 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 Argyrop. 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., southernSzechwan, O.t.
sikimaria Thos., Sikkim, O.t.zappeyi Thos. (Tatsienlu [Kangting)]).
The following forms are closely related tothibetana, and therefore
also to pusilla: Ochotona cansa Lyon (Kansu) with the subspecies
O. cansa morosa Thos. (Shensi); O.c. stewensi Osgood
(Szechwan); O. sorella 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 pusilla-thibetana group (vide
supra) and postpone the creation of a subgenus until more material
becomes 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 inthe 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 Argyrop, 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)itoccurs15km
* This species does not occur today on the right bank of the Volga.
101
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.
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 end 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).
FIGURES 57,58. Lateral view of skull and mandibula of Ochotona
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, at 55°30'N. He claimed
that the northern boundary of its range in the former Shadrinsk County
reached 56°N inthe 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 (in litt. 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. Belyaey,
1934). Specimens caught along the river Dzhamcha and near the village
Chelym in the former Karkaralinsk County are in the ZMAN collection.
102
87
Ifthe 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.1I. Obolenskii).
All this clearly shows that the range of O. pusilla extended considerably
further west in the past. Skeletal remains of O. 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 of O. pusilla inthe 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):114, 1932).
The pika lives most frequently in thickets of peashrub, wild rose,
Filipendula andothers inthe 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 onthe 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 of O. pusilla is very fragmentary.
P. Pallas once obtained a gravid female (30Apr.). 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.
88
Three females out of 5 caught by B.A. Kuznetsov in the Orsk District
vere gravid. The others were already suckling their young. The number
»f embryos and their size in several specimens were as follows:
No. 111— 6 embryos of 15mm length
No, 144— 11 " "35 u n
No, 145— 12 " "49" "
These females were caught in late summer: No.111 on 25 July; No. 144
and 145 on 29 July. We can therefore state definitely thatO. pusilla |
has at least two generations in Summer and autumn.
104 Subspecies of Ochotona 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 Quadrupeduin e Glirium ordine, pp. 31-45,
1811, Lepus pusillus. Pallas P., Zoographia Rosso-Asiatica, pp. 151-152,
1850. Lagomys pusillus Eversman E., Estestvennaya istoriya mlekopitayushchikh Orenburgskogo kraya
(Natural History of Mammals of the Orenburg Territory), pp. 207-208.
1851, Lagomys pusillus Simashko Yu., Russkaya Fauna. Mlekopitayushchie (Russian Fauna, Mammals),
Vol. II, pp. 790-791.
1891, Lagomus pusillus Nehring A., Die geographische Verbreitung der Sdugethiere 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 pusilla 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.1,, 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 40mm. Skull of old specimens little elevated
in the nasofrontal region.
Length of skull 35-38.7mm (M. 36.9); condylobasal length 32.0-36.7 mm
(M. 34.4); zygomatic width 18.4-19mm (M.18.7); width of skull at level
of auditory meatus 18.1-19.8mm (M.18.9); length of upper tooth row 6.9-
7.3mm (M.7.1). Body length 153-182mm (M.167.2); hind feet 26-27.5mm
(M2687)
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) witha
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
105
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 obshchestva estestvoispytatelei, XI, p. 177.
1900. Ochotona pusillus Kashchenko N.F,, Opredelitel' mlekopitayushchikh Tomskogo kraya (Key
to Mammals of the Tomsk Territory), pl. 30.
1932, Ochotona pusilla 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 Fauna of Soviet Central Asia). — Trudy Zoologicheskoge 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 39.7-43.2mm (M. 40.9); condylobasal length 37.2-
40.2mm (M. 38.2); zygomatic width 18.9-22.3mm (M. 20.3); width of
skull at level of auditory meatus 20-22.2mm (M. 20.8); length of upper
tooth row 7.8-8.8mm (M.8.3).
Body length 182-196mm; hind feet 26-30mm (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 astype, 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.
angustifrons, 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 inthe very old skull mentioned above. This skullhas several
individual characters which distinguish it from the other specimens of the
series.
GEOGRAPHICAL DISTRIBUTION. Omsk, Altai, Semipalatinsk and
Zaisan-Tarbagatai districts.
90
107
FIGURES 59-61. Skull of Ochotona pusilla angustifrons Argyrop., No, 2350, 8 June 1929,
Semipalatinsk District of Sergiopol [Ayaguz] Region, Znamenka Village, B. A. Kuznetsov
Drawing by Yu. A. Kostylev
Family LEPORIDAE, HARES
1821. Leporidae Gray J, London Med. Repos., XV, p. 304.
1857. Leporina Blasius J., Naturgesch. der Sdugethiere Deutschlands, S. 409,
1897, Lagidae Schultze E., Helias, XIII, S. 82; Catalogus Mammalium europaeorum, Ztschr. fiir
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.
9]
Dental formula: iz; ie ps; 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 Le pus with the subgenera:
1) Lepuss. str., 2) Eulagus, 3) Poecilolagus, 4) Boreolagus, **
5) Microlagus, 6) Macrotolagus
II Genus Sylvilagus with subgenera
7) Sylvilaguss, str., 8) Tapeti
Ill 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. Oryctolagus 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 Oryctolagus 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 hindlegs 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.
108 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 dorsally; 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, 11/5 - 23/4 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
* mz (as in Ochotonidae) only in Pentalagus Lyon.
** M, Degerbgl 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 andL. borealis).
1867. Eulagos Gray J.E., Annals and Magaz, of Natur. History, Sept. p, 222 (based on Lepus
mediterraneus Wagner, and L. judeae Gray).
1899. Eulepus Acloque Alex., Faune de France, Mammiféres, p. 52 (based on L. europaeus and
L. variabilis).
Ear measured from occiput longer than head, sometimes 1 1/2 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 (1 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 1 1/5 - 1 1/3 as wide as the interorbital space behind the postorbital
processes.
Nasal measured along their oblique outer side shorter, as long as, or
longer than, frontalsuture measured to its anterior point between the
nasals.
109 Width of posterior nasal opening 1 1/5-1 1/3 as wideas 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 1/8 -1 1/4 (Lepus) or 1 1/3-
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 terminal 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 Eulagus (instead of Eulagos) for consistency.
** Tail completely white in Lepus oiostolus Hodgson (I 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, I have not seen the Indian species and1 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
Oryctolagus*. Hares of Mesopotamia, Palestine and Arabia belongtothe
subgenus Eulagus. Hares of the subgenus Eulagus are absent in
== « coc owe
:
er
mec ee meee
FIGURE 62, Measurements of skull of hare FIGURE 63, Measurements of skull of hare
I — total length of skull; II — zygomatic I — basal length; II — condylobasal length;
width; II] — length of nasals (along suture); III — length of bony palate; IV — width of
IV — length of nasals (along outer side); incisive foramina; V-— length of incisive
V — width at level of auditory meatus; foramina; VI — length of pygomatic arch:
VI — lateral length of facial part of skull VII — upper diastema,
(anteriorly to alveolar point, but not to the
end of nasals).
* [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 Macrotolagus 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.
eatat 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 135mm; length of skull
usually more than 90mm (100mm 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, lengthof body usually less than 570mm; _ hind foot less than 135mm long, length
of skull usually less than 90mm.
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 100mm; length of hind foot without claws exceeding 150mm,
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
112
3 (2). Length of skull usually less than 100mm; length of hind foot less than 150mm, usually about
140mm, Clay tones of summer fur on back more intense and darker thaninL. 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
southem 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 winter 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 100mm. Weight 6-6.4kg.
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 123-160mm; length
of skull about 100mm, sometimes less, 89.3, or sometimes more than 101mm. Weight about 3.2kg (2.9-
3.9kg).
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 100 mm (92-105mm); length of hind foot 140-167mm, Weight about 4kg (3.2-4.4 kg).
L. e. caucasicus Ogn.
12 (9). Reddish clay tones on chest and flanks absent and replaced by dull straw-brown (in summer) or
dull straw-gray (in winter), Winter fur with characteristic vinaceous tinge.
Length of hind foot 144-150mm; length of skull usually less than 100 mm (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 P., Zoographia Rosso-Asiatica, p. 148 (Polonia, Lithuania).
NAMES: Common or brown hare (English); Lampe, Feldhase (German); Le liévre 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); kéch' (Komi-Zyryan); toshan (Armenian); urcheli
(Georgian); tkhakumakikh (Kabardian); kuyan (Balkar); tulai (Kalmyk).
PRINCIPAL FIGURES: 1) SchreberI., Sdugethiere, Erlangen, IV, 1792, Taf. CCXXXIII (Lepus
timidus = europaeus); Taf. CCXXXVC. (Lepus variabilis Pall. hybridus - tumak); 2) Bingley
W. , Memoirs of British Quadrupeds, 1809, p. 290 (color pl.); 3) Simashko, Yu. , Russkaya fauna.
Mlekopitayushchie (Russian Fauna. Mammals), Il, 1851, pp. 782-785, pl. 62, fig. 2 (color pl. of winter
fur, named Lepus aquilonius); pl. 60, fig.1; pl. 61, fig. 4 (color pl. and drawing of skull, named
Lepus europaeus); 4) Middendorf A. Th., Ueber die als Bastarde angesprochenen Mittelformen zw.
* Vinaceous buff-avellaneous (Ridgway, 1912),
96
113
114
L. europaeus undL. variabilis, Bullet, phys.-mathem., IX, 14, 1851, S. 29 (drawing of skull,
dorsal); 5) BlasiusI., Naturgesch, d, Sdugethiere 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.,
Atlas éfver Scandinavian Daggdjur, Stockh, , 1873, pl. 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
Schidel der Leporiden, Sitzb. d, Kaiserl. Akademie d, Wissensch,, math, Class, LXXVI, Bd.I, 1877,
Taf. Fig. 1 (skull ventral): 9) Giebel C.G,, Charakter, der Hasenschddel, Ztschr. fiir die ges. Naturwissensch. ,
LIIl, 1880, Taf, VIII-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, tothe 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, pl. 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.1I. 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.1, 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); pl. LXXV, fig, 3 (skull lateral); pl. LXXX and LXXXI, fig. 2 (skull dorsal and ventral); 18) Millais
I.G., The Mammals of Great Britain, Vol. III, 1906, pl. 49 (color.), pl., p. 8 (biological scene); pl., p, 16
(photograph); 19) Lénnberg E., On Hybrid Hares between Lepus timidus andl. europaeus from
South Sweden, Proceed, Zoolog. Society London, 1905, pp. 278-287, fig. 53 (teeth); fig. 54 (A.C. ) (part of
skull); 20) Schaff E., 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 (hare with five legs); 21) Hilzheimer M., Die
Hasenarten Europas, Jahresbericht d. Vereins fiir Vaterl. Naturk. in Wiirtenb, , 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. Soffel K., 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) Schaff E., Die wildlebenden Sdugetiere
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.
pl. XVI (head); p. 262, fig. 47 (skull lateral); fasc. XII, 1912, pl. XIX (3) (feet); pl. XX (7) (feet); fasc. XIV
1913, pl. XXIII (photograph of hare); 25) Ognev, S.1., Mlekopitayushchie Moskovskoi gubernii (Mammals
of the Moscow Province), 1913, pp. 283-298, pl. III, figs. 51-52, 54(photograph of skulls); pl. 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 gubernii (Mammals of the Tauric
Province), part 1, Gryzuny (Rodents), 1916, p.60, pl.IV, fig. 2(skull- L. europaeus aquilonius);
pl. lll, fig, 1 (skull of L.e. transsylvanicus); plate IV, fig.1(skull- L.e. caspius); 28) Filipchenko,
Yu, A.,Izmenchivost' i nasledstvennost' cherepa u mlekopitayushchikh (Variability and Heredity in the
Mammal Skull), Russkii Arkhiv Anatomii, Gistologii i Embriologii, Vol.I, No. 2, Petrograd, 1916, plate
XVI, fig. 2,4 (photograph of skulls - L. e. aquilonius); 29) Thorbom A., British Mammals, Vol. II, I,
1921, pl. 33 (in color); 30) Ognev S.I. Fauna pozvonochnykh Voronezhskoi gubernii (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 I1., 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); 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 Sbornik "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).
MATERIAL EXAMINED: 249 pelts and 291 skulls.
i
114 DIAGNOSIS. Head relatively small. Ear measured from occiput 1 1/5
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: po i=l a: Zen: towee. a.
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 100-120mm; length of ear (measured
from occiput) 120-171mm; length of hind foot 124-167mm.
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 of L. 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 of L. timidus.
Zygomatic arch narrow; in lateral view does not rise so steeply posteriorly
and does not delimit the lower side of the orbit as in L. timidus. Orbit
of L. timidus markedly different fromthat of common hare. Incisive
foramina of common hare generally longer than those of L. timidus.
Yu.A. Filipchenko (1916) gives the following variations of this character:
in L.timidus 22-27.5mm (M.24.30mm); in the common hare 24-29mm
(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. Mittelformenz, L. europaeus undL,
variabilis, Bull, phys, -mathem., IX, No. 14, 1851; I. Blasius, Naturgesch. d. Sdugethiere
Deutschlands, 1857, S, 412-414, 420-421; C.G. Giebel, Charakteristik der Hasenschadel, Ztschr. fir
die ges, Naturw., Bd. LIII, 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
gubernii (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
116
FIGURE 65, SkullofLepus europaeus caspicus FIGURE 66. Skull of Lepus europaeus
Ehrenb., No. M, 3021 o 27Jan. 1913, Khokhlatskoe, caspicus Ehrenb., No, M, 3021
Astrakhan Counties, N. L. Chugunov, S.I. Ognev's
collection
Drawing by Yu. A. Kostylev
Comparison of the mandibula of the common hare and of L. timidus
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
descripmon of 2) Wmuaduc. )
99
riy
FIGURE 67. Skull of Lepus europaeus caspicus Ehrenb,, No, M.3021
TIEN
ae
FIGURE 68. Mandibula of Lepus europaeus caspicus Ehrenb., No. M.3021
Yu.A. Filipchenko (1916) carefully analyzed 80 different cranial
characters. He concluded that only 8 measurements are really characteristic
for the skull of Ly. europa cals. and to afi Mi LOU SV i
Filipchenko gives these 8 measurements in form of indices.
Indices:
1) Lateral length of brain case (from inion (occiput) to posterior margin Lepus timidus Lepus europaeus
of orbit and base of upper margin of zygomatic process of temporal bone)
Se eT en Ee > 80 < 80
Lateral length of facial part of skull (from alveolar point to middle of
anterior or lower corner of orbit)
2) Length of bony palate > 65 <65
Width of incisive foramen
3)Length of tooth row $65 <65
Length of diastema
4) Length of zygomatic arch
= Le > 195 <95
Oblique length of nasal bone
* 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
118
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, toacertain 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 (6) 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:
Cin yf ePe,
n-
in which a - deviation from M, p- number of terms in each class, 3 —
Summation symbol. After calculating the quadratic deviation, the mean
error (m) can be determined because
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
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.,+Ym,? + m,?.We thus obtain the formula:
(M,—WM,) EV m+ m’,
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.
Mei ANE A NE
(M, — M2) = Vm,?+m,?
Lateral length of brain
case. hi aad mes 2.04 +0, 35
Lateral length of facial
part of skull. NA /ff== (0), 315)
Length of bony palate . 1.494015
Width of incisive foramen 1.5040, 20
Length of upper tooth row 2.11 +0, 24
Length of upper diastema 1,374 0,33
Oblique length of nasal
bones ats Sh ly/ 25 (0), 7/th
Length of zygomatic arch 1,78 +0, 51
10]
119
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 onthe average Shorter than that of L. timidus.
Filipchenko (1916) gives the following measurements of the upper tooth
row:
Lepus timidus Lepus europaeus
17.5-21(M. =19.28 mm) 15.5- 18.5 (M. =17.17mm)
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 anindistinct 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 55mm) 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,
pl. XXIX, and wood-brown, pl. 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 9chraceous 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
120) particularly marked on the chest; its general tone intermediate between
121
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
130mm.
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) area 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 65mm 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 arule, 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.
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SYSTEMATIC NOTES. There are 7 subspecies of the common hare in the
the U5. 6)5.
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 andcentral France, North Italy, Corfu;
5) L.e.corsicanus de Winton (Corsica, Sicily, Italy; 6) Lepus e.
transsylvanicus 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 of L. 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 of L. europaeus
follows. The common hare apparently occurs only rarely in southern
Finland, south of Savo, Istava (vide J. Valikangas, 1909, D.J. Wadén,
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).
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 40km 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 inthe 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 of L. 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. ]
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123
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, occursonthe 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 toL. 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 of L. timidus. It becomes
gradually more numerous further south towards Lodeinoe Pole constituting 2 or
3% ofthe populationofL.timidus. AccordingtoS.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. timidus caught (94 common hares and 13,557 of L. timidus).
It is very rare further east, in the Shenkursk District (on the Northern
Dvina). According to S.S. Folitarek and A.A. Maksimov, only 10 common
hares were found among 1706 blue haresexamined 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
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125
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, inthe central parts ofthe 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, according to 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 inthis 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
ahare. 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 ata 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 anirregular
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 UralRegion, in B.A. Kuznesov's opinion (1928). According
to M.K. Serebrennikov (1929) it is very rare in the northern Trans-Ural
forest-steppe. It is much more numerous inthe 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 40km
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 60-70km 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. Inthe 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 35-40km 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. Grevé, 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
The common hare has thus spread widely during the last century ina
northern, eastern and southeasterndirection. 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
TOTO 29
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 main 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.
128
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, andalsoofArtemisia. 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 arenartwus, Xanthium 'spinosum ete. 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 atypical 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 atrain for 2km. The
speed of the train was 55-60km. 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.
Sa
a
Ne
44 TS
Ay CRANKS SS
. oat Ss
t| j
es
Nas
Nore
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 hazelnut. The pellets of the white hare are more oval, with broadly
rounded tips and slightly compressed laterally.
The common hare is a good swimmer. I1.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,
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, 10-15km wide
131
* The differences between the track of the hare inslow and rapid progression, and the description of turns
and loops are given in the chapter on the biology of the blue hare.
111
on the Tsna and Moksha rivers. There were nearly no hares in these
places in 1933, although conditions in general were favorable.
ereets
FIGURE 71. Tracks of the common hare 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
132 in winter, leaving its tracks inthe 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
FIGURE 72. Common hare in summer
Photograph by S.I. Ognev
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, inthickets of high grass
and weeds, overgrown wheat fields, underbrush in ravines, tamarisk thickets
in the semidesert ete: 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.1. Barabash-Nikiforov
(1928) writes that in the Sivash Steppes hares are frequently attacked by
sea eagles (Haliaétus albicilla). They therefore often dig burrows
in the snow which may be 1.5m 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 andenters orchards where
it gnaws the bark of fruit trees, often causing great damage."’
133
o
eS
wip tel ys - ae
| 2 . Mai
ae RTL?
\ ; ; de & eg ne ar ee
fee A NR 85 3
) h Sy mae”
.
@ y .
FIGURE 73. Common hare hiding in winter
Drawing by A. N, Formozov
114
134
A.I. Yurin (in litt. 1934) informed me that the common hare is very
fond of Helianthus tuberosus.
He wrote: ''I found a new method of catching hares. Blinds are erectedin
Helianthus plantations, or near stalks of Helianthus 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 (Salvia silvestris), 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, andthe
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
lis
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, former Voronezh Province
Photograph by S.1. 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,
116
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,
137
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."
7 emewcs rece
FIGURE 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
ofthe biology ofthe 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: 1) 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:
117
1) 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 118
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 moundof 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.
LE)
139
140
'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
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 70km 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: 1)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 inthe steppe
and buries the vegetation, the hares migrate tothe villages. If they feed on
straw and hay, the damage is small, but iftheyenter 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.1. Zharkov, V.P. Teplov and V.1. 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
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, Aquila clanga, andrarely
sea eagles and eagle owls prey on hares. I.G. Pidoplichka (1930) 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 Lepus europaeus
327. Lepus (Eulagus) europaeus europaeus Pall. (1778).
European common 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 (nec 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. Lf{epus] t [imidus] albus Bechstein Joh. Matth., Gemeinniitz, Naturgesch. Deutschlands nach
allen 3 Reichen, I, Editio 2, S. 1096 (Thiringen, Germany).
1801. L [epus] t [imidus] flavus Bechstein Joh. Matth., Op. cit., S. 1096 ( Thiiringen, Germany).
1801, L [epus] t [imidus] niger Bechstein Joh. Matth., Op. cit., S. 1097 (Thuringen, Germany).
1811. Lepus timidus (nec Linnaeus!), Pallas P.S., Zoographia Rosso-Asiatica, pp. 148-149 (Poland
and Lithuania).
1857. Lepus timidus (nec Linnaeus!). Blasius J., Naturgeschichte der Sdugethiere Deutschlands, S.
412, 417 (Mittel-europaische Form L. timidus auct., L. campicola 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. FitzingerL., Sitzungl. d. Kaiserl. Akademie der Wissensch. , Wien, Mathem. Naturwissensch.
Klasse, LVI, S. 161 (Nomina nuda! Hares from Austria are described).
1910, Lepus europaeus Pall, Trouessart E.l., Faune des Mammifeéres 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 hare in Poland; 3) Pallas does
not mention the distribution of the common hare 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 andL. europaeus found in Poland and Lithuania on p, 5 ("...hanc,
mihi vix dubium est, varietatem vere hybridum esse productum, ex adulterioL. variabilis cumL.
europaeo, australiorem Poloniam atque Pannoniam occupante, progenitam et in istis regionibus
multiplicatam...'').
In his 'Zoographia 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 hare
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 100mm; hind feet less than 150mm
(usually about 140mm).
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 3.5-5.8kg. This is much less than the
weight 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 andL.e. hybridus. These
specimens resemble L.e. europaeus inthe 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 Mammiféres d' Europe, 1910,
p. 219).
Measurements of L.europaeus europaeus: length of skull 89-
99mm (M.97.7mm); condylobasal length 84-87.7mm (M. 86.4mm);
zygomatic width 44-48mm (M.46.1mm); length of nasals 44-47mm
(M.44.4mm); width of nasals 16-17mm (M.16.9mm); length of bony
palate 6.1-6.4mm (M.6.2mm); width of incisive foramina 11-12mm (M.
11.4mm); length of upper tooth row 17.0-17.6 mm (M.17.4mm).
G.S. Miller (1912) gives the following cranial measurements for the
typical subspecies: occipitonasal length 94-102.2mm; condylobasal
length 85.2-92.2mm; zygomatic width 45.2-50.2mm; interorbital width
18.4-24.8mm; postorbital width 11.4-15.2mm; width of brain case 31-
34.6mm; length of nasals (along oblique outer margin) 42.2-46.2 mm;
maximum width of both nasals 20.6-24.6mm; diastema 26.8-31.8 mm;
122
length of upper tooth row 17.2-19.2mm; length of lower tooth row 18.6-
20.2mm.
Measurements of specimens. 1) From Hasley, Zealand (Denmark)
2 males ad. and female ad.: length of body 660, 680 and 640mm; tail 80,
82, and 74mm; hind feet 136, 147 and 138mm; ears (measured from
occiput of live specimen) 138, 131 and 125mm. 2) Rheinhessen, Ingelheim
(Germany) male ad.: length of body with head 571mm; tail 98mm; hind
foot 135mm; ear (measured from occiput of dried skin) 120mm. 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 115mm. 4) Verdenberg, St. Gallen
(Switzerland) 2 males ad.: length of body with head 600 and 605mm; tail
90 and 95mm; hind feet 145 and 144mm; ear (measured from occiput)
114 and 118mm.
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 (1811).
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 Mammiféres, I. Paris,
p. 349.
1842, Lepus aquilonius Blasius1., Amtl. Bericht XIX Versamml. Naturf. und Aertzte, Braunschweig,
p. 89 (Central Russia).
143 1850, L. t[imidus] var. hyemalis Tumac, Eversman, E, , Estestvennaya istoriya Orenburgskogo
kraya (Natural History of the Orenburg Territory), part Il, 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 (=aquilonius
Blasius).
1851, Lepus aquilonius Simashko Yu., Russkaya Fauna. Mlekopitayushchie (Russian Fauna. Mammals)
II, pp. 782-785, pl. 62, fig. 2.
1871, Lepus timidus B. hyemalis (=hybridus) Bogdanov, M.N., Ptitsy i zveri chernozemnoi
polosy Povolzh'ya (Birds and Mammals of the Chernozem Belt of the Volga Region), pp. 175-176.
1889, Lepus timidus var. tumak 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 Mammiféres 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.1I., Mlekopitayushchie Moskovskoi
gubernii (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.
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 100mm; length of hind feet more than
150mm (usually 160-170mm),
ADDITIONAL DESCRIPTION. Weight of L.e. hybridus about
4-4.5kg. G.I. Polyakov reported that the heaviest of 40 common hares
killed in the former Bogorodsk County of the Moscow Province weighed
5.2kg. Larger weights are very rare.
G.1. 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
10.2kg. 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 guard 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-
106.5mm(M.101.9mm); condylobasal length 89.1-97mm (M. 91.9mm);
zygomatic width 46.9-51.2mm (M.48.5mm); length of nasal bones 46.1-
49.5mm (M.47.5mm); width of nasals 15.8-21.1mm (M.19.2mm);
length of bony palate 5.3-8.1mm (M.6.6mm); width of incisive foramina
11-13.5mm (M.11.8mm); length of upper tooth row 17.1-20.2mm (M.
18.6mm); hind feet 164-175mm; ear including terminal hair (measured
from forehead) 110-129mm.
SYSTEMATIC NOTES. Pallas (1811) noted the variety Lepus
variabilis var. hybrida. He gave the following description:
"Hyeme non plane alba, quam Rossi sequentis nomine Russak apellere
solent''’. He does not give the range of ''var. hybrida".
M.A. Desmarest (1820) in ''Mammalogie ou description des espéces des
Mammiféres", mentions ''Lepus hybridus Pall., russak des Russes".
He gives the following range for it: ''Habite certains cantons de la Sibérie
et de la Russie".
144
PLATE III
Central Russian common hare (Lepus europaeus hybridus Desm.) (foreground), Steppe common
hare (Lepus e. tesquorum Ogn.) (background)
125
145
I. Blasius (1842) mentions a hare from central Russia under the name
Lepus aquilonius.
Blasius (1857) in ''Naturgeschichte der Saéugethiere Deutschlands",
p.412,417 makes L. aquilonius a synonym of L. timidus
(=europaeus). On later pages Blasius separates a ''northeastern form’
(nordéstliche Form), to which he ascribes L. caspicus, L.
aquilonius, L. medius and L. variabilis var. hybridus.
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
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: 1) 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
146
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. Loénnberg (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:
1) premaxillary more slanting and longer than in the blue hare; 2), thirst
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
relatedto L.e. tesquorum (see below) occasionally occur among
typical L.e. hybridus. Such is, for instance, specimen No. 8.4976
(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: 1)
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. 8.3998 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
147
The Turgai-Orenburg Steppe is populated by the well differentiated
natioL.e. caspicus natioturgaicus Ogn. (see below).
GEOGRAPHICAL DISTRIBUTION. This subspecies is widely distributed
in the central part of eastern Europe. Intermediate forms between L.e.
europaeus andL.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.
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 hybridus andtesquorum. Most of the hares from Skopin
are intermediate betweentesquorum andhybridus, 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 natioturgaicus, 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 chemmozemnoi 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.I, and Gorbachev, S.N., Mlekopitayushchie Yugo-vostoka Orlovskoi
gubernii (Mammals of Southeastem 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 Ognevy 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. ]
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 100mm; length of hind feet 167-185mm.
Average weight about 6-6.4kg*.
ADDITIONAL DESCRIPTION. Anterior part of muzzle dull, straw-gray
(close to drab, Ridgway, 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-whiie 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
with 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 of L.e. tesquorum is never as rich as that of L.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 150mm 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 straw at tips. Tail dorsally blackish brown and white
ventrally. Outsides of ears whitish 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. i145)
the largest hare of 12,903 common and blue hares killed from 1815-1856 weighed about 20 pounds (i.e. ,
8 kg). This probably appliestoL.e. tesquorum, asN,V. Kireevskii hunted chiefly in the Kursk and
Orel provinces,
150
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.102.8mm); condylobasal length 87-97.5mm (M. 92.2 mm);
zygomatic width46-54 mm (M.49.2mm); length of nasal bones 43.2-
50.9mm (M.47.8mm); breadth of nasals 17.2-23.6mm (M. 20.1 mm);
length of bony palate 5.6-7.9mm (M.6.8mm); breadth of incisive foramina
10-15.2mm (M.12.2mm); length of upper tooth row 16.8-20.7mm
(M.18.2 mm).
Length of body with head 595-610mm; hind feet 150-185mm; 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 (1871)
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. timidus), differing from
it in the following characters: 1) 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.
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 of L.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.
hyo tans:
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 of L.e.tesquorum
extends. L.e.transsylvanicus occurs inthe southern Dnieper and
southern Tauric Steppes, but not inthe 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 isa
lighter color than that of the typicaltesquorum. Such a hare was sent
to me by V.V. Raevskii from the southern Don Steppes. The range of
L.e.tesquorum inthe 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. mediterraneus) Tikhomirov A.A and Korchagin A N., Spiski i
opisanie predmetov, nakhodyashchikhsya v Zoologicheskom muzee Moskoyskogo 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 mediterraneus) Nikol'skiiA,N., Pozvonochnye zhivotnye Kryma
(Vertebrates of the Crimea), appendix to Vol. XVIII, ''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., Rumidnische Saugetiere, Sitzb, der Gesellsch. naturf. Freunde
z. Berlin, S, 236,
1908. Lepus europaeus transsylvanicus Hilzheimer M., Die Hasenarten Europas, Jahresb. des
Vereins fiir Vaterl. Naturk. in Wiirttemb. , 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), part 1, 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.
13]
152
Size small: length of skull usually less than 100mm (89.3-101mm,
M.94.2mm); length of hind foot 123-160mm (M.147mm). Average weight
3.2kg (2.9kg-3.9 kg).
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 witha
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 89.3-101mm (M.94.9mm); condylobasal length 78.2-89 mm
(M.83.6mm); zygomatic width 43.2-48.2mm (M.46.1mm); length of
a2
153
nasals 37.2-46.5mm (M.43.1mm); width of nasals 16.8-21.1mm
(M.18.6mm); length of bony palate 5-7.8mm (M.6.2mm); width of incisive
foramina 9.6-13.3mm (M.11.1mm); length of upper tooth row 15.3-18.1mm
(M.16.9mm).
Length of body and head 550-600mm (M.564mm); hind feet 124-160mm
(M.147mm); ear (measured along concavity) 102-118mm (M.107.2mm).
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 intheir 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 RiverValley), pp. 74-75.
1924, Lepus europaeus transsylvanicus Ognev S.1,, Gryzuny Severnogo Kavkaza (Rodents of the
Northern Caucasus), pp. 7, 8.
1925. Lepus europaeus transsylvanicus Beme L, Rezul'taty ornitologicheskoi 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 transsylvanicus TurovS.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 OgnevS.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.1. 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 thatofsummer fur, not darkeras inL.e.
transylvanicus. Only the lower part of back has a gray color. A
133
light gray tinge also on the flanks. Whitish gray colors on cheeks and
sides of muzzle.
Measurements larger thaninL.e.transsylvanicus. Length of
skull about 100mm (92-105mm; M.99.2mm); length of hind feet 140-
167mm (M.151.4mm). Average weight 4kg (3.2-4.4kg).
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 inits
richer color and stronger ocher tone on chest and anterior sides of
forefeet. Color of summer fur intermediate between that of the Crimean
Subspecies andofL.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.
154 As mentioned above, the winter fur of L.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 of L.e. cyrensis. Specimen
No. 2-09, 1907 resembles this subspecies particularly closely.
FIGURES 79-81. Variation of nasals in Lepus 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.1. 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 transitionalto L.e.
caspicus.
There are Specimens among the 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 andL.e.
Cyrenmsis.
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 Lepus europaeus caucasicus: Length of skull
92-105mm (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 16.3-22.2mm (M.19.4mm); length of bony
palate 5-7.7mm (M.6.2mm); 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 560-680mm; hind feet 140-167mm
(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 andL.e. cyrensis (see above) occur in Dagestan on the
Bogos Range. Inthe Central Caucasus, the boundary between the 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 of L.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
Latte. cameasmews™
332. Lepus (Eulagus) europaeus cyrensis Satun. (1905).
Transcaucasian common hare
1832, Lepus timidus Ménétries E,, Catalogue Raisonné des Objets de Zoologie, p. 23 (partim!).
1876, Lepus caspicus Blanford W. T., Eastern Persia, Il, Zoology, pp. 82-83 (Teheran - Isfahan),
135
1896, Lepus europaeus var. SatuninK., Vorladufige Mitteilungen iiber die Sdugethierfauna der
Kaukasuslander, Zoolog, Jahrbiicher, Bd. 9, S. 308.
1903, Lepus sp. Satunin K, A., Obzor issledovaniya mlekopitayushchikh K avkazskogo 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 maloizvestnye 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 cyrensis Satunin, K,A., Fauna Chernomorskogo poberezh'ya (Fauna of the Black Sea Coast),
p. 41,
156 1918, Lepus cyrensis Kollmann M., Note sur les mammiféres rapportés d'Asie Mineure par M. Gadeau de
Kerville, Bull. Mus. , Paris, pp, 201-204. (Asian Minor).
1929, Lepus europaeus cyrensis OgnevS.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 Elisavetpol [Kirovabad] Province. Deposited in the Georgian Museum (former
Caucasian Museum),
MATERIAL 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.
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.I. 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.
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157
Length of skull usually less than 100mm (91-103mm; M.96.6mm);
length of hind feet 144-150mm.
ADDITIONAL DESCRIPTION. Color of summer fur of L.e.cyrensis
very Similar to that of L.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, predominatesinitscolor. 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 incaucasicus. 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 youngcyrensis similar
to that of young L.e. hybridus andL.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, 35km 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 81-86mm; zygomatic width 43.5-48mm; length of
nasals 41-47mm; width of skull at level of auditory meatus 34.2-36.3 mm;
length of row of upper molars 15.3-17.2mm.
Measurements of body small, generally like those of L.e. cyrensis:
length of body with head 490-563mm; hind feet 130-150mm; 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.
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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 hairalongthe 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 (35km 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. S. 7119, female, 24 January 1932, of Kizyl-
Agach, 35km southof 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 Hilzheimer M, , Die Hasenarten Europas, Jahresber. d. Vereins fiir
Vaterl, Naturk, in Wiirttemb., S. 388, Taf. V, Fig. 4.
1910, Lepus medius caspicus Trouessart E.L,, Faune des Mammiféres d'Europe, p. 217.
1916. Lepus europaeus caspicus Ognev S.I,, Mlekopitayushchie Tavricheskoi gubernii (Mammais
of the Taurian Province), part 1, Gryzuny (Rodents), pp. 58-59, pl. 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 Kumetsov, 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
160
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. Inspecimens 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 95.3-103.1mm (M.100mm); length of hind feet 137-
153mm (M.152.2mm). 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: 1) 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.
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,
pl. 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, intermediate between
avellaneous and pinkish cinnamon.
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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 natioturgaicus.
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 amore intense clay-rust color (cinnamon).
The clay-yellow color ofthe 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 natiokalmykorum (''Zoologischer Anzeiger'', 1929, Bd.
84, S.77). The type locality is Ikitsokhura, Akima sands, Kalmyk Region,
(E.1. Orlov). Wealso 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 inseries. 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 whitetips 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 natioturgaicus 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 examined 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, 580mm;
hind feet 150mm; ear 120, 128,135mm. Weight 3.855-4kg,
Length of skull 102.5mm; condylobasal length 90mm; zygomatic width
50mm; length of upper tooth row 18mm.
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
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162
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. natiokalmykorum occurs in the Kalmyk steppes.
It is not known how far north it occurs.
The exact boundary of the range of L.e.c. natioturgaicus is
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. natioturgaicus occurs inthe 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. natioturgaicus
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. natioturgaicus 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.
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 tolai 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 stantsiya,
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 authorconsiders 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 NAMES: '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 Glirium 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 Davuriam, 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 Gobeénsi
ubique ad Tybetum usque vivit... '').
14]
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: 1) Schreber J. Ch., Die Sdugetiere, IV, 1792, S. 878-881, Tab. CCXXXIV (color
plate of entire animal); 2) DvigubskiiI, Opyt estestvennoi istorii vsekh zhivotnykh Rossiiskoi imperii
(Natural History of the Animals of the Russian Empire), 1829, p. 43, pl. 25 (entire animal); 3) Simashko, Yu,
Russkaya fauna (Russian Fauna), II, 1851, p. 778, pl. 61, fig. 1 (color plate of entire animal); 4) Radde G.,
Reisen im Siiden von Ost-Sibirien, 1862, S, 211-215, Taf. VIII, Fig. 2a-c (black pattern of tips of ears);
5) Brehm, Tierleben, Sdugetiere, II, 1914, Taf. III, Fig. 1 (photograph of entire animal).
MATERIAL EXAMINED: 15 pelts and skulls.
DIAGNOSIS. Head relatively large. Ear when stretched (measured
from occiput) 1 1/5-1 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 lengthof 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 87-95.2mm (M.90.6mm); zygomatic width 42.3-
47.8mm (M.44.6mm); length of nasals 36-41mm (M.38mm); width of
nasals 15.3-19.3mm (M.17 mm).
Length of body with head 430-580mm (M.492mm); hind feet 110-135mm
(M.123.6mm); ear (along concavity) 85-115mm (M.94.9mm); tail (without
terminal hair) 75-110mm (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: 1) larger size; 2) greater thickness and width
(especially at the base of the nasals, whichinmost 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
A
FIGURE 84, Tolai hare, Lepus tolai tolai FIGURE 85. Tolaihare, Lepus tolai tolai
Pall. No. M.2323, 16 January 1928, Vicinity of Pall. , skull ventral
Troitskosavsk [Kyakhta], P.S. Mikhno, S. I,
Ognev's collection, skull dorsal
165 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.
FIGURE 86. Tolaihare. 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.
FIGURE 87. Mandible of Lepus 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, andsidesof 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: 1) a straw tinge mixed with gray (avellaneous); 2) a stronger rusty
144
166
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.
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 tolai Pall.
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
167
168
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.
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 witha
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 arichstraw 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 anda
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 strawears with a strong white tinge.
146
169
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 suchas L.e.cyrensis andL.e.
casSpicus. However, the following characters justify the consideration
of L. 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 of L. 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, 30km 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 480mm; tail 112mm (without
terminal hair); ear 85 mm; hind feet (without claws) 115mm. Weight
2 JOike..
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
84-88.2mm; condylobasal length 76.2-78mm; zygomatic width 40.5-
43.6mm; length of nasals 36-40.3mm; widthofnasals 17.2-18.7mm; width
of skull at level of auditory apertures 33.2-34.6mm; length of bony palate
6.2-7.3mm; width of incisive foramina 8.1-9.5mm; length of row of upper
molars 15-15.5mm.
147
170 It would thus be possible to consider the Manchurian Tolai hare as a
separate subspecies. However, I am waiting for more materialto 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:
1) 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,
25km north of Lake Koso Golis in my collection. I also consider asa
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. lehmanni) 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-Uegen, in the
lower reaches of Tuin-Gol, on the northern coast of Lake Ureg 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 of tibetanus 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. **
cy Bi 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 aretwo completely 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
L172
in river valleys (Tuin-Gol, Dzabhan Gol, Murin-Gol rivers, etc.) and lake
basins (Ureg-Nur). One can find three or four during half an hour in
these places, in thickets oftall Lycium halimifolium Mill., shrubs
of Caragana 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 hereon 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 Ureg-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 60-
80 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.
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 Wistenhase (German).
LOCAL NAMES: kuyan, or koion (Kazakh, Kirghiz); taushan (Turkmenian); khar-gush (''Donkey's
ears'' - Persian).
* [Now in the Inner Mongolian Autonomous Region. ]
149
173
PRINCIPAL FIGURES: 1) Blanford W, T., Eastern PersiaII, Zoology 1876, pp. 80-82, pl. VIII (color plate of
entire animal under the name Lepus craspedotis); 2) Blanford W.T., Scientific Results of the Second
Jarkand Mission, Mammalia, 1879, p.63, pl. IV, fig.2- Lepus tibetanus (entire animal in color);
3) Blanford W.T., l.c., 1879, pp. 67-68, pl. V, fig. 1 (entire animal in color); pl. Va, fig. 1-1b (skull) -
Lepus pamirensis; 4) Blanford W.T., l.c., 1879, pp. 68-71, pl. V, fig. 2 (entire animal in color):
pl. Va, fig. 2-2b (skull) - Lepus stoliczkanus; 5) BogdanovM.N., Ocherki prirody Khivinskogo
oazisa (A Survey of the Nature of the Khiva Oasis), 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.I, parts 3 and 4, 1904, pl. LXXXIV, LXXXV, fig. 4
(photograph of skull dorsal and ventral - Lepus tibetanus); 7) Kashkarov D.N., Zhivotnye Turkestana
(Animals of Turkestan), 1932, p, 196, fig. 140 (under the name Lepus tolai lehmanni); 8) Hepter
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 Lepus 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
with 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.
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: 1) 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: 1) bony palate of
desert hare relatively wider than that of common hare; 2) incisive
foramina narrower, sides of their posterior third usually without
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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**.
A specimen of Lepus 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, pl.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 of L.t. lehmanni also resembles that of L. 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. tibetanus 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
176
has an irregular blackish mottling.on a general light straw-yellow
background. In some subspecies, the backhas arust-red tinge ona sand-
straw colored ground (e.g., inL.t.buchariensis). The color of
i. t apamiuar eins hs jiand Eitri gaiais an ic wsyils moreintense: ;Inythe
latter, the color of the back is rather 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.
Ya eat pay er aa S45 fy i)
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 andcreticus,
as subspecies of Lepus europaeus, but also subspecies of true
desert hares suchas L. judeae, tibetanus, lehmanni and tolai.
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 Asianhares closely resembles that of Lepus 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 Asianhares resemble
the Americanhares 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: 1) 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.creticus
(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 Sduger. 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|
hw
a
Wee se a
FIGURES 94-98. Postures of the desert hare, 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 of L. 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. alleni 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)coronoidandcondyloid 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: 1) length of bony
palate; 2) narrow mesopterygoid fossa; 3) markedly inflated and rounded
bullae osseae.
The pale color of L. 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 Asianhares, 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 onmany 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 Gunther
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
(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
tehmeanat seve, 12. t. amine nisi s Blant. Tiaieaoine hariens is
Og, ist. steoliezkanus) Blant.) Ey tibetanus \sub sip. ? (from
Kaper-Wach)wlit.icraspedotis Blant., .t. zanksanileus Satun. ,
hee, decserprorum, Orcn., J. bt. Cemtmhaslatieug satun. 4 au. @.
quercerus Hollister, L.t. przewalskii Satun., L.t. gobicus
Satan... if. 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: 1) 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 closelyrelatedtoL. pallipes
but differs from it in some external characters, suchas 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 Lepus 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 of L. 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 tailis 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 Lepus centrasiaticus (a subspecies oftibetanus)
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
181
* Owing to lack of material, I cannot decide whether Lepus filchneri Matschie, L. aurigineus
Hollist, andL. swinhoei Thos. belong to this group. Gl. Allen (1927) places L. tolai into this
group as a subspecies,
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183
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.
Li teem tr a's iat hess! presumably-occurs! indherfarmnorths) Itaomms
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
of Lepus 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: )) 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, 130-140mm 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.
The skull of L.t. kozlovi 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 centrasiaticus. However, winter specimens of
gansuicus (No. 2840, 8283, 2839, etc.) do not differ from
centrasiaticus.
Finally, it should be mentioned that Lepus kaschgaricus described
by K.A. Satunin is not monotypical. The specimen of kaschgaricus
from the Yarkand Oasis first mentioned in K.A. Satunin's work certainly
belongs to jarkandensis. 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
184
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 reachesthe 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 seena
very small, long-legged, light coloredhare inthe 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 jarkandensis. 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 thatthe 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 Ili, 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 Ili river near the village Ili.
It occurs in the Zaisan Depression from where K.A. Satunin described
anew 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 Lepus lehmanni Sev.
160
185
to V.N. Shnitnikov (1932) it is distributed throughout Dzhetysu [Semireche]
at an altitude of 300-3000m. 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 Salsola.
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 Nitraria, 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.
eee
* [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 cemeteries, 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 harmfulto 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 corsac, desert foxes,
wolves, polecats, and various cats, and also by the following birds of
prey: goldeneagle, steppe eagle. Younghares 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 aralensis 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 OgnevS.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 OgnevS.I, Zur Systematik der russischen Hasen, Zoologischer
Anzeiger, Bd,, 84, H. 1/4. S. 78.
TYPE LOCALITY 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 medium 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 gravy 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 83-88.2mm (M.85.6mm); zygomatic width 38.5-
42.8mm (M.41mm); length of nasals 34.2-38mm (M.36.6mm); width
of nasals 14-16 mm (M.15.7mm).
Length of body with head 387-485 mm (M.455.9mm); hind feet 108-
126mm; tail 75-107mm (M.97.8mm); ears (measured along concavity)
87-100mm, ear (measured from occiput) 100-126mm (M.119.8mm).
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 andL. 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. Ihad 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
differ 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
of Ak-Kuyu, 60 kmnorth of Dzhebel (Transcaspia). Hares from the Kopet-
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
Ly. tibetanus, ang not fo Lk. en rope us.
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
obtainedinearly 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 tiber einige russische Saduger, Folia Zoologica
et Hydrobiologica, Bd, VI, 1934, S. 21-23.
+* S.1, Ognev and W.G. Heptner, ''Mlekopitayushchie srednego Kopet-Daga i prilezhashchei ravniny"
(Mammals of Central Kopet-Dagh and the Adjacent Plain). —Trudy Nauchno-issledovatel'skogo instituta
zoologii, III (1): 71-73. 1929,
164
190
summer fur of the desert hares (e.g., L. t. lehmanni) 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 andL.t.lehmanni.
The taxonomic position of L.t.turcomanus is doubtful. Its color
differs little from that of L.t. lehmanni. Considering L.t.
turcomanus as asynonym of L.t.lehmanni is a simple solution
to this question. There is, however, a difficulty — the skullof 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
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. lehmanni 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.
334', Lepus (Eulagus) tibetanus lehmanni natio zaiSsanicus
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 zaisanicus OgnevS.I., Materialy po sistematike russkikh zaitsev (Materials on
Taxonomy of Russian Hares), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, Vol. XXIII, pp, 483-484,
TYPE LOCALITY 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: 1) 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 of L.t. lehmanni natio zaisanicus more whitish
than that of L.t. lehmanni.
Length of skull 83.8-90mm; zygomatic width 40-43.7mm; length of
nasals 34.1-41.7mm; width of nasals 15-20.3mm.
Length of body and head 443-490mm; hind feet 120-128mm; ear
(measured along concavity) 83-113mm; tail 88-116mm.
SYSTEMATIC NOTES. As this hare is closely related to L.t.
lehmanni, I intendedto make zaisanicus a synonym of lehmanni.
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 anatiooflehmanni.
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 LOCALITY 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
192
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 ofthe hair; tips ofhair 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 84.9-87.2mm; zygomatic width 40.2-44mm; 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 14.3mm.
Measurements of specimens examined by us for comparison with those of
Hollister:
No, 829, Feb, 1936, No. 57 o”, 30 June 1936,
Chegan-Burgazy Near Kosh- Agach
Length (in mm. )
o'ad,, 2 July 1912
According to Hollister
Body
Tail
Hind foot
Ear
Weight (in grams)
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 84mm. It has a pale gray-straw-yellow color onthe 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 witha
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
193
Color of young (body length 250mm, weight 150 grams) as follows: back
pale straw-gray-yellow 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 basesof 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 zaisanicus). 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 Lepus 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 tibetanus) 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 hare
1928, Lepus tolai craspedotis OgnevS,I. and Heptner W.G,, Zoologischer Anzeiger, LXXV, S.
261; Ognev S.1. 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
instituta zoologii, III (1): 71-73. 1929.
168
194
195
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,
DIAGNOSIS. General color of back of summer fur straw-gray witha
well marked dark mottling 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 84-86mm; zygomatic width 38-42mm; length of nasals
36-37mm; width of nasals 15-16mm.
Length of body and head 435-460mm; _ hind feet 121mm; ear (measured
along concavity) 108-119mm; ears (measured from occiput) 139-149mm;
tail 75- 84mm
SYSTEMATIC NOTES. The Kopet-Dagh hares resemble L.t.
lehmanni closely in their color. They differ from it as follows: 1)
grayer tone onthe 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 inthe small material examined. Rows of molars especially
small and weakly developed. Length of upper row of molars 14-14.9mm
invoursspecimens,) inva. t) Vehm anna 15-16.9 mim.
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
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 Germabthey 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 severo-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 Sdugethiere Transcasp. , Tiflis,
pp. 62-64.
1894, Lepus lehmanni Varentsov P., Fauna Zakaspiiskoi oblasti (Fauna of Transcaspia), p. 16,17.
1918. Lepus lehmanni Bil'kevich S.I., Kollektsiya Zakaspiiskogo muzeya (The Collection of the
Transcaspian Museum), Izvestiya Zakaspiiskogo muzeya, I, p. 9.
1928. Lepus tolai desertorum OgnevS.I. und Heptner W.G., Einige Mitteilungen tiber die
Sdugethiere des mitt], Kopet-Dagh etc., Zoologisch. Anzeiger, LXXV, S. 262, Ognev S.I. and Heptner
W.G., Mlekopitayushchie srednego Kopet-Daga i prilezhashchei ravniny (Mammals of the Central
Kopet-Dagh and the Adjacent Plain), Trudy Nauchno-Issledovatel'skogo instituta zoologii, Ill(1):73-75.
1934, Lepus europaeus desertorum Heptner W.G., Systemat. und tiergeogr. Notizen uber Sdugethiere
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.1, 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 76.6-84mm; zygomatic width 37-40.7mm,; length of
nasals 31.8-37.5mm; width of nasals 15.6-18.5mm.
Length of body and head 399-460mm (M.431.3mm); hind paws 102-
120mm (M.118mm); ear (measured along concavity) 98.2-105mm; ear
(measured from occiput) about 120mm or more; tail 79-100mm.
These measurements show that the skullof L.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 Lepus FIGURE 103. Skull of Lepus FIGURE 104. Skull of Lepus
tibetanus desertorum Ogn. tibetanus buchariensis tibetanus subsp., No. M.
M. 3066, from Annau (near Ogn. M. 3660, 22 May 1914, 3065, Sulyukli, Kopet-Dagh,
Ashkhabad), holotype, S.I. Ognev's Khatyn-Rabad, N. A. Pobrinskii, S.I. Ognev's collection
collection S.I. Ognev's collection
197 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 anda
characteristic variegation. Rust tones on occiput and chest well marked.
Color of occiput intermediate bétween 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 inits 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 35km east of Gasan-Kuli, and near
Karabat.
The distribution of L.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
197
198
FIGURE 105. Skull of Lepus tibetanus desertorum Ogn. No. M. 3066,
Annau, near Ashkhabad
FIGURE 106. Skull of Lepus tibetanus buchariensis Ogn. M, 3060, holotype
338. Lepus (Eulagus) tibetanus buchariensis Ogn. (1922).
Buchara desert hare
1823, Lepus tolai Lichtenstein H., Naturg. Anhang m Eversmann's Reise nach Orenburg u. Buchara,
Se Lug,
1922, Lepus tolai buchariensis, OgnevS.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, OgnevS.I., Nekotorye dannye po sistematike russkikh zaitsev
(Data on the Taxonomy of Russian Hares), Biologicheskie izvestiya, Vol.I, 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: 11 specimens.
DIAGNOSIS. Differs from L.t.desertorum inthe 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 79-88.3mm (M.84mm); zygomatic width 38.6-42.2 mm
(M.40.5mm); length of nasals 32.2-41.7mm (M.36.6mm); width of nasals
13.2-16 mm (M.14.5mm).
Length of body and head 460-530 mm (M.485 mm); hind feet 102-119 mm
(M.112.7mm); ears (measured from occiput 95-117mm (M. 123mm) [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
199
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 of L.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 inthe following characters: 1) paler
sand-straw-rust color of the back which is more grayishinL.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; inL.t.lehmanni, the legs are pale gray; 4) color of ears in
L.t.buchariensis paler yellow thaninL.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-Rabad (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
200
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, Lepus 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 ofthe desert hare in
that its summer fur is very soft anddelicate andits 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. 8.5004, 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 83-89mm; zygomatic width 40-41.4mm; length of
nasals 34.3-39.8mm; width of nasals 14.2-16.3mm.
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).
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, ithas 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 Lepus kaschgaricus Satun. from
the Bol'shoi Yuldus and Tien-Shan (obtained by M.N. Przheval'skii) are
closely related to the Pamir hare.
GEOGRAPHICAL DISTRIBUTION. According to N.A. Severtsov, this
form is common onthe Pamir. In G.E. Grum-Grzhimailo's opinion (1886)
it is very common inthis region. M.P. Rozanov (1935) writes that the
Pamir expedition found large numbers of L.t. pamirensis inthe
Markansu area, beyond the Kyzyl-Art Pass in the Trans-Alai.
Subgenus Lepus s. str. Linnaeus (1758)
201 1904, Lepus (subgenus!) Lyon M., The Hares and their Allies, Smithson, Miscell. Collections,
Vol.1I, 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, 11/10 —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 11/8 to 11/4 as long as the bony palate.
Anterior lower premolar slightly inclined backwards.
Geographical distribution. Boreal and arctic regions of Eurasia and
North America ft.
* In arctic forms, tail completely white both in summer and winter,
** With the exception of Lepus 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 Poecilolagus 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 liévre 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'yava (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); teus' (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-Gilyak of the western Sakhalin Coast); ossik, or kanak (white) {Nivkhi-Gilyak 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, pl.47, 1770; 2) Schreberl.,
Die Sdugethiere, Erlangen, Bd, IV, 1792, S. 885-891, Taf. CCXXXV A (summer fur); Taf. CCKXXV B
202 (winter fur); 3) Bingley W., Memoirs of British Quadrupeds, 1809, p. 308 (color plate}; 4) Nilsson Sv. ,
Illuminerade Figurer till Skandinav. Fauna, 1831-1832, Vol.I, H.6, pl.22(Lepus borealis varietas -
entire animal in color), H.6-7, pl. 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; pl. 60A, figs. 2-3 (summer fur in color); pl. 61, fig. 2 (winter fur in color); pl. 61, fig. 3 (skull):
pl. 62, fig. 3 (winter furin color), Lepus variabilis incolor: pl. 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 1'Académ. Imp. de St. -Petersb. , IX, No. 14, 1851, pp. 25-29
(drawing of skull, p. 29); 7) Blasius J., Naturgeschichte d. Sdugethiere Deutsch]., 1857, S. 420-425,
Fig. 229 (structure of teeth and bony palate); 8) Meves W., Holmgren A. E,, Atlas ofver Scandinaviens
Diaggdjur, Stockholm, 1873, pl. XV, Fig. 1-7 (Lepus timidus L,-canescens et borealis in color);
9) Brehm A., Zhizn' zhivotnykh. II (Animal Life, Vol. II), 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, Mammiféres, 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.
Il, (Animal Life, Vol. II), p. 664 (half-tone plate of entire animal); 15) Lydekker R., A Hand-Book of the
Brit. Mammalia, London 1895, pl. XXVIII {entire animal in color); 16) Kashchenko N.F., Rezul'taty
Altaiskoi ekspeditsii (Results of the Altai Expedition), Tomsk, 1899, pp,57-59, pl. 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.1I, pp. 3-4, 1903 (1904). pl. LXXX and
LXXXI, fig.3 L. tschuktschorum (skull dorsal and ventral); fig.4.L. variabilis - (skull); 19)
Gake, V., Zhivotnyi mir (The Animal World), Vol.1, 1901, pl. 29 (color plate); 20) Millais J.G., The
Mammals of Great Britain, Vol. III, 1906, pl. S0(color), pl. 51 (color), pl. p, 32 (biological scene), pl.
p. 34; 21) Hilzheimer M., Die Hasenarten Europas, Jahreshefte d, Vereins fiir vaterd. Naturk. im Wiirtt. ,
1908, S, 385-387, Taf. V., Fig. 8a-c (skull, mandible, teeth); 22) MeerwarthH. 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. scoticus (skull
lateral, ventral and dorsal); p.526, fig, 104 (teeth); 24) Barrett-Hamilton G., A History of British
Mammals, 1912, part. XI, pl. XV (entire animal); p. 230, fig. 2 (skull lateral); p. 232, fig. 3 (palate);
part XII, pl. XIX fig. 8 (hind leg); pl. XX, fig, 8 (hind leg); 25) Ognev S.I. , Mlekopitayushchie Moskovskoi
gubernii (Mammals of the Moscow Province), 1913, pp. 298-307, pl. IIl, pp. 45-50 (seasonal variations of
fur); fig. 53 (skull of young blue hare); pl. 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
arkhiv anatomii, gistologii i embriologii, 1 (2), 1916, pl. XVI, fig. 13 (skull dorsal, ventral); 28) Thorburn
176
20°
204
A. , British Mammals, Vol, II, 1921, pl. 34 (in autumn fur); pl. 35 (in winter fur); 29) Kishida K., An
Explanatory Illustration of Japanese Mammals, ed.II, 1926, pl. 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(Lepus timidus saghaliensis), pl.I (Skulls of Lepus ainu); pl. II
(skull of L. 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,
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 I.I., 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, pl. I-II (photograph of skulls of L, t.
begitschevi andL.t.sibiricorum); 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, 11/10 to 11/8 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 forms sharply different from summer fur (except for
Lepus hibernicus Pall.). Hares of the subgenus Lepus 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.
Meatsyqp.d-beca. ira yp total:s:
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.
Bullae osseae small and moderately inflated, slightly protruding
anteriorly in profile.
Wh
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
206
Length of ear (measured from occiput) 110-118mm,; length of ear (along
concavity 77-115mm; hind feet 132-190mm.
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 ofthe 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 ofthe orbit than does that ofthe 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 betweenthe 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 anexample. 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, pl. 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 ofthe same color. There are three types of guardhairs: 1) 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.
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207
ey
FIGURE 110. Mandible of Lepus timidus kozhevnikovi Ognev. M. 3167,
17 November 1932, Shipov Experimental Forestry near Pavlovsk, S.1. 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.
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 brownroots, 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 contrastto the variable summer color, the winter color is constant,
pure white. The following variations occur, however: 1) 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
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208
persist in such specimens, giving the animals a peculiar smoky, grayish
color*.
FIGURE 111. Blue hare in winter fur.
Drawing by A.N. Komarov
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 of L. hibernicus resembles that
of the blue hare. The only differences are as follows: 1) 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, Ryazan Province (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 (Alopex lagopus beringensis).
181
Nine forms of Lepus timidus are known inthe 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 American 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: 1) 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,. areticus BOse
As was recently shown by Magnus Degerbgl and F. Wimpffen Braestrup
(''Vidensk. Medd. fra Dansk. naturh. Foren", Bd. 98, 1934, pp. 197-205),
the incisors of L. 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 skullof 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 onlya
Subspecies of L. timidus.
According to A. Howell (1936), there are seven subspecies of Lepus
arcticus: 1) 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.
bangsi 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 totschuktschorum. Their skulls are very similar both in
general and in measurements. The only difference between them is the
209
* Skulls of Greenland hares also belong to the ''blue hare type".
1254 182
210
color of the summer fur. Lepus paodromus Merriam is smaller,
and its Summer 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) Lepus 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., 71°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 (1867), 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 numbers 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 possiblythe 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
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212
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 2-12km 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 SiberianIslands. According to
fur traders, it was caught once or twice on the Bol'shoi Lyakhovskii Island.
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 Penzhina and 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.
Inthe 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
Heilungkiang 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 Hébségél 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 [Ayaguz].
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. ]
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213
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 from 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 inthe forests of the Mozyr Polesye (A.G. Fedyushin, 1927)
and farther south, in the forests of the Ovruch District (M. Baklashov,
137%). 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 untilthe present. For instance, ablue hare was foundnearSumami. 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), blueharesrarely occur in the former Zenkov, Poltava and
Konstantinograd [Krasnograd] counties ofthe Poltava Province. Inhis 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.
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214
215
Except for these fragmentary records, we have no data on rare and
accidental captures in the southern and central Ukraine. We repeat that
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 mandibles) of blue hares were found in Karst caverns
which were partly filled with sand. In S.K. Dal's opinion, the presence
of the blue hare inthe Crimea was short lived. It apparently entered the Crimea
during the Wurm glacial, when the ice reached 48-50°N, and the land
bridge connecting the Crimea with the Continent was 200km 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. Dal'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. Inthe centralbelt 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
Buzuluk 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 in
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)).
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216
ZiT
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 (Betula rotundifolia) and willows (Salix 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 Pinus 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 inthe
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
ofdewberry, 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, inthe 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 inthe high, precipitous, eroded
cliffs commonin some parts ofthe 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 Linné. 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 ataciturn 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. Itis 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
188
female run to help the young in response to this sound. There is evena
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 (Asio 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 which 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 ina 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. I would
also consider as cunning the rare cases in which the hare lies ina
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 oran 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
218
* Double and triple steps mean a double or triple repetition of tracks by returning along the line of the track.
189
before restingis 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."
FIGURE 113. Tracks of slow walk of a foraging blue hare
Photograph by S.1I. Ognev
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 ina somersault, as if shot. The hare
climbs precipitous mountains rapidly and safely owing to the build of its legs.
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221
Even when the animal could escape by running over an even space, it may
choose to climb a mountain, rapidly ascending the steep Slope.
FIGURE 114. Track of a hurrying 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.
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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 longtime. In summer they were often seen walking at 6.00a.m.
On several occasions, Stakhrovskii saw hares in the early morning hours
playing with eachother, rollingin 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.
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2
FIGURE 116. Blue hare in winter
Photograph by S.I. Ognev
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 8. 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 3.00a.m.). During quiet, moonlit, winter nights,
they feed and leave footprints even after midnight.
NESTS AND BURROWS. Inthe 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 (in litt., 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 80-100cm deep, for
instance, near the estuary of the Anadyr river (23 November 1930).
Blue hares in northeastern Siberia occasionally build permanent
communal burrows on hills and steep river banks. Such a colony was
described by A. F. Girikova (in litt., 1935).
"I 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). Wesawvery few hares in the forest belt. They appeared
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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 1/2km).
The cliffs here reached a height of 20-30m. 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, whichhad 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 1-2m 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 ona 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 ona small platform above the
exit, which was 16 cm high and 39 cm wide.
"The air vents of another burrow were located alonga crack inthe ground.
The largest burrow (25X35cm) 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 7m. 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 stillthere. 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,
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225
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, especiallynear 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 foodwas 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 Melica nutans, Calamagrostis, Greekvalerian,
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 andnettle. The first two are especially favored by the hares, who
often have to digthe plants out of the snow. They usually bite off the parts
protruding from the snow.
REPRODUCTION. Inthe 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 2-4litters duringthe summer. D.I. Aspisov (1936)
thinks that blue hares normally have 3 litterseachsummer. 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 females have two litters a year, producing 8 young. The natural
death rate among the young reaches 35-40 per cent. The progeny ofa
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.
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 ''autumnhares'' bythe 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
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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 withtheir 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 2-3m 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 dothe 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. Onthe 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 immediately 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 1/9
days of pregnancy. We sawa 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 freeingthe youngone
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 withher 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 ofa 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
ofthe second litter increased by 88.3 %as comparedto 65.8% in a specimen of
the first litter.
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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.I, 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. Lonnberg (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 bythe appearance ofanewundercoat. 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
©. von Loewis, Bemerkungen iiber den Schneehasen Livlands, Zoologisch. Garten, 1877, S. 16-20.
** Brehm's Tierleben, Sdugetiere, Bd. Il, 1914, S. 68,
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229
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ee
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230
(aostdsy ‘[‘q Jeqe) arey anj{q ay Jo yfow Butids Jo ureiZeIq ‘“gIT FUND
ey
fey
ae
Sy
200
229
230
231
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.
"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, andhead, behindthe 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 onthe 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
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
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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.
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 Lepus 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
ofsunspots. 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 onhaystacks, 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
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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 1.5cm thick were eaten completely.
No guns were used. Hares were caught with nets. Groups of 25 people
covered distances of 200m 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, 25times as large inKarelia, 20timesas large inthe 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. InA.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: 1) 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
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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 and1930. 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 Westernregions. 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: 1) 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 Synthetocaulus
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 by Linguatula taenioides
(Pentastomida).
According to M.P. Lyubimov (1935), about half the hares suffer from
coccidiosis. Eimeria stiedae infects the liver and Eimeria
perforans andE.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
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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 1.1. 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 ofhares are animportant 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.
FIGURE 119, Blue hare in beginning of spring molt
Photograph by S.I. Ognev
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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 inner 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 50mm.
237 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 width 48-50. 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 thaninL. 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 86-93.2mm; zygomatic width 45-49mm. Length of
body 440-515 mm; hind feet 132-152mm. :
L.t. transbaikalicus Ognev.
16 (15). Measurements large: length of skull 88-96mm; zygomatic width 47-51mm. Length of
body 523-570mm; hind feet 155-180mm.
L.t. mordeni Goodwin.
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239
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 (NewName for the Alpine hare, Thomas Pennant,
British Zoology, IV, 1770, p. 40).
1778. Lepus borealis Pallas P.S., op. cit., p. 2.
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, editioI, app.1I, pp. XLV-XLVI;
editio II, Vol. II, app.IV, pp. 17, 170 (Nomen preoccupatum!). Bechstein J. M., Gemeinniitzige
Naturgesch, Deutschlands etc,, 1801, editioI, Vol. II, S. 1096 (Lepus timidus albus from
Thiiringia).
1820. Lepus borealis NilssonS., Skandinavisk Fauna, Lund, B.1, p. 211,
1829-1832, Lepus borealis collinus Nilsson S., Illuminerade Figurer till Skandinav. Fauna, Lund,
I, pl. 16, 19.
1829-1832, Lepus borealis sylvaticus Nilsson S., op. cit., I, pl. 22.
1844, Lepus canescens Nilsson S., Ofversigt af Kongl. Vetensk, Academ, Férhandl., Stockholm, I
p. 133.
1884, Lepus variabilis Pleske Th. , Ubersicht der Sdugethiere 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 Naturae" 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 48-50.8mm
(M.49mm).
Length of body 590-600mm; 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: 1) 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.
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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 Lepus 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 Lepus
t.timidus and L. lugubris Kastsch: 1) 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 natiolugubris.
G.E. Barrett-Hamilton (Proceedings Zoolog. Society of London,
part I, p.90, 1900) described the subspecies L.t.altaicus 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 andL.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.lugubris from that of the nominate subspecies. The
ranges of L.t.timidus and L.t.lugubris possibly overlap ina
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 inthis 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 timidus Satunin K, A., Pozvonochnye Moskovskoi gubernii. Mlekopitayushchie (Vertebrates
of the Moscow Province. Mammals), Izvestiya Obshchestva Lyubitelei Estestvoznaniya, antropologii i
etnografii, Vol. XXXVI, p. 14.
* N.F, Kashchenko, Results of the Altai Expedition, Tomsk, 1899, pp. 57-59, tab. Il, fig. 4.
208
240
242
1913, Lepus timidus Ognev S.I., Mlekopitayushchie Moskovskoi gubernii (Mammals of the Moscow
Province), Vol.1, pp. 298-307.
1929, Lepus timidus kozhevnikovi OgnevS.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 Chernaya,
Bogorodsk County, Moscow Province, collection of the Moscow Zoological Museum.
MATERIAL EXAMINED: 89 pelts and 125 skulls.
DIAGNOSIS. Insummer 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 35mm; hairon lower part of chest and sides of belly
up to 60mm.
Length of skull (Figures 120, 121) 97.8-105.6mm (M.100mm); zygomatic
width 50-53 mm (M.52.3mm).
Length of body 450-630mm; hind feet 155-180mm (M.170.8mm); ears
(measured along concavity) 80-100 mm; ears (measured from occiput)
110-118mm (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 of L.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 8.V. Kirikov
includeda female (shot on 20 May 1931 intugais along the river Belaya) which
had the anterior inner margin of the ears markedly black.
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FIGURES 120-125. Skull of subspecies of Lepus 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 natiolugubris 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. transbaikalicus 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
243
GEOGRAPHICAL DISTRIBUTION. The southern and southwestern
boundaries of the range ofthis 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 belongto L.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 toL.t.
sibiricorum*thanto L.t. kozhevnikovi. Hares from the southern
Urals and Bashkiria belong to the central Russian form.
342. Lepus timidus sibiricorum Johans. (1923). West Siberian
blue hare
1877. Lepus variabilis Peters W., Ubersicht iiber die wahrend der Sibirisch, Expedition von Dr.
Finsch gesamm. Sdugethiere, Monatsberichte der K6nigl. Akadem,. der Wissensch, z, Berlin, S, 735
(Samarovo on the Irtysh).
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 (1):25-38.
1923. Lepus timidus natio sibiricorum Johansen G, E,, Po Chulymu (Along the Chulym),
Izvestiya Tomskogo gosudarstvennogo universiteta, p. p. 58-59,
1923. Lepus timidus Martino V. and E,, Materialy po sistematicheskomu i geograficheskomu
Tasprostraneniyu mlekopitayuschikh Kirgizskoi stepi (Material on the Systematic and Geographical
Distribution of Mammals in the Kirghizian Steppe), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk,
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 Shukhov1., 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 toL. t. timidus.
PAM
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 46.7-53.2mm (M.49.7mm).
Length of body 433-550mm; hind feet 145-155mm (M.152.1mm); ear
(measured along concavity) 86-92mm; 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: 1) 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.
GEOGRAPHICAL 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), Lepus 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 andL.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 transbaikalicus Ognev (1929).
Transbaikal blue hare
1910, Lepus timidus Kashtchenko N, F., O kollektsii mlekopitayushchikh iz Zabaikal'ya (A Collection
of Mammals from Transbaikalia), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, XV, p. 271.
1926, Lepus timidus Soronov 1,, Fauna gryzunov v raione lvanovo-Arakhleiskikh khutorov (Rodent
Fauna ih the Region of the lvanovo-Arakhleiskie Hamlets), in ''Regional Studies in Transbaikalia"
(K kraevedcheskim rabotam 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
Trade), inG.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 transbaikalicus OgnevS,I., 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 42mm long; hair of flanks and belly up to 62mm
long.
Length of skull (Figures 123, 129) 86-93.2mm (M.90.7mm); zygomatic
width 45-49 mm (M. 46.7mm).
Length of body 440-515mm; hind feet 132-152mm; ear (measured
along concavity 77-92mm.
SYSTEMATIC NOTES. The above diagnosis shows that the color of
summer fur of L.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. 8.2739, 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. kozhevnikovi from the Moscow Region. Their
color is much grayer than that of the latter. The general color of the back
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246
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214
1254
945 of young Transbaikal hares is rather dull with a straw-brown tinge and
slight straw-white mottling (color close to hair-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
247 Haimin Kungssii Forest Concession inthe upper reaches ofthe 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 gichiganus OgnevS.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 47-50mm (M.49mm) (49 mm in holotype).
Length of body 550-624mm (582mm average for 20 males, 577mm
average for 17 females); hind feet 146-173 mm (M. 164mm) average
for 20 males, and 162mm 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 88-89mm (94mm average for
males and 91mm for females),
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248
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 inthe vicinity of Penzhino
(collectedby V.S. Bazhanov). A comparison between these small
collections and specimens fromthe 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-coloredL.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 1.6-3.0kg* (2.560 kg average).
According to A.I. Cherskii (in litt.) the weight of hares from Maritime
Territory is 2.6-3.4kg, 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: 1) 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.
Lengthofskull94-98mm; condylobasal length 86.2-88.9mm; zygomatic
width 48.2-49.9mm; length of nasals (along oblique line) 37.3-39.8 mm;
length of upper row of molars 18.6-19.2mm.
In my opinion, the Kamchatka blue hares can be considered as belonging
to L.t. gichiganus. Inthe ZMAN collection are some blue hares from
the upper reaches ofthe Imanriver onthe 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, pl.
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,
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249
250
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 of L.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 inthis region. Transitional forms toa
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 of L.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 intheir 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 ofthe 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 gichiganus OgnevS.I., Materialy po sistematike russkikh zaitsev (Material on
the Systematics of Russian Hares), Ezhegodnik Zoologcheskogo muzeya Akademii Nauk, Vol. XXIII,
pp. 491-496,
1927, Lepus timidus gichiganus Gassovski G.N., Gilyui-Ol'doiskii okhotnich'e-khozyaistvennyi
taion (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 silly 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).
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.
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201
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 45mm long, that of flanks up to 65mm long.
Length of skull 88-96 mm (M.93.2mm); zygomatic width 47-51mm
(M.48.7mm).
Length of body 523-570 mm (M.540.8mm); hind feet 155-161 mm*
(M.156.8mm); ears (as measured along concavity) 95-97mm (M.95.6mm).
SYSTEMATIC NOTES. AsIhad 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 3mm and of L.t. gichiganus 2.5mm. We obtained the following
measurements in our material: width of lower incisor of L.t. mordeni
3: 3. Bete si St 24372. "3. 4am (M. 3.14 mm)? of Lt “giechiganws 2 8;
2.9: 3.3mm (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 5mm against 4-4.5mm 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.4.2mm). According to Goodwin,
the width of the largest upper molar of L. t. mordeni is 6.8mm and
5-6mm (M.5.7mm) in L.t. gichiganus. I obtained the following
measurements from our material: inL.t.mordeni 5.7; 6; 6; 6; 6; 6.3;
6.5mm (M.6mm); inL.t. gichiganus 6; 6.1; 6.2; 6.2mm (M.6.1mm).
These data show that the differences of the teeth of the Gizhiga and
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 180 mm long in the holotype of L. t. mordeni.
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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 toL.t.
mordeni.
It is difficult to say how far north this subspecies is distributed. Its
range seems to pass into that of L.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 of L. t. mordeni is not known.
A.de Sowerby (1928) 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, and27km east of the
Yevseyevka Station of the Ussuri Railway. This moves the southern
boundary south by a whole degree. Inthe 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). Sakhalin 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 timidus Sowerby A., The Naturalist in Manchuria, II-III, pp. 173-174 (SakhalinIsland),
1928. Lepus timidus orii Kuroda N., The Mammal Fauna of Sakhalin, — Joummal of Mammalogy,
Vol. 9, No. 3, pp. 223-224,
1931. Lepus timidus saghaliensis AbeI., A Synopsis of the Leporine Mammals of Japan, Journ. of
Science of Hiroshima University, ser. B, div.I, pp. 49-50.
TYPE LOCALITY AND DEPOSITION, Sakhalin Island, Naihoro(western coast of central part of Japanese
Sakhalin), No. 1234, male ad., 6 May 1926, Nagamichi Kuroda's collection.
MATERIAL EXAMINED: 4 specimens.
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
253
L.t.gichiganus as follows: 1) skull large, tail and hind feet shorter;
2) ears of L. t. orii longer than those of L.t. gichiganus; 3)in
summer fur, color of forehead and sides of head pale rust-brown and
reddish, (gray-browninL.t. gichiganus*); color of back, flanks and
haunches grayish brown or reddish brown (dark gray in gichiganus).
Lower part of back of L. t. orii 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 thaninL.t. gichiganus;
3) hind feet longer, 180mm long. This last character differs from the
measurement given by N. Kuroda.
On Hokkaido (Yezo) L. timidus 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 (asin L.hibernicus);
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 Lepus timidus orii Kuroda (in mm)
wo) 7)
Laon
| 3 = 2
sg a 2 a a 2 as)
5 3 et. 4 s a a 9 a
g 2 B Bola pod 3 Bg
“ =]
£3 = % 6 we) a Oo 20) T6cs re)
5 < 5 _ Song 2 2 a3 re)
= % 5 8 e184 Bs | 88 &
q oo - U0 Ss 3 uU ra] Ss -“
5 a me GI Ra Ah = o 2 ae )
Zz ps N = fe I = nu 2 ale:
aad
ay Basal length
+
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 of L. t. saghaliensis Abe (male No. 28)
ei
4 ow 2
=] re)
3 Z ea Ce ta Reta Pe 4
e Es 6 2) a 18 2 por || ze TA A =
et as} 3 ~ a |4 +5) _ a2) a a ‘ot a
. q ~~] 4 » Cal fa t
° iS I 5] “16 ms a (3) oO ia] ca {o)
iy KA) wi iS} a. co Sia 8] 6 a
im <5} ro) | a, oO =)
¥ o |B x als a SS hte! 3 o Slo Oia 3
- ee Yet Lb) Yt Of oH SS Met
ow] H 3 ao | 8 = 9
ele Ele ale lelecedlelezlatiiletlacese
v 4 : ; : rE i
dO = co) s 93/3 wo) he sis | | se & wo 8) | & S| t 8] t% S| to D0
=) co |S a]. a|s GU Siaq|/ts a 3 | a AoOl\e SO] s q
o a = Se Page| ees 2 BIS S| 2 Sho ® 6 99/3 slo a] 9 ct}
4 (a2) = lS ale «a = cS) Chal aya el ese a 4
Pe fos e[= oho] fo] asfas [ssf] fo
The measurements of specimen in the ZMAN collection are as follows
(in mm):
Lengthvofyslaril tr. auf ;. ate uccaie wo «ated ons bh oho one qtiedeTstediee es 94,3
Zyeomati Cawa deh arus vciis ciel false tales ys ceilem-viuer cases bhG\a G08 46.8
Width of skull behind orbit............ Aan oon oad 6 6 15.2
Width lof masals at the, bases. i) 5). 2)! is of 2 wile! als) ofislbeyie orl ele Za
Length of upper tooth row.......... 85 allel 35 Hie 6 tsalo h Wey
Length of mandible................... Dodd one ooo} 65.8
Height of mandible......... Pe Ao Sk Aa ia a Ri Hd 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 toL.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 toL.t.mordeni
347. Lepus timidus kolymensis Ogn. (1922). Kolyma blue hare
1884, Lepus variabilis Bunge A., Naturhist. Beobachtung. im Lena-Delta. Mém. Biolog. de
1'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 AllenG,I. Proceedings New Engl. Zoological Club, V, p. 66 (Bol'shoi
Baranovyi).
1922, Lepus timidus kolymensis Ognev S.1., Nekotorye dannye po sistematike russkikh zaitsev
(Data on the Systematics of RussianHares), Biologicheskie Izvestiya, Vol.1, p. 106.
1926. Lepus timidus kolymensis Ognev S.I., Mlekopitayushchie severo-vostochnoi Sibiri (Mammals
of Northeastern Siberia), pp. 72-77,
TYPE LOCALITY 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 47.7-51mm
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. Itisa
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 Corner of the Taimyr Peninsula),
Materialy po izucheniyu Sibiri, Vol. IV, pp. 15-16.
1936, Lepus timidus begitschevi 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 I, I. 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 onthe back, especially in the lower parts.
Ears pale gray with a weak rust tinge and a few white hairs.
Length of skull 98-108mm (M.103.4mm); condylobasal length 88-95,2 mm
(M.92.6mm); zygomatic width 50.6-54mm (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 575-740mm; hindfeet 143-171mm; 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 I.I. 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 of body 470-550 mm; hind feet 132-157mm (M. 141.6 mm); ear
(measured along concavity) 82-93mm (M.87.7mm).
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 inthe 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 resembles
that of specimens from the Kolyma river. F.F. Shillinger caught a
257
* 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 (25 Jul. 1925). 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
rétained 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.O, Yakovlev, ZMAN
collection
Drawing by Yu. A. Kostylev
New material will have tobe examined in order to decide which
258 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
259
A different form possibly lives along the lower reaches of the Khatanga.
349. Lepus timidus tschuktschorum Nordquist (1883). Chukchi
blue hare
1883. Lepus timidus L. var. tschuktschorum Nordquist O., Antecknigar och studier till Sibirska
sshafskustens Daggdjursfauna Vega Expeditionens Vetenskapliga lakttagelser, Vol. II, pp. 84-90, figs. 8,
9-10; p. 88.
1903. Lepus tschuktschorum AllenI,, 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 OgnevS.I., Mlekopitayushchie severov-vostochnoi Sibiri (Mammals of
Northeastern Siberia), p. 77.
1927. Lepus tschuktschorum (partim? ) Sokol'nikov N. P. , Okhotnich'i i promyslovye zveri
Anadyrskogo kraya (Game and Commercial Animals of the Anadyr Territory), Byulleten' Moskovskogo
obschestva ispytatelei prirody, XXXVI, p. 133.
TYPE LOCALITY 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 62mm long, hair of flanks and underparts
80mm.
Skull massive, with broad nose and zygomatic arches. Length of skull
98-106 mm (116mm according to Nordquist) (M.103mm); zygomatic width
51-54.5mm (57mm according to Nordquist) (M.52.8mm). See Figure 125.
Length of body 610-680mm; hind feet 150-165mm.
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.begitschevi) in its color.
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.II, 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
Pp
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 russkikh 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 russischen Hasen, Zoologischer Anzeiger, Bd, 84, H. 1/4,
S. 72.
1936. Lepus (Caprolagus) Zolotarev N, T., Mlekopitayushchie basseina reki Imana (Mammals of the
River Iman Basin), p. 69.
Ears (measured from occiput) about aslong 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., Mél. Biol. Bullet. de
l'Academ, Imp. de St-Petersb. , III, S. 684-686.
1861. Lepus mandshuricus Maximowicz H., Zoologisch. Nachricht, Mél. Biol. , III, S. 693.
1862. Lepus mandshuricus Radde G., Reisen im Siiden 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 OgnevS,1., Materialy po sistematike russkikh zaitsev (Material on the
Systematics of Russian Hares), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, Vol. XXIII,
pp. 485-491,
21922, Lepus mandshuricus subphasa melanonotus OgnevS.I., op. cit. p. 489.
1923. Lepus mandshuricus Sowerby A, de C., The Naturalist in Manchuria, II-III, pp. 171-172,
1929. Allolagus (mandshuricus) Ognev S.J., Zur Systematik der russischen Hasen, Zoologisch,
Anzeiger, Bd, 84, H. 1/4, S. 72.
1936. Lepus (Caprolagus) mandshuricus 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
2May 1914, Peterthe 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
i)
i)
oo
261 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).
MATERIAL EXAMINED: 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.
262
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.
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 1/2
times as large as distance between anterior alveolus of premolar to tip of
lower incisor.
2963 Bullae osseae very small, little inflated and slightly protruding
forward (Figures 132-139).
Length of skull 82.1-89mm (M.85.4mm); zygomatic width 40-44mm
(M.41.9mm); length of nasals 32.8-37.9mm (M.35.5mm); width of nasals
14-18mm (M.15.7mm).
FIGURES 137-139, Skull of Allolagus mandchuricus Radde, No. M. 3069,
Maritime Territory, S.1. Ognev's collection
230
Length of body and head 420-540mm (M.473.5mm); hind feet (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) massive mandibula, 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 hindfeet. 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 35mm 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. Earsnear
the base rust-gray.
Forelegs rust-straw colored. This color is weakly marked on flanks
as two spots partly entering the inguinal region. Hindlegs 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.
23
266
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 (pl. 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 30mm wide, beneath the throat. Chest pale
pinkish straw (between light pinkish cinnamon and vinaceous-buff, Ridgway,
pl. XXIX and pl. 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
Caprolagus.
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) Oryctolagus; 4) Sylvilagus.
He includes in Sylvilagus 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 taxonomic relationship between the genera Caprolagus and
Allolagus is best demonstrated in table form.
C.hispidus Pearson (1848) (from the southern Himalayas) belongs to
genus Caprolagus. According to Glover Allen (1927), the following
species should be placed in the genus Caprolagus: C.sinensis
Gray (1833-1834) from southern China, andC.sinensis flaviventris
G. Allen (1927) from Fukien (southeastern China).
The monotypic genus Pentalagus Lyon which includes only the
species Pentalagus furnessi Stone (1900) from the Ryukyu Islands is
closely related to Caprolagus.
232
267 common:
Genus Allolagus
Ears measured from occiput about as long as
the head or longer.
Fur soft.
Skull slightly convex dorsally in profile. Highest
point of cranium behind supraorbital space.
Postorbital processes with anterior and
posterior angular processes. Anterior process
sometimes poorly developed,
Rostrum long. Nasals measured along oblique
line about as long as frontal suture.
Incisive foramina about as long as bony palate.
Interorbital space posterior to postorbital
process about 60.4 % of the width of both nasals in
their posterior third,
Genus Caprolagus
Ears much shorter than head.
Fur coarse.
Skull flattened dorsally in profile.
point of cranium in the occipital region.
Postorbital proces with only a posterior
angular process. Anterior processes completely
fused with frontals.
Highest
Rostrum very short. Nasals measured along
oblique line markedly shorter than frontal suture,
Incisive foramina much narrower than length
of the bony palate.
Interorbital space almost as wide as both
nasals in their posterior third,
Pentalagus and Caprolagus have the following characters in
1) ears very short;
i.e., very long;
Dal 3
m. > instead of m. ae
brown both dorsally and ventrally.
on the forelegs. Pentalagus is,
genus.
2) fur coarse;
4) incisive foramina narrow;
of Pentalagus is not as flat asthatofCaprolagus. Itsdentalformulais
3) bony palate primitive,
5) nasals shorter.
Pentalagus is uniformly colored, dull black-
Its claws are unusually large especially
in our opinion, a well characterized
We place the following species in the genus Allolagus: Allolagus
mandshuricus Radde, A.brachyurus brachyurus Temm. (1845)
from Japan (Kyushu, Shikoku — southern part of Hono);
Thos.
part of Hondo Island;
A.b.okiensis
(1906) from Dogo Island; A.b.etigo Abe (1918) — northeastern
and doubtfully A. b. angustidens Hollister (lene)
described from one specimen fron Hondo (Tateyama).
J. Abe ("'A Synopsis of the Leporine Mammals of Japan’! (J. of Sci. of
Hirosh. Univ."', Vol.I, p. 54-55,
1931)), attempted to analyze the
characters of Allolagus mandshuricus and Lepus brachyurus.
He came to the conclusion that L. brachyurus does not belong to the
genus Allolagus,
Allolagus.
We completely disagree with this.
differences between the two species.
As described in ''Zoologischer Anzeiger"'
and that it is more closely related to Lepus than to
It is closely related to the blue hare in his opinion.
J. Abe (1931) compares Allolagus
mandshuricus andbrachyurus*,
and concludes that there are great
feature of Allolagus 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
The skull
1929, the most characteristic
Allolagus
A. brachyurus
mandshuricus
Width of nasals in the middle ......0ccecceecsecs 14.5 — 18 16% *19'*
Width of interorbital space ... 1. eee eee ers erevees 10,2 — 13.2 i — 13
Width of nasals in the posterior third .........2+.++.- 1673 ="19F2 =
2) Nasals about as long as frontal suture or little shorter:
Length of nasals) » s,s, sssteyaccnetoemieneitthamat. « leila taneueh> yom ays S218 37.9 36 — 43
Length, of frontals: 4... cucnesians mpanotinualls selene, «sn iteiehlanay say 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 mandshuricus and brachyurus according
to J. Abe's measurements, there is no difference between the two species:
Aust see A, brachyurus
mandshuricus
Width of posterior nasal foramina.......+.++4-> 1th F2:8 ti2py— 14
Length of bony palate ooo, ame i .: cgepe ane) od en) op cnela) cals (fev e}?: Teas *9
4) Maximal width of incisive foramina about equal to length of bony palate:
Width. of incisive foramina.) coin s¥s) «dellelin eos) oie Taek O49 8.1— 11
Length of bony palate .... 2.22.5 cscecscevsenss (G3 192 Tee 9
5) Incisive foramina about as long as upper diastema:
Length of incisive foramina ......+4-+e++e++ee 19 —T Dies 20:5 = ye2.9
Length of upper diastema ...... eee essere eves 202)! 23 2le —r25
This comparison shows that there are no significant differences between
the skull of A. mandshuricus and A. brachyurus 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 Caprolagus brachyurus niger Noack, from the lower
reaches of the Amur. He did not give a fulldescription, 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 Lepus mandshuricus subphasa melanontus.
I was induced to do this by an interpretation of lower systematic
categories given by V.L. Bianki in ''The Species and Infraspecific Taxa",
((''Vidi podchinennye 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
FIGURES 140-143, Skull of Caprolagus hispidus Pearson
235
271
272
only in some geographical regions. A geographical natio may possibly
develop from an aberration.
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 northern boundary ofthe distri-
bution of the Manchurian hare follows the bank ofthe Kop' river*, continues
alongthe Amur river near Verkhnee Tambovskoe, Srednee Tambovskoe.
Inthe west it follows the Bureya Range, Little Khingan and Great Khingan. Its
range also includes the basins ofthe 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 relatedspecies, does
not have open lairs. It settles in holes in tree trunks which are open on
one side. I found beaten tracks to sucha 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, 1, p. 417 (as a subgenus of the genus
Lepus'),
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.
* 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
FIGURES 144-147, Allolagus brachyurus Temm, No, M.4518. Female ad, , 27 January 1935,
Nagoya, Hondo, Japan, N. Kuroda, S.1, Ognev's collection
Drawing by Yu. A. Kostylev
237
FIGURES 148-151, Skull of Oryctolagus cuniculus L, Specimen No, S,3319, from
near Kalish, MSZM* collection
Drawing by Yu. A. Kostylev
* [MSZM— Moscow State Zoological Museum. ]
238
274
Mesopterygoid fossa much narrower than the length of the bony palate.
Interparietal well developed in adult and old specimens. Postorbital
processes narrow, thin, swordlike, not triangular as inmany 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
AdIolagus *.
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.I, ed. 10, p. 58.
1837. Lepus vernicularis Thompson I., The Athenaeum, p. 468 (nomen nudum‘).
1843, Lepus vermicula Grayl., List Spec. Mammal. British Museum, p. 128 (nomen nudum! ).
1857. Lepus cuniculus BlasiusI., Naturgesch., S. 426-428.
1867, Cuniculus fodiens Gray 1., 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, I, part. 3-4, pp. 402-406.
NAMES: The Rabbit or Cony (English); Das Kaninchen (German); Le Liévre 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
locality. The type is apparently lost,
Principal figures: 1) Schreber1.Ch., Die Saugethiere, Erlang., 1V, 1792, S, 891-897, Taf. CCXXXVI,
A (entire animal in color); 2) Bingley W., Memoirs of British Quadrupeds, 1809, p. 309 (color. pl.);
3) Blasius I. , Naturgesch. der Sdugethiere Deutsch]. 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, Mammiféres, Paris 1885, p, 189, fig. 79 (entire animal); p. 182, fig. 77
(skull); 6) Fokht K., Mlekopitayushchie (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,
Uber das System der Nagethiere, Upsala 1899, Taf.1, 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. III, 1906, pl. 52 (color), pl. 48 (biological scene), pl. 50 (photograph); 11) 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 Sdugethiere 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); pl. XII (head of male and female rabbit); pl. XIII
(same anteriorly); p. 203, fig. 35-37 (diagrams of burrows); pl. 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); pl. XIX (I) (feet); pl. XX (6) (feet); fig, 48, pl. 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
Oryctolagus
239
(mandible); 14) Miller G.S,, Catalogue of the Mammals of Western Europe, London, 1912, p, 488, fig. 101
(skull lateral, ventral and dorsal); 15) Thorburn Archibald, British Mammals, Vol. II, 1921, pl. 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 61-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 cugly. Tears 6: p.1 - 1; Aa. 2 =2)4 Total Gf
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 diastema 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 (fromcorner 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 33-37.6mm (35.4-38.6 according to G.S. Miller); width of nasals
14.6-12:6 mm: *
275
* In the middle. According to G.S. Miller, maximal width of nasals 14-17.2mm.
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 60-72mm (65-70 according to G.S. Miller);
tail 52-69mm (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 (pl. XXIX, Ridgway, 1912), wood-
brown and fawn color (pl. XL)).
A broad, light brown stripe across chest. (between avellaneous, pl. XL,
and light pinkish cinnamon, pl. 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. andO.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. brachyotus Trouessart (estuary of the Rhéne), O.
crassicaudatus crassicaudatus I. Geoffroy (Transvaal), O.c.
curryi Thos. (Orange river), O.c.nyikae Thos. (Niassa, Nyika
Plateau).
GEOGRAPHICAL 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
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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 inthe 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 onhis estate near the Khadzibeiestuary...
"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.
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278
209
J
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 mainly 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.
"In spite of the proximity of the rattling plant, rabbits are very common
in itsiyards.444,
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 inthem. 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
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280
the lower half of the ravine. The stone quarries therefore consisted ofa
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 isacompletely 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
(5-6.00a.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. 1 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, 30-60cm below the surface, which is connected with
the surface by one, generally two (rarely three) galleries 1-2m long. The
size of the central chamber varies. It is usually 40-60cm wide and 25-
40cm 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, andare 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."
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281
Numerous fecal pellets are usually found near the burrows on small
mounds, 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 gestationof the rabbit lasts 30 days. This corresponds
with data obtained inthe 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
.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 populationof 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: 1) 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. Inadditionto
foxes the rabbit ishuntedbyMartes foina, Mustela putorius
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,
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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, andit is inconceivable that we should exterminate them, as
some people have requested.
SECOND SUBORDER. SIMPLICIDENTATA
Maxillae with only one pair of large, curved, rootless incisors covered
with a thick layer of enamel only anteriorly. Number of premolars
2 ori; molars 2 to 3.
1 1 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 in most forms but sometimes very small.
The extinct family. Ischyromyidae is considered intermediate
between the Duplicidentataandthe Simplicidentata. They lived from the
Lower Eocene to the Oligocene.
KEY TO THE FAMILIES OF THE SUBORDER SIMPLICIDENTATA
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 covered with more or less soft hair.
5 (8). Toes of hind legs connected by a bare broad web. Tail squamous,
= 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) aq:
4
10 (11). Molars (including premolars) usually q? only in Ternes (Caucasian-Persian Squirrel) z°
Tail always without horny scales, shaggy, often depressed dorsoventrally. 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)
q
4
11 (10). Molars (including premolars) always ie
Tail without horny 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 Te
4
13 (16). Molars z3 first molar very small.
14 (15). Hindlegsnot 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 hair, 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 1.5mm. Molars covered by
gums, not taking part in chewing. Food is cut by the enormous, sharp, curvedincisors, Anterior surface
of incisors with a deep longitudinal groove.
Seleviniidae*
18 (17). Molars normally developed and protruding from the gums.
284 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
285
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). Claws 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. Mémoires mathém, , 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, Petauristidae 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. Atthe 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: iz; c S p=; ms = (22).
GEOGRAPHICAL DISTRIBUTION. Forest zone of the northern
hemisphere. India, Ceylon, Burma, South China, Sumatra, Java and the
southern parts of the U.S.A.* Inthe 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.
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286
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**: 1)
Jom y's® Thes* (type species J. horsfieldi Waterh. ) - Java, Borneo,
Malacca; 2) Belomys Thos. (type species B. pearsoni Gray) -
Himalayas, Sikkim, Yunnan, Assam, Formosa; 3) Pteromyscus Thos.
(type species P. pulverulentus Gimnth.) - Malacca, Borneo; 4)
Petaurillus Thos. (type species P. hosei Thos.) - Borneo; 5)
Pteromys G. Cuvier (type species P. volans L.) - distribution given
below; 6) Glaucomys Thos.ft (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? (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,, Legons d'anatomie comparée, recuillies et publiées sous ses yeux, I, tab, 1
(Nomen nudum‘ )t
1817. Pteromys Cuvier G,, Le Régne Animal, Vol.1 (description of the genus); See also Editio Il, 1829,
p. 194,
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 Mammiferes, p. 255,
1826. Pteromys Cuvier F,, in: Despréaux L., Dictionnaire des sciences naturelles, dans lequel on traite
méthodiquement des différents étres 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, anew 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 Sciuropterus Group,, Ann. and Magaz. of
Natural History, ser. 8, Vol.I, No.1, 1908, pp. 1-8.
+ 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.
t 1 could not obtain the first edition of ''Legons d'anatomie comparée, There is no description of the
Flying squirrel in the second edition (t. II, p, 152, 1837). Cuvier writes as follows: ''D'autres
mammiféres, savoir, les galéopitéques, les polatouches ou écureuils volants, et les phalangers
volants, ont des membranes entre les pattes, mais sans alongement des doigts; elles ne peuvent
pas servir a les élever, mais elles les soutiennent assez bien en l'air pendant quelques temps de trés
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 ''Régne Animal".
249
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 massive. 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 5.2mm 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 of Pteromys; 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 andthe American Glaucomys*.
The sides of the rostrum converge anteriorly inGlaucomys: therostrum
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.
288
f) = 2
FIGURE 153, Os penis of Pteromys volans L, FIGURE 154, Glans penis of Pteromys
volans L,
1 — lateral; 2 — ventral
From a specimen collected by I. S. Slesarev in the 1 — lateral; 2 — dorsal.
Central Reserve (former Belyi County, former
Smolensk Province). Highly magnified,
Drawing by Yu. A, Kostylev
* 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 Pteromys 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 Pteromys volans L, (lateral)
Drawing by Yu. A. Kostylev
~
Fi ae TN
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. Inthe east Pteromys occurs in
Manchuria, northern and central China, Korea and Japan.
163. Pteromys volans L. (1758). Common flying squirrel
1758. Sciurus volans Linnaeus C., Systema Naturae, ed. X, p. 64.
NAMES: Flying squirrel (English); Flatterhérnchen, das fliegende Eichhérmchen (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
289 Ob); ive-lanki (along the Irtyn); pailanglangi (along the Surgut); poeze (along the Narym and Ket');
sartyin (Bashkir); nyar-brod (Komi-Zyryan); maskalan (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, Samagir); umugi (Udehe on the river Khor);
ornke (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 FIGURES: 1) Schreber J.Ch., Die Sdugethiere, 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, pl, 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 Sdugethiere 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
290
291
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 Sdugetiere 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 Flatterhornchen aus dem Kirgisengebiet, Ztschr. fiir Sdugetiere, 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.1. , 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. 79 (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 coveredwith 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 hind feet both 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.
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 3mm 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
* 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 2.5-3mm.
FIGURE 157. West Siberian flying squirrel
Drawing by K. K, Flerov
FIGURE 158. Head of flying squirrel, Pteromys FIGURE 159. Ventral side of feet of flying
volans L. squirrel
Drawi aN.
ey er 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 inEast 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 infront ofthe eyes. They are up
to 60mm long and there are 12-14 hairs in
each group. Above this group there are
3-4 vibrissae on each side of the forehead.
FIGURE 160. Hind foot of Sakhalin
flying squirrel, Pteromys volans
alchenen Those Posti<omalouskal A few smaller vibrissae are present
[Korsakov], Sakhalin above the eye. One group near the external
Photograph by S. 1. Ognev 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 narrowed anteriorly 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, roundedand 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
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 FIGURE 162. Skull of Pteromys volans
volans L., 18 November 1930, Sharya District, athene Thos. , Post Korsakovskii [Korsakov],
northern Gorki Territory Sakhalin Island
Photograph by S. I. Ognev Photograph by S.1. 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, makingthe 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
of Pteromys volans crosses Russian Lapland, turning south towards
the east. According tol. F. Brandt (''Mammals of North European Russia’
(Pozvonochnye severa Evropeiskoi Rossii), 1856, pp. 31-32) the flying
squirrel was distributed as far as Finnmark. However, thishas 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)
295
296
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 inthe 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
Sci@rus*Vuigaras exalbidt's 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 viliage 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
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 from 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 Yershichidistricts.
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 (1895), 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
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
297
* W.W. Rajewski und L.G. Kaplanow, Zoolog. Anzeiger, Bd. 84H. 5/6, 1929, S. 148.
259
298
other counties of the former Yaroslavl Province. One specimen was
caught 30km 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 fhe 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 (1871), 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°S 5)ata))e
According to L. P. Sabaneev (1874), it occurs rarely on the Urals in the
Goroblagodatskii and Bogoslovskii districts. It is common 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,' 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 1. 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 (1928), 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. G.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.
299 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. Iohansen (1923) reported it
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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 Tunguskarivers. According
to N. P. Naumov (1934), it is common along the Lower Tunguska. According
to A. Middendorf (1861), it reaches 68°N. along the Yenisei. Tell
(Nordenskjéld'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 inthe 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
Lena Basin (I. Polyakov, 1873) and onthe 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.1. Ognev, 1926). According to N. P. Sokol'nikov
(1927), it occurs in deciduous forests 50-60km 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 inthe 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, 50km 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
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301
(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. Sofronovy (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 inbirch groves.
Along the Irtysh they occur in the region of ''ribbon'' pine forests
together with Sciurus vulgaris exalbidus Pallas. According to
I. Ya. Slovtsov (1892), 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.
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, blackbirch, spruce andfir. 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.'"'
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302
In the mountains of Siberia, e.g., on the Altai, the flying squirrel is
found 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 »xpedition,
in a large forest at an altitude of about 1000m.
In extreme northeastern Siberia, e.g., in the Anadyr Terricory, the
flying squirrel occurs in larch forests along the Anadyr.
We know very little of its habits. It begins its activity 1/2 -1hour 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 tound 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 10-15m, beginning at the base of one tree
and ending at the base of another (A.N. Formozovy, 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 40m. 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 squirrel 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 gunandhide. 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 they are
not afraid of man."
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303
A.N. Formozovy (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 ofthe 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 3-10m above the ground. One
tree would sometimes contain several hollows, each occupied by a nest.
The nests were built of lichen (Usnea barbata and USnea
longissima).
In birch groves in Baraba 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 (Dendrocopus major and Picus
martius).
V.V. Gubar' (1930) gives the following data for the Novosibirsk flying
squirrel: ''Flying squirrels are usually found in hollows in high aspens
3-4m above the. ground. Only once did I find an inhabited hollow very low,
at a height of 1.5m. It is simple to determine whether a hollow is
inhabited. When the trunk is tapped withanax, the flying squirrelcomes half
out of the hole or jumps out completely. Confusedby the light, its movements
become very slow. The frightened animal climbs about 1m 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 itis almost invisible onthe bark. Atthetopofthe 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 ona branch. They always land on the trunk
FIGURE 163, Flying squirrel on a bough
Drawing by K. K. Flerov
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304
305
about 1-2m 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,
5-9m 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 5m above the ground. The opening was 6cm wide. The
nest was at the bottom of the hollow, 63cm 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 30-400g. 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.
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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, didthey accept pine cones. The completely ignored
pine nuts.
"The flying squirrel is rather voracious. One pair consumed 1.5 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.1. 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 membrane. 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 Sciurus 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
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 (nec Mus volans Linnaeus,
C., lc., 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 Sdugethiere, I, S, 451 (Russland),
1822, Pteromys sibiricus Desmarest A.G., Mammalogie ou description des espéces de
Mammiféres, 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 vulgaris Simashko Yu., Russkaya fauna (Russian Fauna, II), pp. 597-599,
1857, Pteromys volans Blasius J.F., Naturgeschichte der Sdugethiere 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 Mammiféres d'Europe, 1910, p, 116; Miller G.S., Catalogue of the
Mammals of Western Europe, 1912, pp, 941-946; Ognev S.1., Mlekopitayushchie Moskovskoi gubernii
(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
307 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 of Pteromys 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 38.2-41.3mm (M. 39.92 mm);
condylobasal length 34.9-37.8mm (M.36.4mm); zygomatic width 22.8-
25.2mm (M.24.02mm); length of nasals 11.9-13mm (M.12.33mm); length
of upper molar row 7.1-7.7mm (M.7.28mm).
Length of body 149-172mm (M.164.7mm); tail without terminal hair
98-130mm (M.113.1mm); 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:
1) 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: 1) 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.
308 SUMMER FUR. Back blackish gray (dark mouse-gray) with straw-gray
mottling (between drab and mouse-gray). Color of hairs as follows: 1)
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 12-14mm.
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.ognevi 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). Sbornik 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 zhurnal, XV (3):539. Moscow
1936,
TYPE LOCALITY AND DEPOSITION. Lake Peno in the Western Region (sources of the Volga, former
Ostashkovskii County of the Tver Province). Type No, M4158 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 36.7-39mm; condylobasal length
34.7-36.7mm; zygomatic width 23.9-24.5mm; length of nasals 12.4-
12.9mm; length of upper molar row 6.6-7.5mm. Length of body and head
160-163mm; tail without terminal hair 95-120mm; hind feet without
claws 34-37.mm.
ADDITIONAL DESCRIPTION. Winter fur. Straw-rust tinge, (close to
avellaneous (Ridgway, 1912. pl. 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
more intense in the inguinal region and on the ventral side of the parachutes.
Tail yellowish straw-rust colored (near avellaneous, pl. XL, with pinkish
buff tinge, pl. 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 inthe 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.ognevi differs from P.v.athene as follows: 1) 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. Notknownexactly. 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 russicus 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 Sibirskoi kraevoi stantsii zashchity rastenii, 1, 1930, pp. 78-79.
1934, Pteromys volans gubari OgnevS.I., Obzor letyag (Pteromys), 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. M4036 female, Dec. 1930, V. V. Gubar', type in S. 1. Ognev's collection.
MATERIAL EXAMINED: Holotype and 12 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 volans 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 39.2-41mm (M.40.2mm); condylobasal length 35.7-
36.9mm (M.36.5mm); zygomatic width 23.1-24.5mm (M. 23.8mm);
length of nasals 12.1-13.1mm (M.12.5mm); length of upper molar row
7-7.5mm (M. 7.2mm).
Length of body 135.5-171mm (M.155.3mm); tail 109-140mm )
(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, pl.LI).
Hair on back 15-17mm long. Color as follows: 1) 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
311
312
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.
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.gubari 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.betulinus 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.inecanus as follows: 1) hair shorter and less luxuriant; 2) tail
without straw-rust tones. Its summer fur differs from that of betulinus
as follows: 1) 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 betulinus Serebrennikov M.K., Ein Flatterhémchen aus dem Kirgisengebiet,
Ztschr. f, Sdugetierk, Bd. 4, S. 142-143, Taf. XII.
1934, 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 belulinus OgnevS.1,, 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 (200 km 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.40.4mm); condylobasal length 35-
37.3mm (M.36.58mm); zygomatic width 22.9-25.4mm (M. 24.07 mm);
length of nasals 12-13.1mm (M.12.5mm); length of upper molar row
7-7.3mm (M.7.19mm).
Length of body 116-172mm (M.157.1mm); tail without terminal hair
98-119mm (M.113mm) (122 accordingtoM.K. Serebrennikov); hind feet
without claws 34-39mm (M.35.8mm).
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.
313 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.betulinus andP.v.
gubari are adjacent. The color of P.v.betulinus closely resembles
that of P.v.athene. The winter fur of the Semipalatinsk flying squirrel
differs from that of athene as follows: 1) 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. athene; 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
M.K. Serebrennikov (1929) certainly described a partly albinistic
specimen of P.v.betulinus.
The summer fur of all forms of Pteromys is darker than the winter
fur as arule. Inthe holotype of P. 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. betulinus 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 of Pte romys in western
Siberia. North of Barnaul and Biisk it is replaced by a grayer and very
paletorm, Fv. Subpart.
354. Pteromys volans turovi Ogn. (1929). Central Siberian flying
squirrel
1858. Pteromys volans Schrenck L., Reisen und Forschungen im Amur-Lande, I, S. 116-118; MaakR. ,
Puteshestvie 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, Ill, 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. 14, S.75; Ognev S.I., Mlekopitayushchie Shantarskikh ostrovov (Mammals of
the Shantar Islands), Izvestiya Tikhookeanskoi nauchno-promyslovoi stantsii, 2 (5): 1929, p. 1.
TYPE LOCALITY AND DEPOSITION. Bol'shie Koty (near Lake Baikal). Type No. M. 1241, 6 Oct. 1923,
collected by Professor S.S, Turov. S.1I. 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
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 38-41mm (M.40.3mm); condylobasal length 31.2-37.6 mm
(M.35.8mm); zygomatic width 22.2-24.7mm (M.23.68mm); length of
nasals 12-13mm (M.12.39mm); length of upper tooth row 7-7.7mm
(M.7.20mm). Length of body 170-172mm; tail 98-110mm (M.103.5mm);
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: 1) 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. Inthe 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, 10-12mm long with well
315
275
317
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 andthe 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
of P.v.turovi. P.v.athene is easily distinguished from P.v.
turovi as follows: 1) 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 intheir lighter, grayer color. Their resemblance toP.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, Lake Bolon!' -Odzhal,
Khor river).
355. Pteromys volans athene Thos. (1907). Sakhalin flying squirrel
1861, Pteromys volans Schmidt Fr., Bemerkungen iiber die Sdugethierfauna Siid-Sachaliens, etc, ,
Mélang, 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.
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 aluco OgnevS.]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-
38.2mm (M.37.07mm); zygomatic width 23-25.2mm (M. 23.74mm);
length of nasals 12.3-13.7mm (M.12.74mm); 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 witha 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:
1) 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: 1) 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 yellow-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. Insome specimens it is white witha 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: 1) 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 andP.v.
turovi have been described above. The color of the winter fur of P.v.
athene 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,
Zt
319
No.4, 1921, pp. 208-209) described a new subspecies, Sciuropterus
russicus orii, fromsixspecimens 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 dataon P.v.athene 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 39.5mm; basal length 30.5mm;
zygomatic width 24mm; postorbital width 9mm; interorbital width 7.5mm;
width of rostrum in front of nasals 6.5mm; length of nasals 13mm;
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. orii 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 Pteromys momonga Temm. (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 forspecimens 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 espéce, due
aux soins de M. Biirger, dans les foréts de l'intérieur, ou elle vit dans
les troncs des arbres et se nourrit de noyaux et de semences’’.
Finally, the status of S. 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, 183km 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 I., 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-promyshlennosti vy Kolymskom okruge (A Survey
of the Animal Trade in the Kolyma District), Sankt Petersburg, p. 67.
1903. Sciuropterus russicus AllenJ.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 OgnevS.I., Mlekopitayushchie Severo-Vostochnoi Sibiri (Mammals
of Northeastern Sibiria), pp. 89-93 (partim!).
TYPE LOCALITY 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: 1) 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.41.01mm); condylobasal length 37.1-
38.7mm (M.37.83mm); zygomatic width 24.6-25.6 mm (M. 25.03 mm);
length of nasals 11.1-13.6mm (M.12.73mm); length of upper tooth row
7.2-7.7mm (M.7.4mm).
Length of body 161-180mm; tail 100-141mm; hind feet 31-38mm.
DESCRIPTION. Winter fur resembling that of P.v. zyubari 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 24-26mm. Color as follows: 1) 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. gubari.
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.
Sell Hair on back up to 20mm long. Color as follows: 1) roots broad deep
gray (between deep mouse-gray and deep neutral gray); 2) well marked,
wide, whitishstraw-gray, subapicalbands; 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.
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A ALVId
280
Throat, chest and belly white with a slight yellowish tinge.
Hair on tail up to 33mm 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: 1) 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.incanus differs from that of gubari as
follows: 1) straw tones on back less developed; 2) fur duller and longer.
I 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 (Ihave 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 of P.v.incanus is not
clear. It is not known which subspecies occurs in the northern Yenisei
Territory. Inthe south, the range of P.v.incanus is adjacent to that
of P.v.turovi, inthe southeast to that of P.v.athene, inthe
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 OgneyS.I., Mlekopitayushchie severo-vostochnoi Sibiri (Mammals
of Northeastern Siberia), pp. 89-93 (partim!); Ognev S.1., 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!).
322 1927, Pteromys volans incanus 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. XXXVI, 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 thaninP.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 25.3-25.7mm; length of nasals 13-14mm; length of upper
tooth row 7.5-8mm.
281
323
Length of body 170-207mm; tail 95-110mm; hind feet 35-37mm.
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: 1) 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.incanus 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 inthe
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 silvery P.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
324
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) 38.3mm; condylobasal length
36mm; 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 dorsally 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
Ssayal 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 1 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 157mm; tail 149mm; hind feet 35mm;
ear 19.5mm.
Length of skull 39mm; basal length 29.5mm; maximal width of skull
(apparently measured at the level of the zygomatic arches) 23mm; nasals
12.2X5; interorbital space 7.5mm; width of braincase 17mm; length of
palate 16.2mm; length of palatine foramina 5.5mm; length of bulae osseae
10.2mm; length of row of teeth (without p*) 6.5mm; greatest width of m!
Sei rgalente
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 ofthe Maritime Territory and Korean flying squirrels can not
be basedon skull structure or measurements of skulland 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 ofsuchaclear gray as in
russicus. Upper surface of hands and feet gray, the light hairs on the
toes cream colour, more extended proximally than inthe 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 lighttail, 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) 103mm; hind feet
30mm; ear 16mm.
Length of skull 38.2mm; condylobasal length 34.7mm; zygomatic width
24.2mm; width of skull at level of auditory openings 20.2mm. 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: 1) 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 small.
Frontals with well developed postorbital processes. Bony palate wide.
p “; m ws 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
326
327
1) Sciurinae. The subfamily includes the genus Sciurus, subgenus
Tenes (=Oreosciurus) andthe American subgenera Parasciurus,
Araeosciurus, Neosciurus, Hesperosciurus, 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 alsoMyosciurus.
Distribution: Ethiopian and Oriental Region.
4) Callosciurinae with the genera: Callosciurus, Menetes,
Tomeutes, Rhinosciurus, 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 SUBFAMILIES OF THE FAMILY SCIURIDAE IN THE U.S.S.R.
1 (2). Toes unusually long and slender. Outer margins of fore- and hind legs with stiff pectinate fringes
of hair which may be 5mm long. This hair is especially long on the outer side and at the tip of the 5th toe
of the hindfeet, where it forms a tuft which is longer than the claw.
Toes with black claws; longest claw usually longer than 1cm, sometimes 1.5cm.
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.
Sciurinae.
4 (3). Earsshort, 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,
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. Fourthdigit 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 an Es i 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
(Callosciurwus).
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., Systema 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 -
1/5 shorter than body.
Hair on tail very long, soft and directed laterally in ventral view.
Ears long, at least twice as longas broad. Hair ontips ofears very long,
forming well marked tufts in winter specimens. Cheek pouches not
developed.
Vibrissae arranged as follows: 1) 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:, pylel:. av2e2ydeld- t>.totalss
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., inS.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 Tenes 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 (Sciurus (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 chipmunks,
286
The infraorbital foramina are very small, laterally compressed, with
small angular processes at the base. These processes are especially well
developed inS.vulgaris andinthe AmericanS. niger. They are
absent in Tamiasciurus hudsonicus.
Diastema of moderate length. Diastema about 1/5 to 1/6 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 re spects transitional to the chipmunks), Baiosciurus,
Sciurus, Guerlinguetus. 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 Sciurus Linn.
2 (1). Color of fur not changing or changing very little with the seasons. Belly various shades of
tust-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 Eichhémchen (German); 1'écureuil (French),
LOCAL NAMES: belka sosnovka, elovka, berezovka (Russian hunters); belka chernaya, kedrovka,
Shamanchik (Russian andSiberian 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
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, ulyuki (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); yavunya (Koryak, Nymallan); raulen, uliki, rodlia
(along the Anadyr); ting (Yakut); pogodzhub (Yurak, Odul); lerchieng (Tundra inhabitants, Yukagirs,
Oduls); khuishu, Khui-dzhu, khui-ku-tzu [gray dog] (Chinese).
PRINCIPAL FIGURES: 1) Schreber J. , Sdugethiere, Erlangen IV, 1792, S. 760, Taf. CCXII (drawing
of S. fusco-ater); 2) Bingley W., Mémoires of British Quadrupeds, 1809, p. 277, color plate (entire
animal); 3) Nilsson S., Illuminerade figurer till Scandinaviens Fauna, I. 1832, pl. 11 (entire animal
in color — winter fur); 4) Simashko Yu., Russkaya fauna. Il. Mlekopitayushchie (Russian Fauna. II.
330 Mammals), 1851, pp. 589-594, pl. 27 (skeleton, teeth); pl. 31 (entire animal in color); 5) Blasius J. ,
Naturgeschichte der Sdugethiere Deutsch]. , 1857, S. 272, 274, Fig. 153, S. 271 (halftone plate), Fig, 154,
S. 272 (entire animal); 6) Mewes W., Holmgren A.E., Atlas 6fver Scandinaviens Daggdjur. Stockholm
1873, pl. XI, 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, Mammiféres, Paris 1885, p. 111, 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, pl. 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. XLI,
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.1, pl.4 {entire animal in color); 14) Millais J.G., The Mammals of Great Britain
and Ireland, London 1906, Vol. III, pl. (entire animal in color); 15) Meerwarth H, , Lebensbilder aus der
Tierwelt, I, Leipzig 1908, S, 249-250 (photograph of animal); 16) Schaff E., Die wildlebenden
Sdugetiere 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, Sdugethiere, II, 1914, S, 538 (color plate of Siberian squirrels); 19) Thorburn Arch. , British
Mammals, I, London 1920, pl. 24 (color plate); 20) Barrett-HamiltonGer, 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); pl. XXXVI, p. 700 (tails); pl. 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,
pl. 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.N., Sledopyt-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); pl. A, pp. 244-245 (diagram of spring molt); pl. C, pp. 246-247 (diagram of
autumn molt): 26) Lihring Rud., Das Haarkleid vonSciurus vulgaris und die Verteilung seiner
Farbenvarianten in Deutschland, 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. 1la-1la', 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-25c S.
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, pl. I (color types of
Siberian squirrels, color plate); pl. II (transitions between gray specimens and albinos); pl. III (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
promysel] (Squirrel Hunting), Moscow 1931, p.5 (a squirrel near its nest); p, 6 (a squirrel nest); p. 14
288
331
333
(a swimming squirrel); p. 29 (a squirrel track); p. 32 (a wounded squirrel); p. 44 (a squirrel on a tree); 31)
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 zhurnal, 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., Isakov Yu. 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 18-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.
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).
heats: p) l= ewae 25. i als totale.
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.1. 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 of Tenes anomalus.
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 withtwo
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 small 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. M2? 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 betweenthesetwo. The inner side ofthe 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. M7” 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
336
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.
adultus and senex. Serebrennikov does not give a detailed description
of the last.
Description of cranial characters of different ages of squirrels.
Juvenis. Tothefirst stage belongblind squirrels a few days old,
which 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 21mm. Skull oblong-oval; zygomatic arches little
protruding; face very short compared with braincase. Bones very thin;
fontanel nearly always present between 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 23mm 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 34mm.
Lower anterior milk premolars cutthrough. Mandible 23.8mm long and
11.7 mm high.
Third stage. The first true molars appear. Condylobasal length
38mm. Zygomatic arches 25mm 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.
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 25-26mm. 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, 1).
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 ofthe 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 stageof the subadult lasts from 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 ofthe subadult (Figure
LGOgn 2);
The growth of the teeth and the replacement of the single deciduous
premolar on the mandible 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 ofthe sagittalsuture 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).
1. KEY TO MAIN AGE GROUPS.
A. Upper tooth row incomplete, Permanent molars either absent or breaking through. The tooth row
(including molars and 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
1 Anterior root of p2 projecting far anteriorly,
sometimes reaching beyond alveolus of p': subadult, up to one year old,
p@ triangular, its cross section about half as wide as m
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 pi: adult,
* This table has been modified and corrected in some details.
294
338 Il. KEY TO SUBGROUPS OF SUBADULTS AND ADULTS.
339
A. Tooth row not yet with level surface i.e. , p2 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.
a2, 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 worm 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 connected 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),
e2, 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).
£2, Exposed dentine covers more than half of the occlusal surface, Surface of the teeth
depressed: more than 6 years old (Figure 170, 10).
The color of the squirrels is very variable and the variations are
described under the various subspecies.
The belly of all forms of Sciurus 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
342
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 Sciurus vulgaris have atypical 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 25mm 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 25mm long. B.A.
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 2cm behind the first. These
vibrissae are 30-35mm 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 inthe 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.vexatvovaws Palityete.
The following subspecies occur in Western Europe: 1) Sciurus
vulgaris vulgaris L. —Scandinavia; 2)S.v.leucurus Gmel. —
296
FIGURE 170, Stages in the abrasion of molars of Sciurus 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 (m3) 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 pe strongly projecting forward, First premolar usually absent.
Areas of exposed dentine (the bony substance of the tooth under the endmel) present only on the large outer
tubercles of second premolar.
2. 3-4 months, Upper tooth row complete, only p! absent. pe deciduous. Areas of exposed dentine
on outer tubercles of p“ andm’. A fine line of exposed dentine sometimes present on inner tubercle of pe.
3. 4-5 months, Tooth row usually incomplete or not yet level. Deciduous p~ replaced by permanent
tooth, Deciduous pe 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
Z 1
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
1
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 m3,
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 to enlargedfalcate 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 coalesced with dentine spots on inner tubercles of
all molars, forming horseshoe-shaped figures on m! and m2. 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
with the horseshoe-shaped figures of p’, m! and m2. 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
England; 3)S.v.russus Miller — western and central Europe, from
Britanny to Holland and central France; 4)S.v.fuscoater Altum —
343
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) Sphnnve
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.infuscatus Cabrera —Spain, Las Navas, Avila;
11)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.vulgaris 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 of Sciurus vulgaris are known so far.
343 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
f
FIGURE 171. Glans penis of squirrels
(highly magnified)
rT
FIGURE 172, Os penis of squirrels, 1 -
lateral; 2 - dorsal (highly magnified)
Drawing by Yu. A. Kostylev
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 (1930)
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 onthe left
bank ofthe Taz. Inautumnof 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 1 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 4.5m high."'
299
345
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 Khatangaalongthe 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 Olenekriver (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. Iokhel'son (1898), the squirrel disappears
northwest of Srednekolymsk, near the circumpolar 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 sucha
forest, 170km wide, 50-60km 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, 1-3km 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).
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
346
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, inthe 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,
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 common 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., inthe 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, onthe 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.
347 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 isa
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 mostly 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 I1.D. Kiris (1934), who studied the ecology of
the central Ob squirrel.
1. The characteristic habitat in the Surgut Territoryis 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
349
350
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.
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 toI.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
<upply 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 of Sciurus 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 humidtype. 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 suchas 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 indepressions alongrivers.
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
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forests) to 213km 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: 1) green-moss pine forest; 2) dry-
lichen pine forest; 3) grassy-steppe pine forest.
a
Ce *
Het
(po
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 (Phragmites 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 =Sciurus vulgaris exalbidus Pall.]
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covered here by the leaves of the deciduous trees, pine needles and peat
moss.
The grass cover is dense, including also Solidago virga
aurea L., Artemisia campestris forma glabra, Hypochoeris
maculata L., Pyrola sp.
some places.
Yellow acacia trees form underbrush in
Higher up, the trees surrounding the marsh are replaced by
old pine forests with thin grass cover in which Festuca ovina L,
predominates.
A few of the following plants also occur among the grass:
Alyssum desertorum Statt., Artemisia campestris forma
glabra,
Euphorebia esula L.,
Koeleria glauca D.C., Melandrum album Larcke,
Pulsatilla patens Mil., Stipa
ioannis Czelak., Allium nutans L., Phragmites communis
Trm.,, Selidago’ virga avrea iL;
Silene volgensis Bess., S.
chlorantha Ehrh., Phleum boehmeri Wibel, Scabiosa
ochroleuca’ Dee
hiemale L., etc.
Lie
Uf
“1
“MO
Giz.
FIGURE 174, Squirrel looking out from behind
a tree
Drawing by A.N, Formozov
Hypochoeris maculata L.,
Equisetum
"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.
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
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
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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
2-3m 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 atree. When threatened it hides by pressing
itself tightly to a branch or conceals itself among the dense branches.
Jnr
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 suow, 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,
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355
giving its typical squeak. When it is frightened it rapidly climbs to the top
of atree, 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 longtime. 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-
12km 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 seenasearlyas 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
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we kept a longtime 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
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 - 30cm thick. They are usually 5-15m 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.
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357
The measurements of the nests of Moscow squirrels are as follows:
length 20-50cm (generally 35-40cm); width 25-50cm (generally about
35cm); height 20-40cm (generally about 30cm). Average weight of a dry,
inhabited nest about 450g.
These figures show that the nest is generally spherical. The inner
space is 12-16cm wide. The entrance to the nest is round and is about
5-6cm wide (rarely 8cm). ‘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 WwW 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
twigand, 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 twig without 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),
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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 squirrelrests is carefully plucked, so
that it is always soft.
"The nest is usually spherical, sometimes ovoid. Its outer measurements
are as follows: length 50cm; width 30cm; height 35cm; circumference
100cm. The cavity of the nest is small as the walls are very thick. The
inner measurements are as follows: length 20cm; width and height 10cm.
The length of the cavity is not always in the same direction as the outer
length, but sometimes lies inthe short axis ofthe next. Everynest has twoexits.
The openings are small, and their width (about 5cm) is adapted to the size
of the squirrel. Each entrance is carefully concealed by dry grass, moss,
eter
"The Teleut squirrel usually builds its nest on pine and only rarely
On otheritrees.:).
358 "The Teleut squirrel chooses different habitats for its nests. In the
Srostinsk pine forest, about 50 %ofallnests 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...
"Hach 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. Inwinter itis usedas 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.
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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 theirterritory. 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: 1) 650 acorns in a hollow in a linden tree 3.60m above ground;
the hollow was 40cm deep; 2) 214 acorns ina hollow ina poplar, 6m above
ground (hollow 25cm deep). All acorns were fresh and none were spoiled.
From 30 September onwards the authors observed that squirrels stored
Boletus luteus inthe 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 30cm of snow. The
crossbills break off large numbers of cones and throw them to the ground.
* [Area near Lake Baikal. ]
4* 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.
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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 Vivipara 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 (1935) 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 Cantharellus 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
fuscesice ns.
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 8.45mg, 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 55g of food per day in summer, 70g in autumn, 35g in
winter and 80g 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
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the first dried mushrooms (Boletus luteus) in early August which had
apparently been collected in late July.
ODN
Ad
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 inits teethto the nearest tree, where it pushes
in into a fork of two branches or between the stem and abranch. 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 30cm to 4m 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. subtomentosus,
Boletus chrysenthereon, B. piperatus, Pluteus nana,
Clitocyle sp., Pholiota sp., Gomphidius glutionosus,
Ganoderma 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 andB. rufus, and other species were
poorly represented. On the other hand, only afew Boletus scaber and
B. rufus appeared in stores in 1931. Instead there were Gomphidius
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glutionosus,
Boletus luteus andB. 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 of a Lacerta agilis inthe stomach ofa
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).
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.
January
February
March
April
May
June
July
August
September
October
November
December
Mushrooms,
Mushrooms,
Mushrooms,
Mushrooms,
Mushrooms,
Mushrooms,
Mushrooms,
Mushrooms,
leaves
Mushrooms,
Mushrooms,
Mushrooms,
Type of food
pine nuts
pine nuts
pine nuts
pine nuts
pine pollen, animal food, rarely pine nuts
pine nuts, ant cocoons and other animal food
pine nuts, ant cocoons, sawfly larvae and other animal food
pine nuts, ant cocoons, sawfly larvae and other animal foods tree
pine nuts, sawfly larvae and other animal food
pine nuts and other vegetable food, ant cocoons, insect larvae
nuts, aspen buds and other vegetable foods, sawfly larvae and other
animal foods.
Mushrooms,
pine nuts, sawfly larvae
The squirrel needs water. Observations of this animal in captivity show
that it drinks much.
forest.
In dry years the squirrel keeps close to water in the
In winter it may satisfy its thirst with snow.
aS
364
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:
1) 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 andonits
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 oftubules 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
differently by 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.roburL., QO. mongolica Fischer, Corylus avellana
L., Juglans mandshurica Max, and others. The seeds of these trees are eaten by the squirrel.
** The grooves are studied with a megnifying glass with a magnification of 10 to 15x. The claw is
illuminated from different directions.
1 Korschelt, Lebensdauer, Alter und Tod, Jena, 1924.
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.
Anequal 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 intheir 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 inearly February (S. V. Lobachev, 1932). Inthe northern part of
the Tobolarea 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 zhimi i plodovitost'yu raznykh vidov mlekopitayushchikh). —
Izvestiya Akademii Nauk SSSR, 1930,
B17
1928) and in the Tara District in early March (V.E. Ushakov, 1927). In
the Turukhansk Territory it begins inlate 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); inthe Anadyr Territory in
the middle of February (N. Sokol'nikov, 1927).
FIGURE 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
368
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 5-10cm 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 ona 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. Lobachey (1932) observed during the Upper Vychegda
expedition that the second rut begins when the young of the first litter become
independent in the middle of June. The data of Gol'tsmaier (1935) confirms
Lobachev'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 121g). The female was still feeding the young
of the first litter, as her teats still contained milk. The embryos
were 3X2cm 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 inthis 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 ina
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 mammary glands functioned. If we
assume that this female gave birth on 15 November, it must have had estrus
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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 laststhe 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 formed
couples already in autumn. He also States that the Mari 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)
October
November
December
Subadultus
Est 1
(4.5-12 months old) strus rarely
eee ley 8
ee ee Si
ane
September
Adultus
(1-2 years old)
Estrus possi-
ble exception-
ally
Adultus
(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
372
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 ofembryos, andthe 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 ofembryos. 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 G.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, cur! their
tail between the legs and cuddle up against each other.
Measurements and weight of the newborn common Moscow squirrel:
Sex Weight in grams Body lengthincm Tail length in cm
Male 8 5.6 2.6
i 7.5 5.6 2,1
i 8.5 5,3 253
a 8 5.6 2.4
u. Zi Sau! 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,
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373
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 21st-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 5m 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.
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 (1mm) 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
6-7mm 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 onthe ears. At
this stage the terminal hair of the tail is 20mm 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 1.5g on
the average. The curve of growth and increase in weight is irregular.
According to M. P. Raspopov and Yu.A. Isakov, a 5 1/5-month-old
Moscow squirrel has the weight and size of an adult. The weight of sucha
specimen was 250g, 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 ontrees. After descending to the
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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 of Chenopodium. 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
female 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 ina 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 animal. 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.
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376
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.
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 wel! 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
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377
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
19 Sept. female
Be pinning): 24 Sept. male
9 Sept. female
13-15 Sept. female
; 9 Oct, male
pesca 19 Oct. male
31 Oct. male
End . 1-4 Nov. female
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
B27,
378
379
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 1/2-
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
Months
December
Juvenis Molt begins in the End of molt End of molt in
second half of the late litters
month
Subadultus IMolt begins in the 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 ends at Presumably, end
last days of the
month
about the 20th
of the month
of molt in
females born 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" (Periodichnost' 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
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,
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380
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., inthe
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.
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 of Nucifraga caryocatactes
macrorhynchos to Europe. Migrations of Nucifraga c.
macrorhynchos to Europe make it possible to predict exactly reduced
* Summer migrations of squirrels are usually caused by drought or forest fires.
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381
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 Regionin August. The Siberian squirrel population can thus be
predicted 2-21/2 months before the squirrel hunt.
Loxia curvirostra, Loxia pytyopsittacus and Loxia
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, 100km long in 1917 along
the Northern Dvina and 300km 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 ina locality. The rate of migration is 3-4km/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 andI.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. Accordingto 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 1km 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
12km 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 250-300km 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.
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382
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 ararity. 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 keeptotheir 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. Iokhel'son (1898), hunters in the Kolyma Territory state
that squirrels are attacked by flying squirrels. lIokhel'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,
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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. F
The most serious enemy of the squirrel is the goshawk. It is also hunted
by sparrow hawks, kites andvarious 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. Inthe 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 1-1.5m 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.
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385
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,1I., Mlekopitayushchie Moskovskoi gubernii (Mammals of the
Moscow Province), pp. 265-281.
1925, Sciurus vulgaris varius Turov S,S,,Mlekopitayushchie Ryazanskoi gubernii (Mammals of the
Ryazan Province), p. 59.
1927-1928. Sciurus vulgaris ognevi 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.31.5mm); length of upper tooth row 8.3-9.9mm
(M.9.2mm); hind feet 54-61mm (M. 57.8mm); tail 160-190mm
(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
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
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334
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
HIGUBE Tes) svullct and cheeks reddish rust-buff of varying intensity.
Sours! Tahestts Occiput with darker brownish tones like in the
Hoag a eee Mi middle of the back. Flanks and legs more pure
ete hiaseot motes) 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 giving a black-chestnut tone to the dorsal side ofthe tail. Middle of ventral
side of tail dirty rust-gray. Sides of ventral areawitha fringe of reddish-
rust and black-chestnut tones. Margins oftail witha 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-cinnamonandtawny). 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 (onthe Belaya river)
the boundary meets that of S.v. bashkiricus. The eastern boundary
turns north along the Kama to Perm.
386
335
387
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 OgnevS,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 LOCALITY 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.oOgnevi.
MEASUREMENTS: Length of skull 51.6-53mm; condylobasal length
48-49mm; zygomatic width 31.7-32.5mm; length of upper tooth row 9.6-
10.5mm; length of hind foot 57mm.
DESCRIPTION. Squirrels of the dark type predominate. Back dirty-
gray (between mouse- gray and deep mouse-gray, occasionally witha neutral
gray shade), darker than in darkS.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 of S.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 of S.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.
388
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 OgnevS.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 inS.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 of S.v.ognevi.
MEASUREMENTS. Length of skull 49-52.3mm (M.51.2mm); condylo-
basal length 43-48.7 mm (M.46.3mm); zygomatic width 28.3-32.2mm
(M.31.2mm); length of upper tooth row 9-10mm (M.9.2mm); 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 amongS.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 tail 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 onthe belly. To this type
3
sie belongs specimen No. M.4242, male, of my collection (caught by
33m
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 withawhite 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: 1) 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% inS.v.ognevi).
GEOGRAPHICAL DISTRIBUTION. S.v.formosovi‘eccurs 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 PeninsulaS.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 varius thantoognevi. Along its southern
boundary the range of S.v.formosovi passes gradually into that of
Ss, View Omran 1.
362. Sciurus vulgaris varius Barrett-Hamilton (1899).ex Kerr
(1792). Kola squirrel
1762, Sciurus varius Brisson A,D., Regnum Animale in classes 1X, Lugduni Batavorum, p. 106,
1789, Sciurus vulgaris varius Gmelin in Systema Naturae Linnaei I, ed. XIII, p. 146.
1792, Sciurus vulgaris varius KerrK., 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 Végel 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,I,, Faune des Mammiféres d'Europe, pp. 117-118; Miller
G.S. Catalogue of the Mammals of Western Europe. London, 1912, pp. 906-907.
1928. Sciurus vulgaris varius Serebrennikov M., Synopsis of Russian Squirrels, Doklady Akaemii Nauk
SSSR, p. 421,
TYPE LOCALITY AND DEPOSITION. Brisson, who described Sciurus varius 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.
Summer fur of adult specimens saturated, red-chestnut-rust colored.
Length of skull 49-53.6 mm; condylobasal length 44-48.6mm; zygomatic
width 30.2-31.6mm; length of upper tooth row 9-9.8mm.
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. Inthe 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 of S.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, especiallysoles, withgrayhair. Tail
gray-black, without anyrust shades. Hair roots oftaildarkash-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 of S.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: 1)
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: 1) lighter bluish gray color
of winter fur; 2) large percentage of red-tailed specimens (30% as
against 2-3% inS.v.formosovi); 3) large number of red-eared
Specimens which are very rare in winter specimens of S.v.formosovi.
I compared Swedish specimens of S. v. varius (from near Upsala) in
the ZMAN collection withS. v. vulgaris and found that its summer fur
is very dark, the tail chestnut colored. The back of Swedish specimens is
339
392
dark chestnut, warm-sepia (pl. XXIX, Ridgway, 1912). The tail is still
darker and the flanks more rust colored (with mikado-brown tinge, pl. 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 ofWestern 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 gubernii (Periodic Phenomena in the Life of Mammals, Birds and Reptiles of the Voronezh
Province), p. 379.
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 gubernii (Mammals of the Orel
Province) [according to recent data], p. 6.
1923, Sciurus vulgaris fuscoater OgnevS.I., Fauna nazemnykh pozvonochnykh Voronezhskoi gubernii
(The Fauna of Land Vertebrates of the Voronezh Province), pp. 153-154,
1924, Sciurus vulgaris fuscoater Migulin A. A., Mlekopitayushchie Khar'kovskoi gubernii (Mammals
of the Kharkov Province), ''Priroda i okhota na Ukraine", No, 1-2, pp, 119-128.
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 LOCALITY 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
48.2-50mm; zygomatic width 29.2-32.6mm; length of upper tooth row
10.1mm; hind feet 59-61 mm; tail 160-200mm.
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
393
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 onback, legs andtail, 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.
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. Inthe 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 of S.v.ukrainicus.
364. Sciurus vulgaris kessleri Migulin (1928). Kessler's squirrel
1840, Sciurus vulgaris Nordmann A., Observations sur la Faune Pontique; in: A. Demidoff, Voyage
dans la Russie méridionale, III, pp. 54-55.
1850. Sciurus vulgaris Kessler K.F., Zhivotnye gubernii Kievskogo uchebnogo okruga (Animals of
the Provinces of the Kiev District), pp. 38-40.
1858. Sciurus vulgaris Belke G., Esquisse de 1'Histoire Naturelle de Kamienietz-Podolski, Bulletin
Naturel de Moscou, p, 26.
1877, Sciurus vulgaris TacznowskiL., Liste des Vertébrés de Pologne, Bulletin Zoologique de France,
p. 129,
1915, Sciurus vulgaris Sharleman E,V., Mlekopitayushchie okrestnostei Kieva (Mammals of the
Vicinity of Kiev), Izdatel'stvo Kievskogo ornitologicheskogo obshchestva, pp. 50-51.
1925-1926, Sciurus vulgaris fuscoater KhranevichV., Narys fauny Podillya (A Survey of the Podolian
Fauna), pp. 21-22,
1927-1928. Sciurus vulgaris kessleri Migulin A. A., Obzor gryzunov Ukrainy (A Survey of
Ukrainian Rodents), '"Zakhist roslyn", Nos. 3-4, p. 12.
1930.-S (ciurus) v (ulgaris) kessleri andS. v. ukrainicus Danilovich A. P,, O belkakh
okrestnostei Kieva (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, inthisrespect, 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 inS.v.ukrainicus: Lengthofskull 51-
55.2mm (M.53.7mm); condylobasal length 48-52.8mm (M. 49.5 mm);
342
395
zygomatic width 31-34.3mm (M.32.5mm); length of upper tooth row 9.4-
10.2mm (M.9.8mm); hind feet 53-65mm (M.58mm); tail 180-205mm
(M.188 mm). e
DESCRIPTION. Winter fur of two types: 1) light type. These
specimens closely resemble the rust-brown type of S. vv. ukrainicus
Migul. (see above). S.v.kessleri, however, differs fromS.v.
ukrainicus inthe darker color of the ears. The ears of S.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 and Prout'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 and Prout's brown). Hair ontail 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 ofhead and
back of dark squirrels saturated red-chestnut, 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.ukrainicus
andS.v.kessleri differ in series in a number
of characters (see diagnosis).
The most constant character is the color of the
ears. Those of S.v.kessleri are usually
much darker than those of S.v.ukrainicus.
A.P. Danilovich (1930) recorded both S.v.
kessleri (5 specimens) andS.v.ukrainicus
(2 specimens) near Kiev. This author probably
considered light specimens of S.v.kessleri as
SUSUINE Mase elanlll co ukrainicus. I examined a large series of S.
SE RA GES v.kessleri and did not find a single
kessleri Migulin, No. M. rhe :
3704) troll the formey Kiev ukrainicus amongthem. All specimens from
PiSviaceE the left-bank Ukraine proved to belong to typical
kr aan ews.
Series of S.v.fuscoater 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 of S. v. fuscoater which
shows that the winter fur of S.v.fuscoater differs from that of S.v.
kessleri as follows: 1) 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 ofS.v.
fuscoater chestnut, without any reddish brown tinge, with a gray tinge
343
and a black tail. I never found black-tailed specimens of S.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
squirrel
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 LOCALITY 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 of S.v.ognevi: Length of skull 51-54.7mm
(M.53.2mm); condylobasal length 47.3-48.7mm (M.48.2mm); zygomatic
width 31.2-34.2mm (M.33.2mm); length of upper tooth row 9.2-10.2mm
(M.9.7mm); hind feet 51.3-57mm (M.52.8mm); tail 164-208mm
(M.185.6 mm).
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.
396
344
397
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 witha
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
Seba ah at 1S.
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 inthe north. This
region is inhabited by a type intermediate betweenS.v.bashkiricus
andS.v.ognevi.
365'. Sciurus vulgaris bashkiricus natiouralensis Ogn. (1935).
Ural mountain squirrel
1874, Sciurus vulgaris Sabaneev L. P., Pozvonochnye srednego Urala (Vertebrates of the Central Urals),
pio:
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 I]'menskogo
zapovednika (A Survey of the Mammalian Fauna of the Il'menskii Reserve), pp. 26-28.
1935. Sciurus vulgaris bashkiricus natiouralensis 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, 46,
TYPE LOCALITY AND DEPOSITION. No. M. 1650, sex unknown, December 1926, Miass of the
Zlatoust District (former County of the Orenburg Province), I1'menskie Mountains, S. I. Snigirevskii.
MATERIAL EXAMINED: holotype and 13 paratypes.
DIAGNOSIS. Closely relatedtoS.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 of S.v. bashkiricus, but more chestnut-
yellow, without the rust-red tones characteristic of S. v. bashkiricus.
Measurements like those of the Bashkirian squirrel.
SYSTEMATIC NOTES. Closely related toS.v.bashkiricus,
transitional to West Siberian squirrels. The winter fur differs from that
345
of S.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
?1792, Sciurus vulgaris argenteus 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
nérdlichen 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 gubernskogo muzeya, XXIV, pp. 32-33.
1928. Sciurus vulgaris nadymensis Serebrennikov M., Synopsis of Russian Squirrels, Akademiya
Nauk SSSR, p. 422,
TYPE LOCALITY. Sciurus martensi was described by P, Matschie from the left bank of the lower
Yenisei.
This so-called "species", which is in fact a subspecies of S. vulgaris, wasdescribed on the basis of
A. Middendorf's work "Sibirische Reise", Saugethiere, Végel 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 gew6hnliche Farbung der Eichhérnchen am unteren Jenisej, zwischen der
Stein- Tunguska und Polarkreise, zumal auf dem linken Ufer des Stromes, ist ein helles Grau, welches um
eine, nur bei dem genauesten Vergleiche in die Augen fallende, Tinte heller ist als das Grau der baltischen
Eichhdrnchen, Bei ndherer Untersuchung findet sich dann, dass das Grau der Jenisej Eichhérnchen 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 verrdath. "
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 angefiihrt, die Balge dunkler..."
M.K. Serebrennikov's assumption that the description of martensi 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: 119 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. Ifa 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 51-55.8mm (M.52.4mm); condylobasal length 46-51.2mm
(M.47.2mm); zygomatic width 29.5-32.8mm (M.30.4mm); length of upper
tooth row 9-10mm (M.9.3mm); hind feet 52.1-61.1mm (M.57.9mm); tail
145-192 mm (M. 166.1 mra).
346
3929
400
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-yellow-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 Sciurus 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. 1. Dwigubski (1804) mentionedSciurus vulgaris albus,
which is in part a synonym of martensi, as the Ob is mentioned as one
of its habitats.
The confused synonymy of S.v. martensi withS.v. nadymensis
is discussed above.
The West Siberian squirrel is closely relatedtoS.v.bashkiricus
natio uralensis, and differs from it as follows: 1) lighter buff tone of
back in winter specimens; 2) more luxuriant fur; 3) large size.
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 Darkechestnutatatled acc. eiem- AN aicisuabe 3.7% (4)
iN, Crespo Anleel goad 6lbigls Alo Me ook eZ 93.5% (102)
typicallichestnut=tawed@ 6 cn is «eo 62% (68)
lishircnestnut tamed. tea cileuen elise etree 31.1% (34)
WEG ey seleiglerel 5 6 cosa 0 sydol a cub oN ol ONO orcKc 2.8% (3)
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:
Ty? Chestuut-tailed hii. a7 djsye)) shee) « © 8 OSs 72,2% (13)
typicalichestnut=tanled, 2°. cis) sy>.5\ ofet> 4 5.5% (1)
light chestout-tailed j6 36 oe <saraehen cuss 66.7% (12)
1 PO =F Ek Ts OE eeerae ee tecener o 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 andS.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:
Darkrtailed s, efgrcts pete 49%
Chestnut-tailed ..... 44%
Red-tailed)... 2 4\.,5,5 2k 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 inthe 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.
401
367. Sciurus vulgaris exalbidus Pallas (1778). Teleut
squirrel
1788. Sciurus vulgaris varietasexalbida 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 dem 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: 1) 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 Sciurus vulgaris exalbidus Pallas,
No. 333 female, Korosten forestry, West Siberia, G.K.Gol'tsmaier, Right,
skull of Sciurus 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 54.6-59.2mm (M.56.9mm); condylobasal
length 50-53.2mm (M.52mm); zygomatic width 33.1-36 mm (M. 34.9 mm):
length of upper tooth row 10-11.1mm (M.10.4mm); hind feet 56.1-65.5mm
(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 yellow-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
404
405
that of S. 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 andexalbidus
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 intrueS.v.exalbidus. 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, along the line connecting the
village of Karabinskoe with the estuary of the Kazh river (a left tributary of the
Biya). AccordingtoS.N.Korchev, the Teleut squirreloccurs insmall numbers
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 Sciurus 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
406
In September 1933 N.A. Bobrinskii and S.A. Severtsov caught two
Teleut squirrels* in an oak-birch forest, 50km 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 LOCALITY 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 Sciurus
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 ajuvenilecharacterinS.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 49.5-50mm; zygomatic width 32.0-
34.5 mam.!"’
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 (1-2):80-81. 1932). Kuznetsov writes: ''The second form
found in pine forests of the Kalba Range differs from the Altai
specimens as follows: 1) 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
squirre] (the latter is probably larger).
** This is not correct (see above).
353
407
369. Sciurus vulgaris jenissejensis Ogn. (1935). Yenisei
squirrel
1867. Sciurus vulgaris Middendorf A. Th., Sibirische Reise, Saugethiere, S. 78-83.
1880, Sciurus vulgaris Tel (in Nordenskjéld's book), Ekspeditsiya k ust'yu Eniseya (An Expedition to
the Estuary of the Yenisei), p. 178.
1920. Sciurus vulgaris Solov'ev D.K., Sayanskii promyslovo-okhotnichii raion (The Sayan
Commercial-Hunting District), pp. 255-257.
1924, Sciurus vulgaris Kozhanchikov L. andI., 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 ShukhovI.I., Obzor rasprostraneniya
okhotnich'ikh i promyslovykh mlekopitayushchikh v Eniseiskoi gubernii, 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) Zverey M.D., Obzor gryzunov yuzhnykh chastei
Minusinsko-Khakasskogo okruga (A Survey of the Rodents of the Southern Part of the Minusinsk-Khakassian
District), Sbornik Srednesibirskogo geograficheskogo 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 jenissejensis OgnevS.I., Sistematicheskii obzor belok (A Systematic
Survey of Squirrels), Byulleten' Nauchno-issledovatel'skogo instituta zoologii Moskovskogo
universiteta, No, 2, p. 47.
TYPE LOCALITY AND DEPOSITION. No. 75 male, 23 Oct. 1926, Lower Tunguska, Turukhansk
Territory, N.P. Naumov. Type in the MSZM collection.
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: 1) 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 51.3-53.7mm (M.51.9mm); condylobasal length 45.1-
49.2mm (M.47.2mm); zygomatic width 29.2-32.2mm (M.30.6mm); length
of upper tooth row 9.1-10mm (M.9.5mm); hind feet 56-62.5mm
(M.58.8mm); tail 165-191mm (M.172.2mm).
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 inS.v.martensi, as the inner parts
of the haunches and antebrachia are gray. Soles dark blackish gray.
354
408
409
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.
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 intheir light color, but differ from it in the smaller white
area on the belly. Inner sides ofhindlegs and antebrachia light gray with
a light buff-rust tinge.
Summer fur of three types: 1) 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 zone between
white color of belly and brown color of back. Tail of chestnut and dark-
red type (see above). 3) very light specimens. Color rich rust-red-ocher
(between tawny-russet and mikado-brown). Sides of neck with a light
yellowish ocher stripe. Tail of the true red-tailed type.
B.A. Kuznetsov (in litt., 1935) examined 146,464 pelts obtained by
hunters and found that the Yenisei squirrels fall into the following groups,
according to the color of the tail: 36% black tailed and dark tailed, 46%
chestnut tailed, and about 18% red tailed.
SYSTEMATIC NOTES. Dark specimens of the Yenisei squirrel differ
clearly from dark specimens of S. vv. martensi. Light specimens of the
Yenisei squirrel resemble light specimens of martensi, but can be
recognized by the smaller white area of the belly, dark inguinalregion and
dark inner sides of legs. The differences from S.v.jacutensis will
be given in the description of this subspecies.
The squirrels from Sayan-MinuSinsk are closely related toS.v.
jenissejensis. They differ from it only in their smaller size and less
luxuriant fur (especially on the tail). The Sayan squirrels are thus
apparently identical with the Altai form (see below).
GEOGRAPHICAL DISTRIBUTION. Along the left bank of the Yenisei,
reaching the line Krasnoyarsk - Irkutsk in the south. The eastern boundary
is not known, as the region of Kirensk, Vitim, Vilyui is inhabited by
squirrels which differ from S.v.jenissejensis but are not identical
with S.v.jacutensis (see below).
355
410
370. Sciurus vulgaris altaicus Serebren. (1928). Altai squirrel
1884. Sciurus vulgaris Nikol'skii A.M., Puteshestvie v Altaiskie gory (A Journey to the Altai
Mountains), Trudy Sankt-Peterburgskogo obshchestva estestvoispytatelie, XI, p. 174.
1899, Sciurus vulgaris KashchenkoN.F., Rezul'taty Altaiskoi ekspeditsii (Results of an Expedition
to the Altai), Tomsk, p. 60.
1928. Sciurus vulgaris martensi natioaltaicus Serebrennikov M., Synopsis of Russian Squirrels,
Comptes Rendus de 1'Academie de Sciences de 1'U. R. R.S., p. 422,
TYPE LOCALITY AND DEPOSITION. Altai Mountains, Kok-Su river, estuary of the Yamanuch river.
Type No. 10,075 in the ZMAN collection.
MATERIAL EXAMINED: 64 specimens,
DIAGNOSIS. Color very variable. The variations resemble those of
S.v.jenissejensis. Dark- and black-tailed Altai squirrels are darker
than dark S.v.jenissejensis. Other color types of the two subspecies
are practically identical. There are fewer red-tailed specimens than in the
Yenisei subspecies.
MEASUREMENTS smaller than inS.v.jenissejensis. Lengthofskull
49-53.1mm (M.50.9mm); condylobassal length 45-48.5mm (M. 46.4mm);
zygomatic width 28-31mm (M.29.8mm); length of upper tooth row 8.8-
8.9mm; hind foot 45-60mm (65 in one out of twenty) (M.53.5mm); tail
140-170 mm (M.157.4mm).
DESCRIPTION. Color of winter fur. The Altai squirrel resembles the
Yenisei form in the variety of its color types. It differs from it in its
darker color in series. Dark-tailed squirrels especially differ clearly.
Back of the typical dark-tailed Altai squirrel dark gray (deep mouse-gray)
with fine dark gray-chestnut mottling. Head dark gray (between deep
mouse-gray and dark mouse-gray). Feet blackish gray. Tail black,
sometimes with a light chestnut tinge. Roots of hair on tail dark gray (deep
mouse- gray).
Back of dark-tailed Yenisei squirrels much paler, of purer gray (between
mouse-gray and neutral-gray).
The chestnut- and red-tailed Altai and Yenisei specimens are very
similar, hardly distinguishable in series.
Summer fur. We distinguish three principal types of summer fur. 1)
most specimens are black. Color of back and flanks black or black-
chestnut. Chestnut shade more prominent on head and sometimes also in
middle part of back. White abdominal area small. 2) rust-chestnut
specimens. Color of back between fuscous and natal-brown. Flanks
grayer. Head usually grayer than back. Cheeks gray with light chestnut
tinge (hair brown). Feet black-chestnut. 3) light rust colored specimens.
Back and flanks uniformly rust-brown (approaching mikado-brown).
Muzzle, cheeks and chin markedly dirty gray (between hair-brown and
mouse-gray). Feet dark chestnut. Tail generally reddish rust-chestnut
with a marked black tinge. This red-tailed type is identical with the red-
tailed Yenisei squirrel. According to B.A. Kuznetsov (1932) about 50%
of the Altai squirrels are dark tailed, about 40% chestnut tailed and about
10% red tailed.
SYSTEMATIC NOTES. The Altai squirrel is closely related to the
Yenisei squirrel, of which it is perhaps only a natio. It differs from it
only as follows: 1) smaller size; 2) different percentage of color-types
(see above); 3) darker color of dark types.
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411
GEOGRAPHICAL DISTRIBUTION. Altai, Tarbagatai and Sayan
mountains to the line connecting Krasnoyarsk with the northern spurs of the
Kuznetsk Altai* in the north.
371. Sciurus vulgaris jacutensis Ogn. (1929). Yakut squirrel
1887. Sciurus vulgaris Bunge A., Berichte itiber die Expedition nach den Neusibir. Ins., S. 38.
1898. Sciurus vulgaris Iokhel'son V.I., Ocherk zveropromyshlennosti vy Kolymskom okruge (A Survey
of Commercial Hunting in the Kolyma District), Sankt Peterburg, pp. 60-68.
1900. ''Belka" [squirrel], Slyunin N. V., Okhotsko-Kamchatskii krai(The Okhotsk-Kamchatka Territory),
IS lp. 326.
1903. Sciurus vulgaris calotus AllenI.A., Report on the Mammals collected in Northeastern
Siberia, Bulletin of the American Museum of Natural History, Vol. XIX, pp. 134-136.
1911. Sciurus borealis Brass E., Aus dem Reiche der Pelze, $.583 (Lena; Nomen nudum’ ).
1926, Sciurus vulgaris calotus Ognev, S.I., Mlekopitayushchie severo-vostochnoi Sibiri
(Mammals of Northeastern Siberia), pp. 77-84.
1927. Sciurus vulgaris calotus GassovskiiG.N., Gilyui-Oldoiskii okhotnich'e-khozyaistvennyi
taion (The Gilyui-Oldoi Hunting District), Vladivostok, pp. 523-526.
1928. Sciurus vulgaris fuscorubens (partim') Serebrennikov M. (ex Dwigubski, 1804), Synopsis
of Russian Squirrels, Doklady Akademii Nauk SSSR, p. 423.
TYPE LOCALITY AND DEPOSITION. Village Mugan, 8km from Yakutsk. Type No. M. 2209 in
S. I. Ognev's collection.
MATERIAL EXAMINED: holotype and 24 paratypes.
DIAGNOSIS. The two following color types predominate among winter
specimens: 1) black-tailed — dark-colored specimens. Back blackish
gray, without mottling or brownish tinge. 2) chestnut-tailed. Back blackish
gray with an ash tinge. Percentage of red-tailed specimens small, between
1 to 20% according to locality. Color of back of red-tailed specimens in
winter varying from light ash-mouse-gray to darker mouse-gray with well
marked gray mottling. Summer color variable, between black (most
common) and black-chestnut and even light chestnut. Winter fur fluffy and
luxuriant. White area of belly small.
MEASUREMENTS average. Length of skull 50-52mm (M. 51.6 mm);
condylobasal length 45-49.3mm (M.47.2mm); zygomatic width 29.2-32.2mm
(M.31.1mm); length of upper tooth row 9.1-10.2mm (M.9.8mm).
DESCRIPTION. Dark-tailed and black-tailed specimens predominate in
winter.
1) Black-tailed specimens: Middle of back dark grayish black with a
dark chestnut tinge (between deep mouse-gray and dark quaker-drab).
Flanks more ash colored (deep neutral-gray). Color uniform as there is no
dark mottling. Head dark ash-gray, similar to that of back. Chin and
lower parts of cheeks blackish gray (deep neutral-gray to deep purplish
gray). White area of belly very small as the dark color of the flanks
reaches far down. Groin blackish gray. White color of underparts forms
a short wedge anteriorly which reaches to the middle of the neck. Feet
blackish gray (dusky purplish gray). Soles with a gray-chestnut tinge
(hair-brown). Tail black (between soot-black and dusky purplish gray).
Roots of long hair on tail bluish-gray (dark mouse-gray).
2) Chestnut-tailed specimens are usually dark colored. Their color
is similar to that of black-tailed specimens. The back has a more marked
ash shade (neutral-gray). Dorsal side of tail of such dark chestnut-tailed
specimens almost black. The ash-gray roots are more marked than in
* [Obviously a mistake in the Russian original, should be Kumetsk Ala-Tau, ]
357
412
black-tailed specimens. Hair of ventral side of tail with two gray-buff
bands or one rust-chestnut-gray ring before the broad black hair tip. The
color of the hair on the tail is complex, i.e., roots deep ash-gray,
followed by a narrow blackish gray band, a little marked blackish band, a
rust-buff-gray subapical band and broad black hair tip. In lighter chestnut-
tailed specimens the dorsal side of the tail is generally also black. A
broad stripe formed by gray hair tips in the middle of the ventral side of
the tail. The gray-rust-buif subapical bands of the hair are little marked,
much less than in West Siberian and European chestnut-tailed specimens.
Tail of Yakut chestnut-tailed specimens even darker than in western dark-
tailed specimens.
Bald spine present in dark chestnut-tailed specimens and also in black-
tailed ones.
3) Winter fur of red-tailed specimens. Color of back varying from light
gray (light mouse-gray) to mouse-gray and light neutral-gray with gray
mottling. Ears rust-chestnut (between mikado-brown and tawny). Tip of
ears in many specimens darker, ash-chestnut with a light reddish tinge
(fuscous with brown tinge).
Color of summer fur variable. There are specimens with a black back
and a black or blackish gray tail and specimens with generally dark
chestnut color (between bone-brown and warm sepia), witha brownish tinge
in the middle of the back. Ears and feet black-chestnut. Rust-buff-
colored hair on feet. Color of tail similar to that of back. Color of hairs
on tail as follows: roots dark mouse-gray (deep mouse-gray), followed by
broad dark chestnut bands (warm sepia) with a light rust tinge. Hair tips
black-chestnut. Hair roots on ventral side of tail light ash-gray (mouse-
gray, sometimes light mouse-gray), alternating with short dark chestnut bands,
followed by a rust-chestnut band, a dark chestnut band, a broader rust-
colored band and chestnut-black apices.
In the lightest chestnut-tailed specimens, summer fur on back sometimes
rich, rust colored. Tail similar.
SYSTEMATIC NOTES. In his survey of Russian squirrels, M.K.
Serebrennikov (1928) suggests that the Okhotsk Coast, the Anadyr Territory,
and the lower reaches of the Amur up to the Stanovoi Range in the east are
inhabited by S.v.fuscorubens Dwigubski. In ''Prodromus Faunae
Rossicae" (1804) p.83, Dwigubski describes fuscorubens as follows:
'Hieme argenteo-griseus, duplo major communi". This description is
obviously wrong. Dwigubski gives the distribution of the species as ''In
Siberia orientali" (p.86). This information is certainly not reliable, and
fuscorubens should be considered synonymous with the three East
Siberian subspecies: jacutensis, anadyrensis andrupestris.
In old publications Siberian squirrels (especially the Yakutian squirrel)
were sometimes calledS.v.calotus. This name was, according to
some authors, given by B.H. Hodgson (On a New Species of Mustela?
known to the Nepalese commerce as the Chuakhal Mustela? Calotus nob. ,
Calcutta, Journal of Natural History, II, 1842, pp. 221-223). Other
authors think this name was given by J. Gray (''Synopsis of the Asiatic
Squirrels", Annals and Magazine of Natural History, Vol. XX, series 3,
1867, p. 272).
In his article on ''Mustela calotus'" Hodgson wrote that ''Nepali
merchants" imported gowns trimmed with the fur of a beautiful animal
named ''Chuakhal'', from an unknown locality in the north.
358
413
Hodgson obtained a skin of this animal without skull. He could not
determine to which order the animal belonged. However, it resembled a
squirrel and in Hodgson's opinion was a species of Mustela. The
description of this skin resembles that of the dark Siberian squirrel. On
the basis of the merchants' reports, Hodgson assumed that this mysterious
animal was distributed in the Himalayas and Tibet.
At my request V.P. Uvarov examined the holotype of Mustela
calotus inthe collection of the British Museum. The label of the specimen
bears the name ''Nepal'', which was subsequently emended to ''Thibet''’ by
O. Thomas. According to Uvarov it is without doubt a squirrel. Hodgson's
article contains a plate of a very strange animal with the muzzle of a marten
and the tail and ears of a squirrel.
As Sciurus vulgari'’s does not occur in the Himalayas, Tibet and
Nepal, Mustela (?) calotus Hodgson is obviously a dark specimen of
S.vulgaris without any definite type locality. The Nepali merchants
apparently obtained skins of dark Siberian squirrels. Hodgson's description
was apparently based on one of these skins.
I was surprised that O. Thomas (''Proceedings of the Zoological
Society of London", 1905, p. 346) considered the mountains of Central
Asia (including the Altai) as the terra typica of S. v. calotus, and that
Sowerby accepted this.
The name Mustela (?) calotus cannot be used for the Siberian
squirrel for the following reasons: 1) the description is vague; 2) the
squirrel is placed inthe genus Mustela with a question mark; 3) the range
given is not correct.
But as there is a type specimen of a Siberian squirrel in the British
Museum called Mustela (?) calotus, and as Gray refers to Hodgson's
work in his description of S. calotus, it is correct to name the Siberian
squirrelS. vulgaris calotus Gray (1867) ex Hodgson (1842).
The terra typica ofthis subspecies is givenby J. Gray (1867, p. 212) as
follows: ''Hab. North. China, Thibet (Mr. Hodgson's type), Siberia."
This form does not occur in Tibet, and ''Siberia'' is too general. The
type locality of Gray's S.v.calotus is therefore northern China which
he mentions first. This interpretation makes it necessary to consider S.
v.chiliensis Sowerby (1921) described from the Chihli* Province
(75 miles northwest of Peking) as a synonym of S.v.calotus Gray.
The Yakut squirrel differs from the Yenisei squirrel as follows: 1) more
luxuriant and silky fur; 2) darker color of back in winter fur. It differs
from the Transbaikal subspecies (see below) as follows: 1) smaller size;
2) more luxuriant and silky fur. It differs from S.v.rupestris (see
below) as follows: 1) more luxuriant fur; 2) smaller size.
The Anadyr squirrel is closely related to the Yakut squirrel, perhaps
forming a natio of this. The differences between these two forms are given
below.
B.A. Kuznetsov (in litt., 1935) examined 94,920 pelts of S.v.
jacutensis and gives the following data on color types of the tail; 16%
black-tailed; 34% dark-tailed; 32%dark chestnut-tailed; 18% red-tailed.
In my collection are color variants of squirrels from the Aldan river and
from the upper reaches of the Indigirka. The labels of these specimens
bear remarks based on information from local fur-trading companies by
K.E. Vorob'eva (who placed this material at my disposal).
* [Now Hopei Province, ]
359
According to this information there are fewer red-tailed specimens then
given by B.A. Kuznetsov.
The figures given by K.E. Vorob'eva are as follows:
414
Anadyr squirrels
Black-tailed.. < . pdadiddt. tepduchaniaeh jl meee hh eld 49 %
Daskretalled) oe sie ig ce es Ecce as lane eae 34%
Chestnut-tailed (dark specimens) PORE Bee eae 7%
Chestnut-tailed (light specimens)........... 3%
REUHUAME a ston tememel ee erence wreath ear ente ts 6%
Squirrels from the upper reaches of the Indigirka
BISCK@tallle ds —sciwucitomey oA iate) sce Thy). oa) as (0) ate haan 40 %
PARR =tare dy crac ia, chee kart late ois sys) ion sr reetoneae te 39%
CHERENUE LALOR eet ee or eis eo ecole te ey amt oe te one 19%
Red-tailed (dark specimens with bald spine)... 0.1 — 0.2%
Redwtdiledit. |.) cede aeetels suakysh. Bail) ae. 1%
According to B.A. Kuznetsov (1932) the Lena squirrels differ from the
Yakut squirrels. The Lena squirrels occur in the Vilyui and Vitim districts
of Yakutia, and in the Kirensk, Irkutsk, and Tulun districts of the Siberian
Territory. According to Kuznetsov, there are many different color types:
black-tailed, dark-tailed, chestnut-tailed and red-tailed specimens. Series
generally contain: about 11% black-tailed specimens, 23% dark-tailed,
49% chestnut-tailed and 17% red-tailed ones. These data are based on
212,458 skins. The color of the black-tailed specimens is similar to that
of the Yakut squirrels, but the fur of the Lena strain is not so bluish and
dark. Color of tail lighter and more drab.
Color of dark-tailed specimens lighter than that of black-tailed ones
and color of spine lighter. Color of chestnut-tailed specimens still lighter.
Red-tailed specimens of the Lena strain differ from Yakut specimens
in the weak rust tones on the spine. Tail and tufts on ears rich rust
colored.
In order to determine whether there is a separate Lena race (or more
correctly a natio) specimens from the Vilyui-Vitim districts on the left
bank of the Lena should be compared with specimens from the right bank
of the Lena.
GEOGRAPHICAL DISTRIBUTION. It is difficult to determine the
boundaries of the range of this subspecies for lack of material. It
apparently occurs in the upper reaches of the Lena, along the Vilyui,
Muna. This is correct if we consider the Lena squirrel as identical with
the Yakut subspecies. It also occurs along the Olenek. According to R.
Maak (1886) it is very rare in this region, near 68°N. It occurs along the
Aldan, Khanym, Indigirka, Yana (A. Bunge, 1887), Kolyma and Omolon
rivers. This subspecies apparently also includes squirrels from Penzhin-
Kamchatka. The southern slopes of the Stanovoi Range and the Dzhugdzhur
Range are inhabited by the Okhotsk-Sakhalin subspecies. It is not clear which
subspecies occurs in the Gilyui-Ol'doi hunting territory. According to
G.N. Gassovskii (1927), squirrels are very common in pine forests in this
region and it is possibly inhabited by a subspecies either closely related to
or identical with the Yakut subspecies, or perhaps transitional to the
Okhotsk-Sakhalin subspecies.
360
415
416
371. Sciurus vulgaris jacutensis natioanadyrensis Ognev
(1929). Anadyr squirrel
1883. Sciurus vulgaris Nordquist O,, Anteckingar och. studier till Sibirska ishafskustens
Daggdjursfauna, p. 80.
1927, Sciurus vulgaris calotus Sokol'nikov N. P., Okhotnich'i i promyslovye zveri Anadyrskogo
kraya (Game and Commercial Animals of the Anadyr Territory), Byulleten’ Moskovskogo
obshchestva ispytatelei prirody, XXXVI, pp. 134-136,
1929, S(ciurus) v(ulgaris) jacutensis natioanadyrensis OgnevS.I., Uber einige Nagetiere
Ost-Sibiriens. Zoologisch. Anzeiger, Bd. 83, H. 1/4, S. 83.
1935. Sciurus vulgaris anadyrensis OgnevS.I., Sistematicheskii obzor belok, vodyashchikhsya
v SSSR (A Systematic Survey of the Squirrels of the U.S.S.R.), Byulleten' Nauchno-issledovatel'skogo
insitituta zoologii Moskovskogo universiteta, Vol. II, p. 48.
TYPE LOCALITY AND DEPOSITION. No. 4449, Anadyr Territory, N. P. Sokol'nikov. Holotype in the
Zoological Museum of Moscow University.
MATERIAL EXAMINED: holotype and 23 paratypes.
DIAGNOSIS. Closely related to the Yakut squirrel. Winter fur differing
from that of the Yakut squirrel as follows: 1) more marked dull chestnut
tinge on back, often forming well marked bald spine; 2) more marked
chestnut or chestnut-gray mottling which is almost absent in Yakut
squirrels; 3) more luxuriant fur.
MEASUREMENTS. Lengthof skull 50.3-54 mm (M. 52.4 mm); condylobasal
length 45.6-48.5mm (M.47.2mm); zygomatic width 30.2-31.8mm
(M.31.1mm); length of upper tooth row 8.8-10.1mm (M.9.2mm); hind
feet 50-62mm (M.57.1mm); tail 145-180mm (M. 162.8 mm).
ADDITIONAL DESCRIPTION. Color of winter fur closely resembling
that of S.v.jacutensis. Unfortunately, I had only one summer pelt
of the Anadyr squirrel at my disposal. Back dark-chestnut. A narrow
red-ocher stripe (ochraceous tawny-tawny) on flanks, at the limit between
the dark dorsal area and the white belly. Tail of the dark* type.
In the Zoological Museum of the Moscow University are red-tailed
squirrels caught by N.P. Sokol'nikov in the Anadyr Territory. He
apparently collected rare color types. N.P. Sokol'nikov (1927) calls the
red-tailed squirrels in winter fur ''color variants'' and writes that he has
several such specimens.
Color of back of red-tailed squirrels of the Anadyr Territory in winter
dirty buff-gray (light drab) with a buff-rust tinge (wood-brown, sometimes
with a rust tinge). Bald-spine specimens occur. Back generally with
chestnut-gray mottling.
In the ZMMU** is a collection of aberrant specimens of S.v.
jacutensis natioanadyrensis. Two of them are true albinos, and
three are flavescent, pale buff-yellow with buff-gray-chestnut tails.
SYSTEMATIC NOTES. Closely related to the Yakut squirrel, but the
color of this is purer and more bluish than that of the Anadyr squirrel and
without chestnut shades. Bald-spine is rare among Yakut squirrels.
GEOGRAPHICAL DISTRIBUTION. Larch forests of the Anadyr Territory
(called in Russian ''listvyaga''). One ofthese forests begins 50-60km above
the village Markovo and stretches for 170km along the Anadyr river.
Another forest follows the Mainariver, at its left tributaries (the Algans). This
* N. P, Sokol'nikov (1927) describes the Anadyr squirrel as follows: 'In summer the squirrel is reddish,
resembling the European squirrel but darker... "'
4* [ZMMU — Zoological Museum of the Moscow University, ]
361
forest is much smaller than the former. These forests are narrow, not
wider than 1-3km. N.P. Sokol'nikov (1927) writes: ''The forests of the
Anadyr area are separated from the Siberian forest belt. This squirrel
is therefore isolated and localized. It cannot go anywhere and its
population cannot increase by immigration of squirrels from other areas."
372. Sciurus vulgaris fusconigricans Dwigubski (1804).
Transbaikal squirrel
1804. S(ciurus) vulgaris fusconigricans Dwigubski J., Prodromus Faunae Rossicae, pp. 83-84.
1862. Sciurus vulgaris Radde G,, Reisen im Siiden von Ost-Sibirien. S, 133-146.
1910, Sciurus vulgaris Kashchenko N.F., Okollektsii mlekopitayushchikh Zabaikal'ya (A Collection
of Mammals from Transbaikalia), Ezhegodnik Zoologicheskogo muzeya Akademii Nauk, Vol. XV,
pp. 273-274,
1922, Sciurus vulgaris varius Dybovski Ben., Arch. Tow. Naukow, Lwowie, III, p. 8.
1928. Sciurus vulgaris fusconigricans Serebrennikov M., Synopsis of Russian Squirrels, Doklady
Akademii Nauk, p, 423.
1929. Sciurus vulgaris Kumetsov B,A., Zverovye promysly vostochnogo Zabaikal'ya (Animal
Commerce in Eastern Transbaikalia), Trudy po lesnomy opytnomu delu, Tsentral'naya lesnaya opytnaya
stantsiya, Issue VI, pp. 87-89. )
1935. Sciurus vulgaris fusconigricans OgnevS.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, Vol. II, p. 48.
TYPE LOCALITY. Barguzin, Transbaikalia ("ad Bargusin'' according to Dwigubski). Holotype lost.
MATERIAL EXAMINED: 16 specimens.
DIAGNOSIS. Back in winter specimens dark bluish gray with fine, well
marked gray-black mottling in black-tailed and dark-tailed specimens.
Color of chestnut-tailed specimens lighter and grayer. Specimens with
black or black-chestnut back predominate among summer specimens.
Dark chestnut specimens are common. Dark brownish chestnut specimens
(red-tailed ones) are apparently much less frequent. Winter fur not as
luxuriant and fluffy as in the Yakut squirrel. White abdominal area small.
MEASUREMENTS large. Length of skull 52-55.8mm (M. 54.5 mm);
condylobasal length 48-52mm (M.50.4mm); zygomatic width 31-33.2mm
(M.32.9mm); length of upper tooth row 9.2-10.2mm (M.9.6mm); hind
feet 51-60 mm (M.54mm); tail 160-185mm (M.175mm).
DESCRIPTION. Winter fur usually very dark, deep gray (between
deep neutral-gray and deep mouse-gray) with fine well marked grayish black
mottling. This distinguishes the subspecies from dark S.v.jacutensis
which has no such mottling. Top and sides of head dark gray with well
marked mottling. Ears black. Feet blackish gray. Tail of the black or
dark type. Chestnut-tailed specimens of the dark type are rare.
In chestnut-tailed specimens of the dark type'the back is much lighter
than in dark- or black-tailed specimens. Back dirty chestnut-gray, nearly
mouse-gray. White abdominal area markedly reduced.
The color of summer fur shows three types. To the first type belong
squirrels of black or black-chestnut color and with black or dark tails.
The second type has the following characters: back deep dark chestnut
(between bone-brown, natal-brown and fuscous); flanks grayer; ears and
tail black-chestnut; tail of the dark-chestnut type.
The third type is rare and consists of red-tailed squirrels. Back dark
chestnut-brown (between walnut-brown, russet). Flanks grayer (natal-
brown). Anterior part of head and cheeks with well marked grayish
362
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jo Inj JautuIns — ainBtz raddy
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Inj JaqUIM — oinB1z tamoy ‘adAy 1OT[OOT ey
‘soul, SNOTINYOIULU sTIEB[NA Snintod¢) [euinbs uermyoue~w oy |,
IIA ALV1d
363
chestnut tones. Tail brown-chestnut with mouse-gray hair roots. Feet
chestnut-brown.
According to B.A. Kuznetsov (in litt. 1935), Transbaikalian squirrels
usually include about 38% black-tailed specimens, 26% dark-tailed, 33%
dark chestnut-tailed, and only about 3% red-tailed specimens. These data
are based on 51,108 skins.
SYSTEMATIC NOTES. S.v.fusconigricans differs from S.v.
jacutensis as follows: 1) larger size; 2) less luxuriant winter fur;
3) dark mottling on back of dark-tailed and black-tailed winter specimens.
It differs from the Manchurian squirrel (see below) as follows: 1) smaller
size; 2) more intense color of back in winter specimens; 3) presence of
red-tailed specimens.
GEOGRAPHICAL DISTRIBUTION. Widely distributed in Transbaikalia,
along the Barguzin, Upper Angara, Vitim, Olekma, and Shilka rivers, on
the Muiskii Range, andinthe Chita and Verkhneudinsk* districts. According
to B.A. Kuznetsov (1929), it is not found in isolated pine forests along the
Onon river. It occurs in larch forests on the Yablonovyi Range. M.K.
Serebrennikov (1928) records this subspecies from forests near Lake
Khubsuguland from Northern Mongolia. To judge from my material, this
region is inhabited by a form more closely related to the Altai squirrel,
or transitional between the Altai and Transbaikalian squirrels. Such
transitional specimens occur on the northwestern spurs of the Khingan.
They are intermediate between S.v.fusconigricans and the larger
Manchurian squirrel.
373. Sciurus vulgaris mantchuricus Thos. (1909). Manchurian
squirrel
1858, Sciurus vulgaris Schrenck L., Reisen und Forschungen im Amur-Lande, I, S, 118-124 (partim!).
1859. Sciurus vulgaris Maak R., Puteshestvie na Amur, Obzor zhivotnykh (An Expedition to the
Amur. A Zoological Survey), Sankt Peterburg, p, 103 (partim' ).
1870, Sciurus vulgaris Przheval'skii N. M., Puteshestvie v Ussuriiskom krae (A Joumey to the Ussuri
Territory), Sankt Peterburg, pp. 255-256.
418 1909. Sciurus vulgaris mantchuricus Thomas O,, Annals and Magazine of Natural History, ser. 8,
Vol. IV, p. 501.
1912, Sciurus vulgaris mantchuricus OgnevS.I., O kollektsii mlekopitayushchikh iz
Ussuriiskogo kraya (A Collection of Mammals from the Ussuri Territory), Ezhegodnik Zoologicheskogo
muzeya Akademii Nauk, Vol. XVI, pp. 500-502,
1923. Sciurus vulgaris mantchuricus Sowerby A.de C., The Naturalist in Manchuria, Vol. II-III,
p. 146,
1928. Sciurus vulgaris mantchuricus Serebrennikov M., Synopsis of Russian Squirrels, Doklady
Akademii Nauk, p. 423.
1935. Sciurus vulgaris mantchuricus OgnevS.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 universitata, Vol. II, p. 49.
TYPE LOCALITY AND DEPOSITION. Great Khingan. Holotype in the British Museum.
MATERIAL EXAMINED: 30 specimens.
DIAGNOSIS. Back in winter specimens (Plate VIII) intense gray with
fine, well marked dark mottling. Color generally lighter than in Transbaikal
squirrels. There are sometimes bald-spine winter specimens with a light
chestnut tinge on the lower part of the back. Most specimens are black- or
dark-tailed. Dark chestnut-tailed specimens are rare. Red-tailed
* [Now Ulan-Ude. ]
364
419
specimens do not occur*. The summer specimens show four color types.
Most common are specimens with black or black-chestnut back. The other
two are tricolored squirrels which occur more rarely: 1) squirrels with
dull brown-gray back and rust-red lateral stripe; 2) specimens with
saturated chestnut-brown back and narrow, gray lateral stripes. Rust-
brown-ocher squirrels are very rare. Most summer specimens are black-
tailed. Lighter specimens include dark-tailed and rarely dark-chestnut-
tailed specimens. Fur of winter specimens shorter and less luxuriant than
in Transbaikal specimens.
MEASUREMENTS large. Length of skull 53-57 mm (M. 55 mm); condylobasal
length 48.3-51.3mm (M.50mm); zygomatic width 30-33.7mm (M. 32mm);
length of upper tooth row 9-10.1mm (M.9.8mm).
DESCRIPTION. Winter fur. Back gray (between neutral-gray and
mouse-gray), generally lighter than inS.v.fusconigricans. Back with
well marked dark gray mottling. Ears and feet dark gray. Tail of the
black or dark type. Dark chestnut-tailed specimens are rare. Bald-spine
specimens are also rare. They have a light chestnut tinge caused by
chestnut-white hair tips and light chestnut bands before the black subapical
bands of the hair on the lower part of the back.
Summer fur shows four types. To the first type belong squirrels with
black or black-chestnut back. This color varies between deep chestnut-
black and brilliant black. Drab tones are occasionally present on cheeks
and around the eyes. Flanks of the same color as the back. Ears and feet
black. Tail of the black type, rarely of the dark type.
The second color type consists of tricolored squirrels (Plate VIII, upper
figure). Dorsum dull brown-gray (between natal-brown and warm-sepia).
Area around eyes, sides ofmuzzle and cheeks buff-drab (hair-brown) with
fine chestnut-gray mottling. A-rich ocher-red-rust stripe (orange -
cinnamon) on the flanks between the brownish gray back and the white belly,
from the neck to the inner sides of the legs. Outer sides of legs, shoulders,
antebrachia, haunches and tibiae of the same color as the back. Inner
sides of legs ocher-red-rust. Feet brownish gray-chestnut. Dorsal side
of toes chestnut colored. Tail of the dark type, black in some specimens
and rarely of the dark chestnut type.
The third type also includes tricolored squirrels. Head, back, outer
sides of legs and feet chestnut-brown (argus-brown with chestnut tinge).
Dirty gray (mouse-gray) with chestnut mottling predominates on the sides
of the muzzle, cheeks and around the eyes. A narrow gray stripe (deep
mouse-gray) on the flanks between the chestnut-brown back and the white
underparts, from the sides of the neck to the inner sides of the legs. Tail
of the dark type. Black color of root of tail sometimes with a well marked
chestnut-brown shade.
The fourth type of summer fur includes squirrels with the following
colors: Back rich rust-brown-ocher (between tawny and russet). Muzzle
lighter and yellower. Region around the eyes grayer. Forehead of the
same color as the back. Occiput with well marked chestnut-gray tones.
A narrow yellow-rust stripe (cinnamon-buff) on the flanks between the
rust-brown-ocher back and the white underparts from the sides of the neck
to the inner sides of the legs. Feet rust-brown-ocher, closely resembling
that of back. Tail of the dark type, rarely of the dark-chestnut type. This
* Or are very rare. I never saw such specimens.
365
420
color type is rare in summer specimens. Black or black-chestnut summer
specimens predominate, constituting 80% of the population. Tricolored
specimens are less frequent, and light specimens are very rare.
SYSTEMATIC NOTES. The differences betweenS.v.mantchuricus
andS.v.fusconigricans have been given above. The Manchurian
squirrel differs from S.v.rupestris as follows: 1) larger size; 2)
smaller percentage of bald-spine winter specimens; 3) complete absence
of red-tailed specimens.
GEOGRAPHICAL DISTRIBUTION. Manchuria, northern part of the Kirin
Province (de Sowerby, 1923), on the Great Khingan (O. Thomas, 1909), in
the Ussuri and the Maritime territories, to about 48-49°N. Squirrels
from the Lesser Khingan and Northern Korea apparently belong to this
subspecies.
374. Sciurus vulgaris rupestris Thos. (1907). Sakhalin-Amur
squirrel
1861, Sciurus vulgaris Schmidt Fr., Bemerk. iiber die Sdugethiere Siid Sachalins etc., Mélange
Biologique de 1'Académie Impériale de Saint-Pétersburg, Vol. IV, p. 107.
1889, Sciurus vulgaris Nikol'skii A.M., Ostrov Sakhalin i ego fauna pozvonochnykh zhivotnykh
(The Vertebrate Fauna of Sakhalin Island), pp, 123-125.
1907. Sciurus vulgaris rupestris Thomas O., The Duke of Bedford's Zoological Explorations in
Eastern Asia, IV, Proceedings of the Zoological Society of London, Vol.I, p. 410.
1923. Sciurus vulgaris rupestris Sowerby A.de C., The Naturalist in Manchuria, Vols. II-III,
p. 147.
1928. Sciurus vulgaris rupestris Serebrennikov M., Synopsis of Russian Squirrels, Doklady
Akademii Nauk SSSR, p. 424; OgnevS.I., Mlekopitayushchie Shantarskikh ostrovov (Mammals of
the Shantar Islands), Izvestiya Tikhookeanskoi nauchno-promyslovoi stantsii, II1(5): 15 et seq., 1929.
1929, Sciurus vulgaris dulkeiti OgnevS.1L., Uber einige Nagetiere Ostsibiriens, Zoologischer
Anzeiger, Bd, 83, H, 1/4, S. 76-85.
TYPE LOCALITY AND DEPOSITION. The Korsakov Station on Sakhalin Island. Holotype in the
British Museum.
MATERIAL EXAMINED: 100 pelts, most of them with skulls.
DIAGNOSIS. Color very similar to that of S.v.mantchuricus.
Winter fur more luxuriant and percentage of bald-spine specimens higher.
Black or black-chestnut specimens predominate among summer specimens.
Black- and dark-tailed specimens are most common. Dark-chestnut
Specimens are rare. Red-tailed specimens extremely rare. *
MEASUREMENTS smaller thanthoseofS.v.mantchuricus. Length
of skull (Figure 187) 50.6-55.3mm (M.52.6 mm); condylobasal length
46.1-50.3mm (M.47.9mm); zygomatic width 28.3-32.5mm (M. 30.8 mm);
length of upper tooth row 9-10.2mm (M.9.7mm); hind feet 56-67mm
(M.59.8mm); tail 165-206mm (M.184mm).
ADDITIONAL DESCRIPTION. Color of winter fur closely resembling
that of the Manchurian squirrel, generally more luxuriant. Bald-spine
specimens are apparently more frequent.
* B, A. Kuznetsov (in litt. 1935) gives the following figures with regard to tail color in the Amur strain:
25% black-tailed, 34% dark-tailed, 35% dark-chestnut-tailed and 6% red-tailed, These data are based
on 53,287 skins. The Amur strain includes squirrels from the Amur Region, the Ussuri Territory,
Sakhalin and the Shantar Islands. According to B. A. Kuznetsov, red-tailed specimens of the Amur strain
occur only in the middle reaches of the Amur.
366
421
Black and chestnut-black specimens apparently predominate in summer.
Percentage of black- and dark-tailed specimens very high. Chestnut-
colored specimens occur together with black summer specimens. Back of
chestnut-colored specimens deep chestnut-brown (warm -sepia) with a black
tinge and chestnut-black mottling. Roots of guard hairs and hair of
undercoat gray (neutral-gray). Hair tips of undercoat dull rust colored.
Gray roots of guard hair followed by a black or chestnut-black band, a
rust-brown band (near mikado-brown) and black or black-chestnut hair tips.
There are also longer hairs with gray roots and black tips. Brown-rust
tone more marked on the head than on the back. Forehead, region around
the eyes and cheeks with a marked gray tinge (near deep mouse-gray).
Feet black or black-chestnut. A wide blackish gray stripe between the
chestnut-brown-black back and the much reduced white abdominal area
from the sides of the neck and along the flanks. This stripe causes the
tricolored fur, with chestnut-brown back, grayish black flanks and white
underparts.
Tail of most summer specimens of the black or dark type. In my
collection is a young specimen with a peculiar tail color (No. M. 3796,
Sakhalin, Argi-Pagi, 13 July 1932, A Rekst). Tail dull yellowish drab-
rust. Roots of guard hairs dull buff-gray and hair tips gray-rust. There
are also hairs with broad black-chestnut tips. The combination of these
colors creates the dull yellowish drab-rust color.
Red-tailed specimens are very rare, e.g., from the Goryun river (a
tributary of the Amur), and a specimen from the Great Shantar Island.
Back of these specimens rust-ocher-chestnut, similar to color of tail.
Only one red-tailed specimen was found among 160 Amur squirrels (not
counting the specimen from the Great Shantar Island).
SYSTEMATIC NOTES. In 1929 I described the subspecies S.v.
dulkeiti from the Shantar Islands (''Zoologischer Anzeiger", loc. cit,
1929). On the basis of published data (O. Thomas, M.K. Serebrennikov)
I thought that S.v.rupestris from Sakhalin Island is a small insular
form which differs from the Middle Amur form. After examination of
squirrels from Sakhalin I found that this was wrong. S.v.rupestris
is not smaller than the Shantar and Middle Amur squirrels; it is
identical with them.
S.vV.rupestris is closely related toS.v.mantchuricus from
which it differs in its smaller size.
GEOGRAPHICAL DISTRIBUTION. Sakhalin and Shantar islands and the
Amur Territory, to about 48-49°N in the south. Farther southS.v.
mantchuricus occurs. The exact boundary between these two closely
related subspecies is not known. There may be no sharp boundary. South
of 48-49 N the size of the squirrels becomes larger. Between Bira and
Bidzhan rivers there occur specimensofs.v.mantchuricus withlarge
skulls.
Subgenus Tenes Thos.
1778. Sciurus Gmelin J, , Systema Naturae Linnaei, editio XIII, Vol.I, p.148(S.anomalus),
1811, Sciurus Pallas P. Zoographia Rosso-Asiatica, 1, pp. 186-187 (S.caucasicus).
1909. Tenes ThomasO. The Generic Arrangement of the African Squirrels, Annals and Magazine of
Natural History, III(8):468(S. persicus; footnote),
1935, Oreosciurus OgnevS.I., Sistematichekii 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, pp. 50-51,
367
Species of this subgenus differ from true Sciurus in the following
characters: 1) color not varying with the seasons; 2) belly not white like in
true squirrels but rust-chestnut of various shades, or buff-yellow; 3) hind
soles with six calluses (S. vulgaris has only four); 4) teats 10
(S. vulgaris has 8); 5) one pair of upper premolars (S. vulgaris
has two pairs); 6) Structure of crowns of premolars differs from that of
S.vulgaris (see below); 7) shape of os penis differs from that of true
squirrels (see Figure 196).
Type species: Sciurus persicus Erxl.
GEOGRAPHICAL DISTRIBUTION. Transcaucasia, Asia Minor, Syria,
Palestine and part of Iran.
422
165. Sciurus (Tenes) persicus anomalus Gmel.(1778).
Caucasian squirrel
1778. Sciurus anomalus Gmelin J., Systema Naturae Linnaei, editio XIII, Vol.I, p. 148.
1811. Sciurus caucasicus Pallas P,, Zoographia Rosso-Asiatica, I, pp. 186-187 (''frequens est in
fagetis et quercetis subalpinis et montanis Caucasi... "').
1840. Sciurus caucasicus, Nordmann A., Observations sur la Faune Pontique; A. Demidoff, Voyage
dans la Russie Méridionale, III, p.55; Simashko Yu., Russkaya fauna (Russian Fauna), II, 1851,
p. 593; Bogdanov M.N., Etyudy russkoi okhoty, Okhotnich'i i promyslovye zveri i ptitsy Evropeiskoi
Rossii (Studies on Hunting in Russia. Game and Commercial Mammals and Birds in European Russia),
Okhota i konnozavodstvo, Nos. 5-6, 1874, p, 121.
1843. Sciurus russatus Wagner J. A. , Die Sdugethiere von Schreber, Suppl. III, Erlangen, S.155, 156
(without indication of locality; presumably the Asian part of Turkey. The description of the color
corresponds to that of anomalus),
1884, Sciurus persicus (partim') Lataste F., SurSciurus persicus, Le Naturaliste, pp, 1-4.
1920. Sciurus anomalus SatuninK.A., Mlekopitayushchie Kavkazskogo kraya (Mammals of
Caucasia), Vol. II, pp. 49-64.
1928, Sciurus anomalus, Serebrennikov M. , Synopsis of Russian Squirrels, Doklady Akademii Nauk,
p. 425.
NAMES: The Caucasian or Georgian squirrel (English); das Transkaukasische Eichhérnchen (German);
l'Ecureuil de Caucase, 1'Ecureuil anomal (French).
LOCAL NAMES: tritina, tria (Georgian); tsikvi, tsiku (Kakhetia); abkhidkha (Abkhazian); kovkasi
skyur (literary Armenian); mkrosh (vernacular Armenian); garmzy sychan (Kurds along river Tertera).
TYPE LOCALITY AND DEPOSITION. Transcaucasia (without exact indication of locality), Type lost(?).
PRINCIPAL FIGURES: 1) Schreber J., Saugethiere, Erlangen, IV, 1792; 2) Simashko Yu., Russkaya
fauna, II, Mlekopitayushchie (Russian Fauna, I]. Mammals), 1851, pl. 32, fig. 1 (in color); 3) Vinogradov
B.S., K poznaniyu gryzunov Zakavkaz'ya (The Rodents of Transcaucasia), Zapiski Kavkazskogo muzeya,
A(3) (damaged nuts); 4) SatuninK.A., Mlekopitayushchie Kavkazskogo kraya (Mammals of Caucasia),
Vol, II, 1920, p. 48, figs. 130-131 (skull ventral and dorsal); p. 51, fig. 132 (skull lateral); p. 53, fig. 133 (molars).
MATERIAL EXAMINED: 10 specimens,
DIAGNOSIS. Back chestnut-gray-buff with blackish chestnut mottling.
Lower part of back of many specimens (especially in winter fur) black-
chestnut. Flanks chestnut-brown. Underparts rust-yellow-brown. Tail
chestnut-brown-rust. Terminal part of tail in some specimens witha
black band before the ocher-yellow tip (plate IX).
424 Length of skull 50-53mm; condylobasal length 46.3-49mm; zygomatic
width 28-31mm; length of upper tooth row 9.5-10mm; hind feet 52-56mm;
tail 130-159 mm.
ADDITIONAL DESCRIPTION. The skull of the Caucasian squirrel differs
from that of S. vulgaris as follows: 1) coronal suture with two pointed
processes of the frontals; coronal suture of S. vulgaris rounded or nearly
straight, sometimes with two rounded processes; 2) frontal suture longer
368
423
FIGURES 190-193, Skull of Caucasian squirrel, Sciurus (Tenes) persicus anomalus Gmel. ,
No, M.2079, 24 Jan, 1927, village Dzhar of the Zakataly District, M. V. Shidlovskii, S.I, Ognev's
collection
Drawing by Yu. A. Kostylev
369
424 than that of S. vulgaris; 3) zygomatic arches narrower (see Figures 189-
195).
FIGURES 194, 195, Skull of the Caucasian and common squirrel
Left — skull of Sciurus (Tenes) persicus anomalus Gmel. (ventral), No. M, 12078,
S.I. Ognev's collection. Right — skull of Sciurus vulgaris kessleri Migulin, No, M.3704,
male, 25 Nov. 1927, former Kiev Province
Photograph by S. I. Ognev
S. persicus andS.vulgaris differ markedly in the structure of
the teeth. The first upper premolar is absent in the Caucasian squirrel.
The second upper premolar is larger and more rounded than inS. vulgaris
in which it is bevelled off anteriorly to some extent. Inner tubercle of
second upper premolar better developed and more rounded than inS.
vulgaris. First lower premolar with two anterior tubercles which are
425 higher and more rounded than the more widely-separated tubercles of
S.vulgaris. Incisors shorter and weaker than in the common squirrel.
Os penis. The baculum of the Caucasian squirrel is larger and more
curved distally than that of S. vulgaris (see Figures 171-172 and
Figure 196). Os penis of S.anomalus 11mm long and its diameter at
the base 2.6mm; inS.vulgaris length 8.1-9 and width at the base 2.1-
2.2mm.
Color. In winter specimens color of back of two types: 1) back chestnut-
gray-buff with blackish chestnut mottling; 2) anterior half of back of the
same color and the lower part of back, from loin to base of tail brilliant
black-chestnut with light whitish buff mottling. Color of hairs on back as
follows: Hair roots deep gray (hair-brown) followed by a narrow black-
chestnut band, a pale whitish buff band and chestnut-black hair tip. Some
longer guard hairs with black-chestnut tips scattered throughout the fur.
370
426
In squirrels with black-chestnut lower part of back, hairs in this region
are colored as follows: hair-roots deep gray (chaetura-drab), followed by
a narrow black-chestnut band, a
whitish band and black-chestnut tip.
1 Forehead, upper parts of cheeks and
occiput rust-chestnut with black-gray
mottling (between sayal-brown and
mikado-brown). Eyes surrounded by a
light buff-white-gray ring. Flanks
2 and shoulders chestnut-brown (between
hazel and russet).
Dorsal side of legs of the same color
FIGURE 196. Os penis of the Caucasian squirrel as the back. Outer side of hind legs
I Jaterethl 2 Sl doraln(Highly cnapaified), rust-brown (sayal-brown with tawny
Drawing by Yu. A. Kostyley tinge).
Soles buff-yellow- gray, between
cinnamon-buff and avellaneous. Each hind sole with an irregular black
spot along the inner margin. Ears of the same color as the soles, with
small terminal tufts, 10-12 mm long.
Underparts from chin to groin saturated rust-yellow-brown (between
cinnamon-buff and cinnamon).
Dorsal side of tail chestnut-brown-rust (between russet and mars-
brown). Tip of tail ocher-yellow (ochraceous-tawny) with a black
subapical band in some specimens. This dark band is caused by black
hair with ocher-yellow tips at the tip of the tail. Middle of ventral side of
tail pale rust-yellow or yellowish drab. Ventral side laterally more
intense rust-red. Hair of margins of ventral side of tail with black
subapical bands and pale yellowish tips.
Color of summer fur little different from that of winter fur. Back in
summer generally lighter, brown, and without the black tone of winter
specimens. Underparts of summer specimens usually of more intense
color with more marked rust tone.
Soles of forefeet with five well developed calluses. The two posterior
caluses are more rounded and higher than the three anterior calluses at
the base of the toes. One large, elongate callus at the outer margin. One
rounded, small callus near the inner margin. Soles of forefeet almost bare
Hind soles of winter specimens with sparse. short hair. Claws of forefeet
shorter than in true squirrels (Figures 197, 198).
Beats 10: pil-ieeva.2-27e 2-2: total 10. In Scars oar 1 Ss
p. Li a. 2-2:7 1. lle “totaliee
SYSTEMATIC NOTES I placed the Caucasian, the Persian and Syrian
squirrels in one subgenus, Oreosciurus, in1935. I had unfortunately
overlooked a footnote in O. Thomas! paper on African squirrels (op. cit. ,
1909, p.469): ''S. persicus with only 4/4 cheek teeth, but similar to
true Sciurus inall other characters, should form a special subgenus,
which might be called Tenes.'' This quotation shows that Thomas took
into account only the reduced number of upper premolars, ignoring a
number of important characters which distinguish this peculiar group of
squirrels. The name Tenes has to be adopted because of priority, in
spite of the incomplete and superficial diagnosis given by Thomas.
During examination of a series of Caucasian squirrels I noticed that, in
Abkhazian squirrels, the color of the flanks is a saturated red-rust
371
- (mikado-brown-orange-cinnamon, pl. XXIX), the belly intense ocher
(cinnamon) and the groin shows yellowish spots (cinnamon-buff, pl. XXIX).
Tail intense chestnut-red (russet, pl. X) with a rust tinge at the tip
(mikado-brown-orange-cinnamon). Back with marked black tone and buff -
yellow mottling.
The Abkhazian squirrels are possibly a western race of the Caucasian
squirrel with more saturated color, but this is unlikely as there are
squirrels in Artvin which differ very little from squirrels in Central
Transcaucasia.
The Persian squirrel, Sciurus (Tenes) persicus Erxl., is
closely relatedto anomalus. I examined seven specimens from the
vicinity of Balkh near the Irano-Turkish border (collected by Nesterov)
and one specimen from Iran (collected by N.A. Zarudnyi) in the ZMAN
The summer fur of the Persian squirrel is light, gray-buff-rust, grayer
and lighter than pinkish buff (pl. XXIX Ridgway, 1912). Top of head more
ocher, (near pinkish cinnamon, pl. XXIX). Cheeks, throat, chin, flanks,
belly and feet buff-yellow with a light rust tinge (pinkish buff, pl. XXIX).
Tail dorsally ocher-rust (cinnamon-buff) with yellowish hair tips
(cinnamon-buff). Median ventral line of tail pale yellowish rust (pinkish
buff). Sides of ventral area of tail more rust colored (cinnamon-buff).
427
FIGURE 197. Ventral side of forefoot of FIGURE 198. HindfootofSciurus (Tenes)
Sciurus (Tenes) persicus anomalus persicus anomalus Gmel.
Gmel,. , No, $,1352, Murav Dagh Photograph by S. 1. Ognev
Winter fur on back generally dirty drab with marked whitish mottling.
Skull of the Persian squirrel very similar to that of anomalus. Coronal
suture without the posterior processes characteristic of anomalus.
Number of upper molars as inanomalus.
372
I examined two Syrian squirrels (Sciurus (Tenes) persicus
syriacus Ehrenb.), one from Syria and the other from Palestine. The
animal closely resembles the Caucasian squirrel. Color of underparts
lighter and yellower. Tips of hair on tail yellowish. Mottling on back in
winter fur more whitish than inanomalus. The Syrian squirrel is
intermediate between anomalus andpersicus. This is well shown by
the Palestinian specimen in the ZMAN.
428 In my opinion, squirrels of the group persicus-anomalus should
be considered as one species with several subspecies. This would also
correspond to the zoogeographical data. The Persian squirrel should
include the following subspecies: Sciurus (Tenes) persicus
persicus Erxl. (Gilan Province), S. (Tenes) persicus anomalus
Gmel. (Transcaucasia), S. (Tenes) persicus syriacus Ehrenb. *
(Syria, Palestine), S. (Tenes) persicus fulvus Blanf. (Shiraz,
Southern Iran).
The color ofS. (Tenes) persicus anomalus Gm. is the most
saturated, and its range is the most northern. The subspecies becomes
markedly lighter towards the south. Syriacus and fulvus are especially
light.
GEOGRAPHICAL DISTRIBUTION. Widely distributed in Transcaucasia,
along the southern slopes of the eastern part of the Great Caucasus,
throughout the Little Caucasus and in Western Transcaucasia (inthe Artvin
District). K.A. Satunin (1920) thinks that the range corresponds with his
zoogeographical division of Caucasia, that is the ''forest district of Eastern
Transcaucasia''. According to more recent data, S. persicus
anomalus also occurs in the ''West Transcaucasian"' district to use K.A.
Satunin's terminology. S.p.anomalus is absent in a large part of the
forest district of Eastern Transcaucasia in its southeastern and northeastern
parts. It does not occur on the Northern Caucasus. This is discussed in
detail by E.A. Biichner ("Bulletin de 1'Académie Imperiale de Saint-
Petersbourg'', Nouvelle série, I (XXXIII) 1890, pp. 125-132). All reports
of the occurrence of squirrels on the NorthernCaucasusrefertoGlis glis
which is called "squirrel" or ''small squirrel'' by the local population.
The knowledge of the distribution of squirrels in Western Transcaucasia
is very incomplete. G.I. Radde wrote on the distribution of squirrels in
Svanetia: ''Squirrels do not occur frequently here. In 1864 I took ten days
to pass through the inaccessible ravine of the Ingur and Lakhamuli in
Dzhvara. Over 70 versts (Svanetia - Mingrelia) I saw a squirrel only once,
in a beech forest and not in a pine forest." (‘Museum Caucasicum, '' 1899,
1, p: 62)
K.A. Satunin (1920) writes on the distribution of the squirrel in the
Sukhumi District: ''On the way from Sukhumi to Andreevskoe in 1913, my
son saw a squirrel near the road but did not succeed in catching it. He
may therefore have been mistaken.'' K.A. Satunin does not give any
information on the distribution of squirrels in Abkhazia. There are now
specimens from Abkhazia in the ZMAN, from the southern slope of the
ravine from Sukhumi to the Pskhu range, from the Kodor District and also
from near the village Parneauty (collected by G.P. Adlerberg). Farther
* Sciurus historicus J, E. Gray is a synonym of this form (Synopsis of the Asiatic Squirrels, Annals and
Magazine of Natural History, 1867, p. 273).
373
429
FIGURE 199, Sciurus (Tenes) persicus
anomalus Gmel, Sketch from nature
Drawing by K. K. Flerov
1254
south along the Black Sea coast,
squirrels were observed near Ardanuch
by K.D. Deryugin (K.A. Satunin, 1920).
Specimens collected by P. V. Nesterov
near Artvin and Gurzhan, former Artvin
District are in the ZMAN collection.
A. Nordmann (1840) recorded this
species from Adzharia. We have no
reports from the Kutaisi District,
although it probably occurs there. It
is common from the Surami Range
eastward and occasionally in the former
Tiflis Province (K.A. Satunin, 1920).
He writes: "I once saw a squirrel in
a garden in Tiflis in Mikhailovskaya
Str., where, untilrecently, the gardens
along the Kura formed a nearly
continuous belt.
''In Kakhetia and the Zakataly District
the squirrel is common and sometimes
even numerous. My son caught two
specimens near Lagodekhi. The
squirrel is very common in central
Transcaucasia, in the western part
of Tiflis Province, along the eastern
spurs ofthe Meskhiiskii Range and the
mountains near it. I found large
numbers on the Borzhomi Estate
(villages Tori and Bakuriani) and in
Ardanuch. In the east its range
includes the forests at a similar height
on the northern and eastern slopes of
the Little Caucasus. I never found it
on the southern slopes (Zangezur
County) during two journeys in 1892
and 1912. Everywhere I found Glis
glis L., but no squirrels."
In the ZMAN collection are
specimens of Caucasian squirrels from
the vicinity of Zakataly and Lagodekhi
(collected by Mlokosevich). In my
collection are specimens from the
Zakataly District (collected by M. V.
Shidlovskii).
We have no exact data on the
distribution of the squirrel on the
Talysh, except for the contradictory
observations by G.I. Radde.
MODE OF LIFE. Beech forests
are the main habitat of the Caucasian
squirrel. It avoids pine forests with
underbrush, and prefers forests with
high trees.
374
431
432
The habits of the Caucasian squirrel resemble those of the common
northern squirrel. According to K.A. Satunin, the sight, hearing and sense
of smell are highly developed. Its choice of food indicates that its taste is
developed. According to Satunin, the voice of the animal is sharp, metallic
and sounds like ''chit-chit-chit". It gives voice immediately when in danger.
The squirrel is active throughout the day. It climbs rapidly on trees.
During stops it holds itself to the bark with the claws of all feet. The
Caucasian squirrel leaps from branch to branch making jumps of 3-5m.
The tail plays an important part, acting as a parachute. Satunin writes:
"T sometimes observed that when the animal arrived in an unknown place,
it climbed to the crown of a tree or onto a broken branch so that it could
have a wider view. It sat for a long time in the same place, surveying the
area as if considering its future actions."'
The Caucasian squirrel is an excellent swimmer but does not like to
enter water.
FIGURE 200, The Caucasian squirrel
Drawing by K. K. Flerov
It usually feeds on nuts, mainly the Persian walnut, chestnuts, acorns,
hazel and beech nuts, seeds of fruit, and berries. As it lives mainly in
beech forests, beech nuts are apparently its mainfood. Satunin writes:
"In Tiflis I once saw a squirrel on a tree with ripe apricots. The animal
tore away the juicy pulp and discarded it to reach the stone which it ate
like a nut."
According to K.A. Satunin the Caucasian squirrel collects large stores of
food for winter, mainly nuts. These are hoarded in hollows in trees,
clefts, burrows among the roots of trees, etc.
The Caucasian squirrel may rarely eat eggs in nests of small birds, and
even nestlings. Non-Russian scientists accuse the Russian common
squirrel of this. However, such cases are rare and isolated.
K.A. Satunin writes: ''As far as I know squirrels do not build nests in
Caucasia, but live only in hollows of trees. According to L. L. Mlokosevich,
squirrels near Lagodekhilivein elms (Ulmus), linden (Tilia) and maple
(Acer insigne), rarely inoak. The nests are built. of moss, dry leaves
and other material. The same hollow is used for many years. L.L.
Mlokosevich found squirrels for more than five years in the same elm
tree."
According to the literature, the Caucasian squirrel has 3-7 young.
L.L. Mlokosevich found only 3 young in the nests. According to
PLATE IX
The Caucasian squirrel, Sciurus (Tenes) persicus anomalus Gmel.
Drawing by V. A. Vatagin
376
K.A. Satunin (1920) gestation lasts 3-4 weeks, but I doubt this. By analogy
with the common squirrel, the gestation period cannot last less than a
month. According to K.A. Satunin, in Transcaucasia the young are born
in April.
bos
AVS
Drawing by K. K. Flerov
FIGURE 201. The Caucasian squirrel
The Caucasian squirrel is of little commercial value. Until the First
World War the animal was not hunted. According to B.A. Kuznetsov,
several hundred skins were collected between 1930-1932, but commerce
has much increased and up to 30,000 pelts are now collected per year
Subfamily SPERMOPHILOPSINAE
Animals living in sand deserts. Back of uniform color.
Ears very short forming only small folds around the openings.
Tail less than half the length of the body, covered with moderately long
hair.
Toes unusually long and slender. Secona and fourth toes of hind feet
longest and of about equal length. Second and third toes of forefeet of about
equal length, the third slightly longer than the second.
433
377
434
Claws black, very long. Longest claws usually longer than 1cm (up
to 1.5cm).
Cheek pouches not developed.
Crowns of molars inclined outwards, more or less rounded. Enamel
pattern of crowns complicated, lophodont. Crowns of upper molars with
transverse plicated loops, their rounded part directed inwards.
The surface of the molars also with two rounded-oval enamel areas,
the anterior larger and better developed than the posterior. Lower molars
with more bunodont structure.
Dentalformula of molars: p =; nay
3
Glans penis with a large carinate, laterally compressed process, its
tip inclined downwards (Figure 208).
The carinate tip of the glans is more strongly flattened at the right
side. It is laterally widened, and inclined to the left. The structure of
the glans penis of Spermophilopsis resembles that of Geosciurus
(Subfamily Xerinae) (Figure 209).
The geographical distribution of the subfamily has been given above.
The subfamily Spermophilopsinae is closely related to the African
subfamily Xerinae, from which it differs in cranial and morphological
characters.
The anatomy of Xerinae and Spermophilopsinae should be studied to
determine their relationship. A preliminary study of Spermophilopsis
and Xerus was recently made by B.S. Vinogradov (1936) who showed
that there is a close resemblance between these two genera. It would,
however, be premature to place Spermophilopsis inthe subfamily
Xerinae.
Genus Spermophilopsis W.Blasius (1884)
1883. Arctomys Lichtenstein H., Naturhist, Anhang in Eversmann's Reise nach Orenburg und Buchara,
Sete
1827-1834, Citellus Lichtenstein H., Darstellung neuer Saugethiere, Taf. XXXII.
1852, Spermophilus Brand J.F., Zool. Anhang in A, Lehmann's Reise nach Buchara und Samarkand,
S. 303,
1884, Spermophilopsis Blasius W., Uber eine bisher zu den Zieseln (Spermophilus) gerechn,
vermutl, neue Nagetiergattung aus Turkestan, Tageblatt der Natur. Versamml. zu Magdeburg, Sect.
Zool. , S. 324/5.
The species of this genus are adapted to-life in sand deserts.
Back ofuniform color, pale, resembling the background. Tail less than
half the length of the body, covered with dense fur.
Terminal part of tail black ventrally, the dorsal side black only at the
margin of the tip (Figure 210).
Ears very short, forming only small folds. Ear tufts never present.
Cheek pouches not developed. Vibrissae arranged as follows: 1) paired
groups of long vibrissae on sides of muzzle; 2) paired groups of soft, white
vibrissae at the upper lip; 3) at margin of lower lip; 4) paired groups
under the eyes; 5) one vibrissa near the outer corner of the eye; 6) paired
groups of four vibrissae on the cheeks behind and below the eyes; 7)
a single group on the chin; 8) groups of eight vibrissae at the outer ventral
margin of wrist; 9) one vibrissa on the outer side of antebrachium; 10) one
378
pair on the chest; 11) another pair posterior to group 10; 12) two
symmetrical vibrissae outside those of group 11; 13) symmetrical groups
of one or two vibrissae widely apart at the sides of the belly, laterally and
posteriorly to group 12.
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FIGURE 202, Measurements of skulls FIGURE 203. Measurements of skulls of
of Spermophilopsinae and Marmotinae Spermophilopsinae and Marmotinae
I — width of rostrum; II — length of I — condylobasal length; II — length of bony palate;
nasals; III — zygomatic width; IV — III — diastema; IV — length of upper tooth row;
temporal width; V — width of skull V — length of bullae osseae; VI — width of bullae
posterior to postorbital processes. osseae.
Drawing by Yu. A. Kostylev
Reats: paddle faZia2 lol roles total Ge
435 Posterior processes of premaxillaries wide. Occipital part of each
process as wide as both nasals in their posterior part.
Coronal suture rounded posteriorly, the middle part almost straight
with two irregular, angular lateral processes.
Interorbital space very broad, about one third of condylobasal length.
Anterior part of frontal area flattened and slanting anteriorly. Posterior
part slightly elevated with two small tubercles.
Brain case short and wide.
Anterior part of zygomatic arch at level of zygomatic process markedly
elevated and widened, like in squirrels.
Infraorbital foramina very small, laterally compressed as in squirrels.
Angular processes at the base of the infraorbital foramina.
Diastema about as long as upper tooth row.
Type species: Arctomys leptodactylus Licht.
GEOGRAPHICAL DISTRIBUTION. Northern Iran and Afghanistan,
Transcaucasia and sporadically in the Kyzyl-Kum and Kara-Kum. An
isolated area in the southern Balkhash Region.
379
436
166. Spermophilopsis leptodactylus Licht. (1823). Long-clawed
ground squirrel
1823. Arctomys leptodactylus Lichtenstein H., Naturhist. Anhang zu Eversmann's Reise nach
Orenburg und Buchara, S. 119.
NAMES: Das langkrallige Ziesel. Des diinnzehige Ziese] (German).
LOCAL NAMES: alaka (eastern Turkmenistan); alangart (western Turkmenistan); tishkan (Semirech'e,
Dzhety Su); karakuiryuk [black tail] (Khiva Oasis, according to M. N. Bogdanov),
PRINCIPAL FIGURES: 1) Lichtenstein H., Darstellung neuer Sdugethiere, Berlin, 1827-1834,
Taf. XXXII; 2) Bogdanov M.N., Ocherki prirody Khivinskogo oazisa (Description of Nature in the Khiva
Oasis), 1882, p. 26 (drawing of entire animal); 3) Fausek V., Biologicheskie issledovaniya v Zakaspiiskoi
oblasti (Biological Investigations in Transcaspia), Zapiski po obshchei geografii Geograficheskogo obshchestva,
Vol. XXVII, 1906, pp. 41-42, figs. 28-30 (structure of legs); 4) Bobrinskii N. A. , Opredelitel' okhotnich'ikh
i promyslovykh zverei (Key to Game and Commerical Animals), 1928, p. 44, fig. 28 (paws); second edition,
1932, p.56, fig. 37; 5) V.G.Heptner, O melkikh pushnykh vidakh Turkmenii (Small Fur-Bearing Animals
of Turkmenistan), Pushnoe delo, 1929, No. 8, pp, 58-64, fig. 1, p. 62 (running suslik), fig. 2, p. 63 (burrow);
6) Vinogradov B.S. and S,1I. Obolenskii, Vrednye i poleznye v sel'skom khozyaistve mlekopitayushchie
(Harmful and Useful Mammals in Agriculture), Moscow, 1932, p. 29, fig. 18 (drawing of entire animal from
photograph of dead specimen); 7) Kashkarov D.N., Zhivotnye Turkestana (Animals of Turkestan), 1932,
p. 16, fig. 4 (entire animal); 8) Lavrov N.P. and S.P. Naumov, Rasprostranenie i biologiya tonkopalogo
suslika v Turkmenskoi SSR (Distribution and Biology of the Long-clawedGround Squirrel in Turkmenia),
Zoologicheskii zhurnal, XII(2): 81, fig, 1, 1933(entire animal, sketch from photograph in nature); 9)
Vinogradov B. A., Mlekopitayushchie SSSR. Gryzuny (Mammals of the U.S.S.R. Rodents), Leningrad,
1933, pl. IV, fig. VII (photograph of entire animal); 10) Serebrennikov M. K., Album osteurop. ,
westsibirisch. und turkestanisch. Sdugetiere, III, Ztschr. f. Sdugetierkunde, Bd. VIII, 1933, Taf. III, Alb. 5
(entire animal); 11) Lavrov N.P. and S$, P.Naumov, Stroenie mekha i lin'ka tonkopalogo suslika (Fur and
Molt inS permophilopsis), Zoologicheskii Zhurnal, XIII(2): 313, 1934. fig. 1 (hair structure);
pp. 314-315, 317, 318, 323-324, figs. 2-9 (structure and arrangement of hair); p, 326, fig. 10 (molt);
12) Vinogradov B.S., Pavlovskii E.N. and K.K. Flerov, Zveri Tadzhikistana (Animals of Tadzhikistan),
1935, pp. 202-203, figs. 72-73 (entire animal, photograph of burrow); 13) Vinogradov B.S., Argyropulo A, I.
and V.G.Heptmer, Gryzuny Srednei Azii (Rodents of Central Asia), 1936, p. 23, fig. 1 (photograph of entire
animal), p. 24, fig. 2 (head of young), p. 26, fig. 4 (skull), p. 27, fig. 6 (penis), p. 28, figs, 7-8 (teeth),
p. 29, fig. 9 (photograph of habitat),
MATERIAL EXAMINED: 161 pelts and skulls,
DIAGNOSIS. Interorbital space very broad, more than 30%, of
condylobasal length. Skull very broad posterior to postorbital processes.
Width of skull at this level more than 40% of condylobasal length.
Zygomatic width more than 75% of condylobasal length. Posterior part of
nasal processes of premaxillaries very wide. Occipital part of nasal
processes as wide as both nasals in their posterior part or a little
narrower. Interparietal large and of varying shape, with well defined
border. Bullae osseae very large and inflated. Their anterior part
divided from posterior parts by a well marked transverse groove. Around
the auditory meatus they form tubular processes. Length of upper tooth
row less than length of bullae osseae. Crowns of molars rounded and with
folds of the lophodont type. Diameter of first upper premolar 1/3 to 1/4 of
that of second premolar (Figure 204-207).
Toes very long and slender. Second to fourth toes of hind legs longest
and of about equal length; first toe long, only little shorter than fifth.
Second and third toes of forefeet of about equal length and longer than the
others; third toe little longer than second; first toe very small with well
developed claw; fifth about half the length of the fourth. Claws very long,
and black. Longest claws usually more than 1cm long (to 1.5cm).
Summer fur short, coarse, stiffand sparse, of yellowish buff-sand color,
sometimes witha light rust tinge (intermediate between light pinkish cinnamon
and cinnamon-buff, pl. XXIX). Fine gray-chestnut mottling sometimes present.
380
iota 4
FIGURES 204-207. Skull of Spermophilopsis leptodactylus Licht. No. M.3214 male, 4 May 1925.
Sands near Ashkhabad, S.1. Ognev's collection
Drawing by Yu. A, Kostylev
381
437
438
Winter fur (Plate X) verydelicate, brilliant, silky. It color varies between
rich buff-rust-sand tones and more rust-drab tones (light pinkish cinnamon,
pinkish buff, cinnamon-buff, pl. XXIX). Fine drab-chestnut mottling
present to a varying extent in winter fur of the different subspecies.
MEASUREMENTS. Length of skull 50-59.2mm; condylobasal length
44.2-54mm; zygomatic width 31-38.6mm; interorbital space 13.4-
20.1mm; length of upper tooth row 9.8-12.2mm. Body length 210-278 mm;
tail length 43-102mm; hind foot 49-67mm.
Teats 8.
DESCRIPTION. The skullof Spermophilopsis resembles that of
the squirrel, having a similarly wide interorbital space and area posterior
to the postorbital processes. The same cranial characters distinguish
Spermophilopsis from Citellus. The longitudinal diameter of the
last upper molar is about half as long as the distance between the posterior
margin of its alveolus and the anterior margin of the mesopterygoid fossa.
The mandibula resembles that of the squirrel. It differs from that of ground
squirrels in its short and massive angular process. In Citellus the
angular process is lower, its dorsal margin has a large notch and it is more
inclined downwards, sidewards and backwards. The interparietal of
Spermophilopsis is present as a separate one evenin very old specimens.
In squirrels and ground squirrels this bone is present only in very young
specimens.
The skullofSpermophilopsis differs from that of the squirrel
as follows: 1) postorbital process more posterior; 2) nasals narrower and
less widened anteriorly; 3) posterior part of nasal processes of
premaxillaries rounded; inSciurus vulgaris they have outer lateral
processes.
Crowns of molars inclined outwards and rounded. Enamel pattern of
crowns lophodont. Upper molars with transverse plicated loops the rounded
part of which is directed inwards. Surface also with two rounded oval
enamel areas, the anterior better developed and larger than the posterior.
Lower molars bunodont, especially the anterior premolars. The teeth of
Spermophilopsis closely resemble those of African species of the
genus Xerus (cf. M. Weber, Sdugetiere, B. II, 1928, S.257, Figure 172,
Tih, WV).
Color of summer fur on back and head light, uniformly buff-sand
colored with a rust tinge (near cinnamon-buff, light pinkish cinnamon,
pl. XXIX, sometimes with a yellowish, light ochraceous-buff tone, pl. XV).
In specimens from Kushka the rust tone is more saturated and the hairs have
better marked chestnut-brown bands which produce a fine mottling. Color
of flanks lighter and yellower than that of back. Legs buff-sand colored.
Soles dorsally paler and more whitish. Chin, throat and belly white. Hair
in these areas white to the roots. Dorsal side of tail buff-rust-sand colored.
Many hairs on the sides of the tail with blackish subapical bands and buff-
yellow apices. The black subapical bands become broader and more
intensely colored in the terminal part of the tail, where they form a
subterminal black area with a whitish border. Ventral side of basal third of
tail buff-sand-gray, remaining two thirds of tail black with a whitish fringe.
Claws very long, black-chestnut, strong, slightly curved, with yellowish
tips. Ears very short, consisting of short folds around the auditory
meatus.
382
440
Winter fur sharply different from summer fur. Hair of underfur in
summer very short, barely visible. The fur consists mainly of stiff, short
guard hairs (i.e., guide hairs, guard
hairs and intermediate hairs). The
winter fur consists mainly of the
underfur which is very soft and
straight, wavy only on the hindquarters.
Guard hairs of winter fur straight,
stiff and elastic. The guide and guard
hairs of the winter fur are more than
twice as long as those of the summer
fur. N.P. Raspopov and S.P. Naumov
(1934) state that in winter the hair of
the underfur on the back is nine times
as long as that of the summer fur. On
the other hand the guide hairs and guard
hairs in summer are considerably
thicker than in winter. The hairs of
the underfur in summer are thinner
than in winter fur. This is clearly
shown in tables (p. 440).
FIGURE 208, Penis of Spermophilopsis Winter fur much denser than
summer fur. N.P. Lavrov and S.P.
Naumov calculated the number of hairs
per 0.25cm? and obtained the following
figures: 1) summer fur in middle of
back, 475 hairs (217 guide and guard
hairs, 258 intermediate hairs and
hairs of underfur); winter fur in
Z middle of back, 1241 hairs (132 guide
a — left; b— right; ¢ — dorsal (according to
B. S. Vinogradov).
1 } and guard hairs, 1109 hairs of
FZ underfur); 2) winter fur in middle of
belly, 1962 (1075 guard hairs and
FIGURE 209. Glans penis of Euxerus guide hairs, 887 hairs of underfur);
ery tinepus summer fur in middle of belly, 155 (69
1 — left; 2— ventral (according to R. Pocock), guard hairs and guide hairs, 86
intermediate hairs and hairs of
undercoat). Winter fur on back three times as dense as summer fur, on
belly six times as dense as summer fur. Color of winter fur varying
accordingto locality. The palest specimens occur in the Central Kara-Kum. The
general color of the back of these specimens is brilliant rust-sandy-gold
(cinnamon-buff, pl. XXIX). Flanks paler (pinkish buff, pl. XXIX). Belly
white. The color of guard hairs in these specimens as follows: roots
depigmented, followed by a short blackish gray band, a whitish central
band and a rust-yellow tip. Tips of many hairs whitish yellow.
Length of winter hair in mm (average)
Middle of Middle of | Middle of Middle of |Middle of Middle of
Buttocks Buttocks Buttocks
back belly back belly back belly
Rr
383
441
Thickness of winter hair (in p)
hickest t
Type of hair e \e me ah. Middle of hair
Sampling zone Thickness of base
34.2
19,2
38
Middle of back ...
Middle of belly ...
Length of summer hair (in mm)
Type of hair
Zone from
which the
measurement was taken
Intermediate | Intermediate
hair (1st type) | hair (2ndtype)
Guide hairs Guard hairs Underfur
Middle of back ~ 2°: :.
Middle of belly .....
Thickness of summer hair (inp)
Type of hair
Zone from
which the
measurement was taken
Underfur
Gide hat asian Intermediate Intermediate
ard ha
caer i e . hair(1st type) | hair (2nd type)
150 103 SER L/ 20.5
90 10:5 57.5 17S
Top of head and back in specimens of the yellow color type buff-yellow-
gray darkened by the black-chestnut subapical bands of the hair. Flanks
purer rust-buff-yellow. Throat, neck and belly white. Color of individual
guard hairs as follows: roots depigmented, followed by narrow blackish
bands, whitish middle parts which become more rust-colored and yellow
distally, well marked black-chestnut subapical bands and buff-yellow-rust
hair tips. Shape of winter guard hairs as follows: proximal 3/4 straight,
not thicker proximally; terminal 1/4 of hair arrowlike or lanceolate.
Color of winter fur more golden and lighter than summer fur.
Color of young summer specimens different from that of adults, paler
and almost without any dark mottling. Fur very pale, delicate, yellow-
sandy, sometimes with a light pinkish tinge (between pinkish buff,
pl. XXIX and vinaceous-buff, pl. XL).
The number and arrangement of the vibrissae are constant throughout
the year. They are very distinct on the short and coarse summer fur.
In winter, the vibrissae can be found only by careful examination of the fur
in previously known places. Spermophilopsis hasa great variety of
vibrissae arranged in the following order. On the head: 1) paired groups
of 18-22 vibrissae on lateral part of the muzzle and between nose and eyes;
these vibrissae are called whiskers and they are 1.5-35mm long; the
shortest are near the nostrils and the longest at the sides, above the upper
lip; 2) paired groups of soft white vibrissae along the margin of the upper
Middle of back .....
Middle of belly
384
lip; each group consists of 10-15 hairs 0.7-3mm long; 3) single groups
along the margin of the lower lip; dense paired groups above the eyes, each
consisting of 6-10 (generally 7) hairs 7-20mm long; 5) one long vibrissa,
13-21 mm long near the outer corner of the eye; 6) paired groups of 4 hairs
on the cheeks, posterior to and below the
eyes; 7) a single group on the chin, consist-
ing of 3-4 vibrissae 20-22mm long. On the
forefeet: 8) groups of 8 hairs 11-21 mm long
on the outer ventral side of the wrist: 9)
one long hair (20-25mm) on the external
surface of the antebrachium. On the body:
10) a pair of symmetrical vibrissae on the
chest, 23-26 mm long (in females, these
vibrissae are between the first and second
teats); 11) a pair of vibrissae, 23-24mm
long, posterior to group 10; 12) two
symmetrical vibrissae, 15-16mm lceng,
lateral to group 11; 13) 1 or 2 vibrissae
18-23 mm long on the belly posteriorly and
laterally to group 12. These vibrissae are
at the sides of the belly in females, lateral
to the 2nd teat. N.P. Lavrov and S.N.
Naumov (1934) state that concentration of
the vibrissae on the underparts is
characteristic for Spermophilopsis.
Four pairs of vibrissae on chest and belly.
S permophilopsis (ventral) Sciurus vulgaris has only two abdominal
Drawing by A. N. Formozov pairs of vibrissae, only rarely three pairs
(according to I. D. Kidris).
The legs of Spermophilopsis are very typical. The soles of the
forelegs in summer fur are covered with sparse fur through which large
calluses are visible at the base of the toes. Stiff pectinate fringes of hair
(5mm long) grow along the outer margin of the sole, the lateral margin of
the fifth toe and part of the sole. Such fringes of white, flexible hair grow
also on the inner sides of the second toes. Hind soles with dense, thick
fur which reaches the base of the toes. Heels covered with dense fur.
Calluses of soles concealed by hair. Outer margin of hind feet with a well
developed pectinate hair fringe. Its hairs are especially long at the tip of
the fifth toe, forming a tuft along the claw which is longer than the claw
itself. Shorter hair fringes at the inner margin of the first toes. In winter
fur, the hair cover of the soles is better developed and the calluses of the
soles of the forefeet are almost completely covered with white hair. The
feet of Citellus fulvus, which is ecologically related to
Spermophilopsis, are bare. The tufted soles of Spermophilopsis
are apparently an adaptation to the hot sand. S.P. Naumov and N.P.
Lavrov (1934) state that the bare-soled C. fulvus lives mainly on fixed
sand, and even more often in steppes where the soil is cooler because of
the rich vegetation. Furthermore, C. fulvus goes into aestivation in
summer when the ground is very hot.
The hairy fringes of the toes are important. When the animal walks
or runs on the sand they stretch sideways, enlarging the supporting area
and preventing it from sinking into the loose sand.
442
FIGURE 210. End of tail of
385
SYSTEMATIC NOTES. In addition to specific characters, the skull of
Spermophilopsis has much in common with that of the squirrel.
Spermophilopsis is closely related to the African Xeridae in the
structure of the skull and the stiff hair of its summer fur. There are,
however, some differences in the form of the skull. The distribution of
vibrissae has been described above. The morphology of
Spermophilopsis proves that it is systematically far removed from
true Citellus.
GEOGRAPHICAL DISTRIBUTION. The northern boundary of the range
in the west passes north of the Krasnovodsk spit, south of Kara-Bogaz-Gol
Gulf. Farther east it occurs throughout the Chil'-Mamed-Kum Desert.
Further on the boundary is not known and then it passes into the Tashgauz
(on the Amu Darya). According to M.N. Bogdanov (1882), the northern-
most occurrence of this species in the Kyzyl-Kum is a locality near
Irkibai, near the Yana Darya river. It has not been found so far in the
valley of the Chu river. The habitat of the animal in the Dzhety Su is
possibly separated from its range in the Kyzyl-Kum. According to V.N.
Shnitnikov (1917), Spermophilopsis occurs onthe shore of Lake
Balkhash (between 46°30' and 47°10' N) and also on the Karatal between the
Ili river and the Karatal. The southern part of the range includes the
southern parts of Turkmenia and apparently the Asterabad Province in
Persia. Inthe southeastern corner of Turkmenia Spermophilopsis
occurs near Kushka, Kerki and in northern Afghanistan. There are
unfortunately no exact data on this southern boundary.
According to S. P. Naumov and N.P. Lavrov (1933),
Spermophilopsis apparently occurs throughout the Dardzha
Peninsula. They found it in the eastern part of the peninsula. Radde and
Walter (1890) recorded it from near the Mikhailovo fortifications. It also
occurs on Cheleken Island. According to V. Levchuk (1906), it is common
in the sands on this island. According to furriers (1933),
SpermophilopSsis apparently occurs throughout the Dardzha Peninsula.
They found it inthe eastern part of the peninsula. Radde and Walter
(1890) recorded it from near the Mikhailov fortifications. It also occurs
on Cheleken Island. According to V. Levchuk (1906), it is common in the
sands on this island. According to furriers (1933), Spermophilopsis
occurs throughout the sands south of the Dardzha Peninsula, to the south of
Chikishlyar. It is common in the vicinity of Kara Kala. According to
N.P. Lavrov and S.P. Naumov (1933), the animal is absent south of these
regions, in the southwestern corner of Turkmenia, between the railway,
the Caspian Sea and the Atrek river. It occurs only sporadically in sandy
and sandy-loamy areas in this region. In the northwestern parts of its
westernrange, N.P. LavrovandsS. P. Naumov found it inthe central part of the
Krasnovodsk spit. They write: 'Spermophilopsis is distributed
throughout the Chil'-Mamed Kum sands, according to our observations in
the western part of this region, and reports of fur trappers and Turkmens
from its eastern part.'' According to M.N. Bogdarov (1882), it lives in
the Kyzyl-Kum desert as far as the Syr Darya in the north, though it does
not reach this river inmanyplaces. The northernmost record during
M.N. Bogdanov's expedition was the Koskuduk well west of Irkibai, 2km from
from the Yana Darya. it is numerous onsand farther south as far as the
Amu Dar'ya. It is common in the KyzylKum, Dzhaman Kum and the sands
between the Murun (near Aristanbel - Kuduk) and Uch-Uchak. On the left
443
444
386
445
side of the Amu Darya it occurs in the sands to Kum! Daryalyk, near
Tashaus and Kun'-Urgench. It is absent on the Mangyshlak, Ust'Urt,
in the Barsuks, the Sam sands, the Astantai-Mantai, Isen Chagyl. N.P.
Lavrov and S.P. Naumov (1933) give detailed information on the distribution
of Spermophilopsis in Turkmenia: 'It apparently occurs throughout
the Kara-Kum... It was found at the southern outskirts of the Kara Kum
between Akhch Kuim in the west and Chardzhui in the east. In the center of
the Kara Kum at the line Ashkhabad - Ierbent - Sernye its mounds are
everywhere except in the low cultivated land along the railway. L.M.
Shul'pin informed us that in 1930 he found Spermophilopsis all along the
way in the central Kara Kum, north of the Bashi railway station. Furs are
processed in Darganat and Tatauz in the northern parts of the Kara Kum.
"In the southeastern Kara Kum, that is, the area between the Tedzhen
river and the Amu Dar'ya, and between the railway and the border,
Spermophilopsis was found in the followingplaces: K.K. Flerov found it
near the railroad stations of Teshkepri, Takhta-Bazar and Kushka on the
Kushka branch-line. According to fur-trading companies, it occurs near
Iolotan and Seraks. Shnitnikov recorded it from Kerkov. We found it south
of the Uchadzhi Station.
"It is still not known whether it occurs in the sands of the southern part of
of the above region. According to the forester N.I. Pustyshkin (of the
Chimchakly District), susliks of more northern areas, Uch-Adzhi and
Chimchakly, inhabit sand dunes. It may be assumed that they also occur
in the sand dunes of the southern region.
"They are absent in the greater part of the southwestern corner of
Western Turkmenia (bordered by the railway, the Caspian Sea and the Atrek
river). They occur only sporadically in sand and sandy-loamy patches.
The species is also absent in most of the northwestern corner of Turkmenia,
which is higher with a pebbly-stony soil. It is not found in most of the
cultivated, irrigated plain at the foot of the Great and Little Balkhans and
the Kopet Dagh. This plain is narrow, but widens in the Tedzhen and Mary
oases. Along rivers it forms extensions northward and southward. The
animal is absent in localities along the Amu Darya."
In 1925, V.G. Heptner and I found large numbers of Spermophilopsis
on sands near Ashkhabad and the Annau station. It is widely distributed
in Buchara. The species was described by Prof. Lichtenstein from a
specimen collected by Prof. Eversmann near Kara Ata, 140km from
Bukh. N.M. Dukel'skaya (1926-1927) found it in the Karakul District
(Zeravshan Region). N.A. Bobriskii found it in southern Buchara. In
the ZMAN collection are skins from Khadzhi Davlet, Karakul, Bogoabdal
(Buchara), Lower Pyandzh, Iman Baba (Mugrab), Bii Kal on the Amu Darya
(Buchara). V.N. Shnitnikov (1917-1936) reports that in Semirech'e the
Species is distributed along the shore of Lake Balkhash in the former
Kopal County (between 40°30! and 47° N). Small colonies live on the right
bank of the Karatal, in the Tantabai area (46°N and 47°E) and also in various
localities between the Ili river, Karatal and the shore of Lake Balkhash.
The animal occurs sporadically and in small numbers in these localities.
MODE OF LIFE. Spermophilopsis lives mainly on sand. In the
vicinity of Ashkhabad the sand hills reach as far as the arid semidesert.
Spermophilopsis keeps to the sand in this region, not entering the
semidesert. However, the range of Spermophilopsis is not restricted
to sands. N.P. Lavrov and §.P. Naumov (1933) found it on soil without any
sand, but they found it mainly on sand (sand dunes). They also found it in
387
pebbly-loamy foothills and clayey-sandy plains. Spermophilopsis
inhabits areas with solid soil only in the vicinity of sand.
Spermophilopsis does not occur in takyrs, solonetz* regions,
clay-Artemisia steppes and mountain-Artemisia steppes.
The animal is cautious and timid. It does not let man approach to
within shotgun range. It apparentlyhas good sight, as it notices danger from
afar. It is silent; I heard its voice only once, when, surprised it gave
a chirping cry and buried itself quickly in the sand. It may move as far
as 500 steps from its principal burrow. S.I. Ognev and V.G. Heptner
(1929) write: ''It bounds along with outstretched feet, raising its hind-
quarters and holding its tail upwards. Its movements are intermediate
between those of the squirrel and the hare and do not resemble those of the
Russian susliks which run close to the ground. When escaping from a
hunter, it tries not to lose sight of him. Having reached the top of a dune
it pauses, rises on its hind feet and raises its tail. The white color of its
underparts can then be seen from a long distance making it possible to see
the animal which is otherwise invisible because of its color which matches
the sandy ground. After looking about, the animal runs away, pausing
again on the next dune to have another look. The animal makes several
such inspections until it reaches its shelter. There it stops for a moment
and then enters. It spends a long period in the shelter and does not come
out again without making sure that there is no further danger.’
We studied the daily life of Spermophilopsis near Ashkhabad in
the spring and early summer of 1925. We found that the animals come out
late in the morning when it is already warm, and remain active until noon.
During the hottest part of the day they remain in their burrows. They
reappear towards evening, when the heat has subsided, but they are much
less numerous, less active and more cautious than in the morning.
N.P. Lavrov and S. P. Naumov (1933) provide detailed data on the daily
life of susliks in Turkmenia during different months. In the second half
of February they rarely came out of their burrows, showed little activity
and did not move more than 10-15m from their burrows. Their daily
activity began at about 10a.m. In the evening they returned to their
burrows at 5-6p.m. (in clear weather and a temperature of 12-15°).
In the last days of February the weather was clear and warm. The
temperature rose to 20-25°. The animals were more numerous, they
moved from their burrows and returned at 7 p.m. In late March the
animals began to come out at9a.m. and remained outside until 11-12 a.m.
After midday the number of animals outside decreased, and increased
again only after 4-5 p.m. The animals sometimes moved 200m from their
burrows. N.P. Lavrov and S.P. Naumov observed that the animals began
to come out earlier in late spring. In the middle of April they appeared at
6-7 a.m. In May D.N. Kashkarov (1929) saw them in the central Kara-
Kum as early as 5a.m.
According to observations by V.G. Heptner and myself (1925) near
Ashkhabad, the burrows of Spermophilopsis are wider than burrows of
European susliks. They are located mainly on slopes of dunes, either on
bare ground or under plants of Aristida pennata. Burrows are also
often built in soil cultivated with cereals, Peganum harmala and
Sophora pachycarpa. They do not inhabit typical solonetz, but their
446
* [A type of alkaline soil. ]
388
447
448
burrows may be near such soils. In front of the burrow is a heap of
excavated sand.
N.P. Lavrov and S.P. Naumov (1933) write: 'Spermophilopsis
lives mainly on sand hills and in depressions covered with silt and
surrounded by hillocks or barchans. Sandy soil is not necessary for
burrowing. Near the Sernye hillocks and the Dzhebel station we found
small numbers in the foothills. This is a plain consisting of solonetz
serozems with debris-gravel deposits and a small admixture of sand.
FIGURE 211. A burrow of Spermophilopsis in the desert near Ashkhabad
Photograph by S. I. Ognev
"Four kilometers west of the Dzhebel station we found a colony of
Spermophilopsis onthe Uzboi, close toa salt lake. The burrows were
on the slope of the bank among dense thickets of tamarisk, Elaeagnus,
Caragana and willow... On the opposite shore of the lake
Spermophilopsis lived ina plain of sand mixed with fine earth and
rubble.
"The more fixed the sands were, the more numerous were burrows on
slopes and hillocks, and the less numerous in valleys and depressions.
"In places with few shrubs the entrances of the burrows are usually
under shrubs of Haloxylon, Calligonum etc., which apparently
protect the burrow against damage and the animal from its enemies.
"On hillocks in the western part of the Krasnovodsk spit all entrances
were directed NE - SE. This is apparently related to the winds which
usually blow from the west.
389
"Most burrows have one entrance. We rarely found burrows with two or
more entrances.'' These authors did find burrows with three and even
four entrances. The burrows are very complicated. Lavrov and Naumov
continue: ''We excavated a burrow on 28 February, 1932 near the Dzhebel
station, south of the railway. It was on a small, sand hillock about 1m
high, and had two entrances which were under asmallSalsola
arbuscula, 3.5m apart. The main gallery was 4.8m long and went
through the hillock horizontally. It had lateral galleries which branched
and rejoined the main gallery. Some lateral galleries descended or
ascended, ending blind. These blind ends were always more than 28cm
below the surface. The galleries were usually 6-8.5cm wide and the blind
alleys 5-5.5cm. The branches or junctions of two or more galleries
formed chambers, but there were no nests. These chambers were about
15cm wide. The burrows contained only a few old stalks of Carex
physodes andCalligonum seeds. No remnants of food or feces were
found near the burrow. There was only a heap of sand in front of the
entrance.
Another burrow was excavated on 5 April near the Uch Adzhi station.
This was in a depression densely overgrown with Carex and surrounded
by mounds. The burrow had one entrance 12cm wide and 8.5cm high.
Lateral galleries, 17cm wide, branched off in both directions 0.5m from
the main gallery. The main gallery then became 27cm wide but its height
remained the same. Ata distance of 3.5m from the entrance the main
gallery ended in a broad chamber the ceiling of which was 13cm below the
surface. The lateral galleries had blind ends at least 37cm below the
surface, the deepest being at 120cm. We found a few seeds of
Calligonum and Haloxylon and capsules and stalks of Carex inthe
burrow. We also found some feces of sheep which had apparently been
introduced into the burrow by accident. We did not find any nests."
In addition to these permanent burrows, N.P. Lavrov and S.P. Naumov
often found temporary burrows. They dig them in various places,
generally in loose sand. They use these dwellings only for a short time,
for one night or for some hours during the day. The temporary burrows
are apparently used by specimens which have moved far away from their
principal burrow. The number of temporary burrows is in inverse
proportion to the amount of food ina place. The greatest number of
temporary burrows is found in loose sand. They are less numerous in fixed
sand. It is difficult to determine the number of temporary burrows as
they are easily destroyed in the loose sand.
It is thus difficult to determine the number of Spermophilopsis in
an area. Many burrows have to be considered as temporary, i.e., not
inhabited. N.P. Lavrov and S.P. Naumov (1933) observed the greatest
number of inhabited burrows in places with plenty of food plants. Thus
in late February 1932 almost 100% of the burrows near the Dzhebel station
were inhabited. In good weather, the ground squirrels could be observed
near almost all burrows. Where food was poor only 70% and in some
places only 20-30% were occupied. According to B.S. Vinogradov (1936),
crickets (Phyllobothrium vlassovi) are common inthe burrows.
According to V.G. Heptner (1929), in spring Spermophilopsis
feeds mainly on Carex physodes, digging the rhizomes out of the sand.
Burrows are everywhere during this time. Towards May the desert
becomes scorched, Carex has seeds, andSpermophilopsis has to
449
390
uiseie~A “yA Aq 3utmeiq
Inj 19}0IM UT “WYOTT sn[Azoepojday stsdojtydoursads
X ALVId
391
450
go far from its burrow for food. Later, it feeds on Artemisia, which
remains green even in July. The animal also eats shoots of conifers.
Fruits of various shrubs (e.g., Calligonum) become more important
as food in summer. In winter the animal feeds only on such fruits, and
climbs on the shrubs to obtain them. Spermophilopsis feeds on parts
of plants in the soil throughout the year. It is especially fond ofthe rhizomes
of Carex. It may occasionally take animal food, e.g., beetles. In
D.N. Kashkarov's opinion, the main food of the animal is tubers of Poa
bulbosa. N.P. Lavrov and S.P. Naumov (1933) observed that bulbs of
Tulipa patens and Allium are an important part of its diet from
February to April. They found that the stomach contents of several
specimens caught near the Dzhebel station consisted of a white mass witha
strong onion smell.
In Turkmenia estrus begins in the middle of February. In 1932 the
estrus was at its height during the first days of March. Estrus is
apparently not affected by bad weather. N.P. Lavrov and S.P. Naumov
write: ''We rode through the sands of Chil' Mamed Kum from 1 to 6 March.
Most of the time there was a snowstorm and the temperature dropped to
zero at times. Between the snowstorms, we found fresh tracks in the snow.
Following these, we found that some animals had made trips of almost 1 km.
Some animals came out of burrows and went along the slope of a sand ridge.
Sometimes, they would turn aside and dig in the snow near a bush. We
examined such places and always found burrows. The animals would
sometimes enter the burrow, but generally they went on to the next
burrow after cleaning the entrance. It seemed that these tracks had been
made by males in estrus, looking for females. They apparently did
not belong to feeding animals. Males caught in this period had much
enlarged testes.'' The estrus lasts several weeks. In the middle of
April one finds sterile females, females in estrus, and females in advanced
pregnancy. The duration of gestation is not known. Each litter consists of
three to six young.
Spermophilopsis molts twice a year. Spring molt starts onthe head.
The interorbital region and tip of the muzzle molt first. The winter fur
is retained longer on the occiput. The fur on the belly becomes thin, but
no new fur is observed. The molt then spreads tothe shoulders, tothe spine,
the inner sides of the haunches, the chest and anterior part of the belly.
The anterior part of the flanks then begins to molt. At this time the whole
body undergoes molt, but the process is slower on the hindquarters.
According to N. P. Lavrov and S.P. Naumov (1934), most animals caught
in late March in the sands north of Kazandzhik and Kizyl Arvat were molting.
Animals caught in early April in the southern part of the Kara-Kum (the
Uch-Adzhi station, Repetek) were in full molt. On the other hand, molt is
sometimes much delayed. Several specimens in the ZMAN collection caught
in late April near Karakul (20-27 April 1930) still had their full winter fur,
without any traces of molt. In one specimen caught near Karakul in 24 June
1930, molt was only beginning. Such a delay is abnormal. Most specimens
complete their molt in May. Several specimens caught near Ashkhabad and
Karakul on 4, 8-9 May 1930 had winter fur on the lower part of the back,
near the base of the tail and on the outer sides of the haunches.
According to N. P. Layrov and S.P. Naumov (1933) the base of the
tail molts first and the molt then spreads to the tip. The long hair at the
sides of the tail molts in the same sequence. Old hair falls out more
392
451
rapidly than the new hair grows, and at some stage of the molt there is
only a patch of frayed winter hair at the tip of the tail. The new hair
grows in the opposite direction, i.e., from the tip to the base.
The skin grows dark during the molt, but the darkening does not reflect
the actual process of molt. The skin on chest and belly, which are covered
with white hair, keeps its color during the molt, as it does not contain
pigment.
N.P. Lavrov and S.P. Naumov did not find different molting dates for
males and females.
The molt of the young of the same year into adult fur (which is shorter,
coarser and with dark mottling) is completed in the second half of July.
Molt begins on the head and ends on the hindlegs and buttocks as in adults.
This could be determined from the large series in the ZMAN.
The hair on the feet changes so gradually that it is difficult to determine
exact data.
Autumn molt begins on the outer side of the tibiae and on the buttocks.
It then spreads to the flanks and back towards the head.
The head molts last. The summer fur does not show signs of molt in
August. The summer fur of a specimen caught on 2 October 1926 near
Saryyazal (along the Kushka railway siding) was very thin on the hind
quarters and new winter fur was visible underneath. Specimen No. M.703
of my collection (caught 18 December 1925 near Repetek) has its full
winter fur.
Spermophilopsis does not truly hibernate. It remains in its
burrow in winter, waiting for the warm weather.
The animals may have several such ''sleep periods" during the winter,
depending on the weather.
From June to September, many squirrels in the sands near Ashkhabad
block the entrances of their burrows with sand from the inside and do not
go outside. This aestivation is apparently similar to that of Citellus
fulvus. It would be interesting to find out whether Spermophilopsis
goes into aestivation throughout its range, and what percentage of the
population remains active throughout the summer.
Spermophilopsis is diurnal and is therefore not hunted by nocturnal
predators. It is hunted by foxes in some localities. N.P. Lavrov and
S.P. Naumov (1933) found many skulls and bones in the Krasnovodsk sands.
Hair of Spermophilopsis was repeatedly found in feces of foxes.
Spermophilopsis is hunted by the following birds: buzzards, harriers
and probably some eagles.
Spermophilopsis causes no harm to agriculture as it lives far
from cultivation.
Spermophilopsis causes damage by destroying sand-fixing plants.
Seeds of Calligonum and Carex physoides, which are important
for this purpose are frequently eaten by it. Near the Dzhebel station N.P.
Lavrov and $.P. Naumov repeatedly found seeds of Calligonum which
had been sown by the Service of Protective Forests of the Railroad to fix
the sand near the railway. They state: ''We were surprised how
Spermophilopsis succeeded in finding the seeds buried in the sand.
Nearly all seeds are destroyed by Spermophilopsis.' The animals dig
up large areas of fixed sand to obtain the roots and bulbs of plants. Holes
made for this purpose are close together. The whole feedingarea is covered
with holes (12cm deep and 7-10cm wide) and with heaps of earth.
393
D.N. Kashkarov (1932) counted 12,400 such holes per hectare in the Central
Kara Kum.
452 The winter fur of the sand marmot is silky and soft. The skin is tough.
The pelts may be dyed, and the hair absorbs and retains the dye well. The
pelt is of more value than that of Citellus. Commercial use of
Spermophilopsis began in 1930, but is still very limited.
The Turkmens capture it with traps. Russians rarely shoot it. Shooting
is not a good method as wounded animals often reach their burrows. They
have to be shot from a long distance as they are very shy.
Subspecies of Sbermophilopsis
376. Spermophilopsis leptodactylus leptodactylus Licht.
(1823).
1823. Arctomys leptodactylus Lichtenstein H., Nathurhistor, Anhang zu Eversmann's Reise nach
Orenburg und Buchara, S, 119,
1827-1834. Citellus leptodactylus Lichtenstein H., Darstellung neuer Sdugethiere, Berl. , Taf. XXXII.
1852, Spermophilus leptodactylus Brandt J. F., Zoologischer Anhang zu A. Lehmann's Reise nach
Buchara und Samarkand, S,. 303.
1882. Spermophilus leptodactylus Bogdanov M.N., Ocherki prirody Khivinskogo oazisa (A Survey
of the Nature of the Khiva Oasis), pp. 27-28, fig. on p. 46.
1890, Spermophilus leptodactylus Radde, G. A. Walter and W. Blasius, Die Saugethiere
Transcaspiens, Tiflis, 1890, S. 38.
1918. Spermophilus leptodactylus Bil'kevich S.I,, Kollektsiya Zakaspiiskogo muzeya (The
Collection of the Transcaspian Museum), Izvestiya Zakaspiiskogo muzeya, I, p. 7.
1927. Spermophilopsis leptodactylus leptodactylus Obolenskij S., A Preliminary Review of
the Palearctic Sousliks, Compte rendu de 1'Académie des Sciences, Leningrad, p. 193.
1929. Spermophilopsis leptodactylus (leptodactylus) (Licht. ) subspecies distinguenda Ognev
S.1, and V.G. Heptner, Mlekopitayushchie srednego Kopet-daga i prilezhashchei ravniny (Mammals of
Central Kopet Dagh and the Adjacent Plain), Trudy Instituta zoologii, 3 (1):131-134,
1933, Spermophilopsis leptodactylus Lavrov N.P. andS.,P. Naumov, Rasprostranenie i biologiya
tonkopalogo suslika v Turkmenskoi SSR (The Distribution and Biology of Spermophilopsis in
Turkmenia), Zoologicheskii zhurnal, 12 (2):80-116; by the same authors: Stroenie mekha i lin'ka
tonkopalogo suslika pustyni Kara~Kum (Fur Structure and Molt in Spermophilopsis in the Kara-Kum
Desert), Ibid, 13 (2): 311-332, 1934,
TYPE LOCALITY AND DEPOSITION. Vicinity of Kara Ata, 140km northwest of the Old Town of
Bukhara. Holotype probably in the Berlin Museum,
MATERIAL EXAMINED: 151 pelts and skulls.
DIAGNOSIS. Summer fur coarse, stiff and thin. Summer fur on back
light, uniformly buff-sand-colored witha rust shade near cinnamon-buff,
light pinkish cinnamon (pl. XXIX), sometimes with a yellowish tone (light
ochraceous-buff, pl.XV). Summer fur sometimes with a light rust tone
and fine gray-chestnut mottling. Winter fur delicate and silky. Color
between rich buff-rust-sandy and rust-drab. Fine drab-chestnut mottling
poorly developed.
453 MEASUREMENTS: Lengthofskull 53-58.2mm (M.54.5mm); condylobasal
length 47.8-53.5mm (M.49.51mm); zygomatic width 33-36.6mm (M.
(M.34.42mm); interorbital space 14.9-18mm (M.16.07mm); length of
upper tooth row 9.8-12.2mm (M.11.1mm). Length of body and head 207-
305mm (M.237mm); tail without terminal hair 43-102mm (M. 67.1 mm);
hind feet 51-67 mm (M.57mm).
SYSTEMATIC NOTES. Even in large series of Spermophilopsis,
the characters of the subspecies are difficult or impossible to recognize.
This is due to the unevenness of the series and the unavailability of
394
454
specimens from different seasons. V.G. Heptner and Ognev (1929)
suggested that the specimens from the Kara-Kum belong to a separate form
which differs from the nominate subspecies in the richer color of the winter
fur. The winter fur of this form is ocher, while that of the nominate
subspecies is a duller buff-yellow with marked fine mottling. Study of
large series did not confirm that these differences are constant. Summer
specimens from Ashkhabad, Kerki and Bukhara are also indistinguishable
from the nominate form. S.1.schumakovi (from near Kushka) differs
clearly fromS.1. leptodactylus inits more saturated drab-rust
color with fine chestnut-brown mottling (see below). Specimens from the
central Kara-Kum (Sernye hillocks) are probably a separate subspecies, as
suggested by N.P. Lavrov andS.P. Naumov. The back of these specimens,
in winter fur is rust-sandy-golden (cinnamon-buff, pl. XXIX), the flanks are
paler (pinkish buff, pl. XXIX) and the belly is white. Lavrov and Naumov
did not obtain summer specimens of this race. The status of this form
will have to be determined when such specimens are available.
J. Scully (Mammals and Birds Collected in North Afghanistan. Journal
of the Asiatic Society of Bengal, Vol. LVI, part II, No.1, 1887, pp. 70-71)
described Spermophilus bactrianus from Afghanistan (Khamiab).
As Scully states, this animal is closely related to Spermophilopsis
leptodactylus. His detailed description shows that the Afghan
squirrel differs from S.1.leptodactylus as follows: 1) 6 teats,
(8 inleptodactylus); 2) shorter tail; the tailofbactrianus is
shorter than the hind foot; tailofleptodactylus longer (see above).
S.bactrianus is probably a different species closely related to the
pe leprodact yluws:
The geographical distribution has been given above. It should be added
that the nominate race occurs throughout the range except in the central Kara-
Kum which is inhabited by a pale, as yet undescribed subspecies, and the
southeastern corner of its range (the Kushka District) where S.1.
schumakovi occurs. It is not known which form occurs in the
Semirech'e, from where it was recorded by V.N. Shnitnikov (see above).
The Semirech'e squirrel possibly also belongs to a separate race.
376. Spermophilopsis leptodactylus schumakovi Satunin
(1908). Kushka, or Shumakov's ground squirrel
1908. Citellus (Spermophilopsis) schumakovi SatuninK.A., Materialy k poznaniyu
mlekopitayushchikh Kavkazskogo kraya i Zakaspiiskoi oblasti. VII, Novyivid suslika iz Zakaspiiskoi
oblasti (Material on Caucasian and Transcaspian Mammals. VII. A New Species of Ground Squirrel
from Transcaspia), Izvestiya Kavkazskogo muzeya, Vol. III, pp. 254-260.
1918, Citellus schumakovi Bil'kevich S.I., Kollektsii Zakaspiiskogo muzeya (Collections of the
Transcaspian Museum), Izvestiya Zakaspiiskogo muzeya, f, p. 7.
1927. Spermophilopsis leptodactylus schumakovi Oboilenskii S.I., Rukovodstvo k opredeleniyu
suslikov Palearktiki (A Key to Palearctic Ground Squirrels), Izdatel'stvo Oblastnoi stantsii zashchity
rastenii, Leningrad, p. 3; Obolenskij S., A preliminary review of the Palearctic ground squirrels,
Compte rendu de 1'Academie de Sciences de 1'U. R.S.S., 1927, 193.
1933, Spermophilopsis leptodactylus schumakovi Vinogradov B,S,, Mlekopitayushchie
SSSR, Gryzuny (Mammals of the U.S.S.R., Rodents), p. 17.
TYPE LOCALITY AND DEPOSITION. Vicinity of the Kushka fortress, Transcaspia. Holotype in the
collection of the Caucasian Museum (formerly the Georgian Museum), No. 70A. a.
MATERIAL EXAMINED: 10 pelts with skulls.
395
455
DIAGNOSIS. Differs from typicalS.1. leptodactylus inthe more
saturated gray-rust tone of the summer fur (between pinkish cinnamon and
cinnamon-buff, pl. XXIX). Its most typical character is the presence of
more intense chestnut-brown bands on the hairs, giving a more chestnut-
gray tinge to the fur. These bands are blackish gray on top of the head,
giving a darker tone to the head. Dorsal side of tailofS.1.schumakovi
darker and more rust-colored than inS.1l.leptodactylus.
Dark chestnut-gray mottling of buff-yellow-gray winter fur more marked
thaninS.1. leptodactylus. The winter furofS.1. schumakovi
is thus darker and duller.
Skull larger. Length of skull 56.3-59.2mm (M.58.4mm); condylobasal
length of skull 51.6-54mm (M.53.2mm); zygomatic width 36-38.6 mm
(M.37.6mm); interorbital width 15.5-20.1mm (M.18.2mm); length of
upper tooth row 11.3-12mm. Length of body and head 237-278mm; tail
62.2-74mm; length of hind feet (without claws 56-60.5mm.
GEOGRAPHICAL DISTRIBUTION. Restricted to the southeastern
corner of Turkmenia, the Kushka District.
Subfamily MARMOTINAE Chipmunks, Ground Squirrels and Marmots.
Terrestrial mammals living in underground burrows. Only the chipmunks
climb on trees although they live in underground burrows.
The fur of the back of marmots, ground squirrels, Citellus fulvus
and prairie dogs is of uniform color. In other genera of the subfamily, the
fur on the back shows more or less developed speckling or well marked
stripes (Callospermophilus, Citellus tridecemlineatus,
Eutamias, Tamias).
Tail 1/6 - 1/2 or more of the body length, densely covered with hair
of moderate length. The terminal hair of the tail is longest.
Ears very short, forming small folds projecting only a few millimeters.
The ears of chipmunks are longer. Forefeet with four toes, hind feet with
five. The third toe of the forelegs is longer that the others. The third
or the third and fourth toes of the hind legs are longer than the others, both
of about the same length.
Claws of ground squirrels and marmots long, less curved than in
squirrels. Claws of chipmunks relatively short and curved.
Cheek pouches present.
Molars brachydont, transversely elongate, irregularly rectangular. The
tubercles of the molars form transverse, elevated enamel folds.
0. 0. 0. 4 1, 23 0, 0. 3.4 Lo 8
ET Ca Re OR RT IY na Te: ©
Glans penis very narrow terminally, taperingtoa slender fingerlike
appendage which is slightly inclined downwards.
Os penis of Citellus small and of peculiar form. Baculum with
widened, club-shaped base, ending in a flattened, widened, leaf-shaped
part or ina plate with a serrated margin. Os penis of Marmota not so
widened and with fewer denticles on the terminal plate. The terminal plate
of the baculum of the genus Cynomys is even less widened and has fewer
denticles.
Formula of molars:
396
The geographical distribution and the genera of this subfamily have been
given above.
KEY TO GENERA AND SUBGENERA OF THE SUBFAMILY MARMOTINAE IN THE U.S.S.R.
1 (2). Back with five well marked black or dark chestnut stripes beginning at the occiput along the
whole back, and alternating with light buff-yellow stripes
Genus Eutamias Trouessart,
2(1). Back without dark stripes, of uniform color or speckled,
3 (4). Size large. Length of body and head of adults not less than 380mm, length of hind feet without
claws more than 70mm. Pupil round
Genus Marmota Frisch,
4 (3). Much smaller animals, Length of body and head usually about 200mm, to 375mm in very old,
large Citellus fulvus. Length of hind feet without claws usually less than 60mm, reaching 60mm in
oldC. fulvus. Pupil usually oblong
Genus Citellus Oken.
5 (6). Hind soles bare or only margins with narrow fringes of hair
Subgenus Colobotis Brandt
6 (5). Hind soles with dense short hair.
7 (8). Nasal processes of premaxillaries narrow in their posterior part, about as wide as one nasal in its
posterior third,
Interorbital space narrow, about 1/5 of the condylobasal length
Subgenus Citellus Obolenskij (ex Oken)
8 (7). Nasal processes of premaxillaries very wide in their posterior part, about as wide as both nasals
in their posterior third,
Interorbital space broad, its width about 23 % of the condylobasal length
Subgenus Urocitellus Obolenskij
Genus Eutamias Trouessart (1880)
1769. Sciurus (partim!) Laxmann M.E., Sibirische Briefe, S.69-71; Gmelin I. Fr. , Caroli Linné
Systema Naturae, Tom I, 1788, pp. 150-151,
1826. Myoxus (partim,') Siebold Ph. F., De historia naturalis in Japonia statu, etc, , Wirgerburgi, p. 17
(M. lineatus).
1851, Tamias, Simashko Yu. , Russkaya fauna tom II, Mlekopitayushchie (Russian Fauna, II, Mammals),
p. 601,
1859, Tamias BairdS.F., Mammals of North America, Philadelphia, p. 295(T. pallasii).
1880, Eutamias Trouessart E,L,, Catalogue des Mammiféres vivants et fossiles, Fasc. III, p. 29
(E. asiaticus Gm.),
1898-1899, Tamias Trouessart E.L, , Catalogus Mammalium tam viventium quam fossilium, Berolini,
p. 429,
1904, Eutamias Trouessart E.1l., Catalogus Mammalium, Supplementum, p. 330.
Mainly terrestrial rodents which climb well on trees.
The most typical character of the genus is the presence of five well
marked, longitudinal, black-chestnut stripes alternating with light yellowish
white stripes on the back. Tail without terminal hair 2/3 to 3/4 of the
length of the body. Tail about half the length of the body in some American
species.
397
457
458
Hair of the tail of moderate length. Ventral side of tail without lateral
combing".
Ears of moderate length, about one third longer than wide. Tips of
ears without tufts. Cheek pouches developed.
Vibrissae arranged as follows: 1) the longest, thickest and darkest
vibrissae between eyes and nose; 2) 3-4 vibrissae above the eyes; 3)
small group of 2-3 hairs on the cheeks; 4) on chin and throat; these
vibrissae may be absent; 5) on the outer sides of the forelegs, lateral and
posterior to the wrist and on the elbow; 6) in the genital area in males.
Teste: »pwhelk sad2=2iet 1-1. Toval 5.
Frontal processes of premaxillaries widened. Width of each process in
its posterior part equal or almost equal, sometimes markedly wider (in
Eutamias speciosus) than width of both nasals in their basal part.
Coronal suture sometimes nearly rectangular. Posterior part of coronal
suture sometimes with a sharp lateral projection which points ventrally and
slightly laterally.
Interorbital space of moderate width, between 26-28 % of condylobasal
length. Frontal area slanting and convex. Braincase rounded-oval.
Anterior part of zygomatic arch at level of zygomatic process of maxilla
flattened horizontally.
Infraorbital foramina larger than in other genera of Sciuridae, oblong-
oval and without angular elevations at base of the infraorbital foramina.
Diastema very long, almost twice as long as upper tooth row.
Dental formula; i= : co eal 5) < ; ms (=22);
[in the North American genus Tamias: i z: cS = ; 2 m5 (=20)].
Type species: Sciurus (Eutamias) sibiricus Laxmann.
GEOGRAPHICAL DISTRIBUTION. Palearctic region from the boreal
zone of the Kholmogory forests of the former Arkhangelsk Province to the
Bering Sea, reaching the Tundra in the north.
The southern boundary reaches as far as the forests. It turns in a west-
east direction, reaching about 48°N in Central Asia. It reaches the Central
Asian deserts and includes Manchuria, Korea, Shantung and the central
parts of China. In China the chipmunk enters the Oriental Region.
Chipmunks also occur on Sakhalin, Iturup and Yezo [Hokkaido].
In North America species of the genus Eutamias * mainly occur in the
western part of the continent, from the Yukon and British Columbia to
Ontario, and as far as Arizona, New Mexico and Texas in the south.
167. Eutamias sibiricus Laxm. (1769). The chipmunk
1769. Sciurus sibiricus LaxmannM.E., Sibirische Briefe, Géttingen-Gotha, S. 69-71.
NAMES: Chipmunk, Pallas! Ground Squirrel, Siberian Ground Squirrel (English); Burunduk, das
gestreifte Backenhérnchen (German); le Suisse (French).
LOCAL NAMES: "polosatik"' [striped animal] (northeastern Europe); ''polozok" [crawler] (Northern
Dvina); ''zemlyanoi zaichik"' [ground hare] (Baraba Steppe); 'medvezh'ya sovest'" [bear's conscience] (a
jocular name used by Siberian commercial hunters); koch'shar, k'zer or kuozver (along the Sos'va, Mansi
and Vogul languages); orda, orda vitsya (the word "'vitsya" means stripe, Komi, Zyrian); urdo (Votyak);
* G.S. Miller records 16 species and 57 subspecies for North America (List of North American Recent
Mammals, Washington, 1924, pp, 199-208).
398
vaarte (Verkhoturye); kustavai (Khanty, Ostyak); kuiarek, yul-ala (Bashkir); kuegeruk (Tatar); kucheger
(near the Surgut); sheiek (near the Naryn); koop (near the Ket); denka (near the Yenisef); dzhireki
(Mongolian); guriki or girokhi (Buryat); ul'chuki maruvki (Buryat); makhotoi, muruku (Yakut);
ulkichan (Manegir and Birar Tunguses); ulchuki or ulchi-oron (on the Sayan Mountains);
tar (Nivkhi, Gilyak, western coast of Sakhalin); tagkhr (Nivkhi, inner part and eastern coast of Sakhalin);
atakhniki (Ainu); ugukki (Nanai, Orochon); uldshe (Mangun); on'ocho (Kiler near the Kura); iitan (Chukchi);
moryumki*, Haopangtza (Chinese).
PRINCIPAL FIGURES: 1) Dvigubskii I. , Opyt estestvennoi istorii vsekh zhivotnykh Rossiiskoi imperii
(A Natural History of the Animals of the Russian Empire), 1829, pl. 18 (entire animal — poor figure); 2)
Simashko Yu. , Russkaya fauna (Russian Fauna}, Vol. II, 1851, pp. 601-603, pl. 32, figs. 2-3 (in color);
3) Middendorf A. Th, , Sibirische Reise, Sdugetiere, 1867, S. 83, Taf.1I, Fig. 10 (embryo); 4) Fokht K.,
Mlekopitayushchie (Mammals), Sankt Peterburg, 1885, p.376, (drawing of entire animal); 5) Brehm's
Tierleben, Sdugetiere, II, 1914, Taf. S. 516 (in color}; 6) Sowerby A. de C., The Naturalist in Manchuria,
II-III, 1923, pp, 148-149, Tab, VII, photographI; 7) Kishida K., An Explanatory Illustration of Japanese
Mammals, 1925, 2nd edition (in Japanese), 1926, pl. V (entire animal); 8) Bobrinskii N. A., Opredelitel'
okhotnich'ikh i promyslovykh zverei (Key to Game and Commercial Mammals), 1928, p, 48, fig. 32 (entire
animal), see also 'Opredelitel' okhotnich'ikh i promyslovykh zverei SSSR" (Key to Game and Commercial
Mammals of the U.S.S.R.), 3rd edition, 1935, p.53, fig. 44 (same figure as in the preceding books);
9) Bakutin M.G. , Nablyudeniya nad burundukom (Observations on the Chipmunk), Izvestiya sibirskoi kraevoi
stantsii zashchity rasteniil, 1930, pp. 69-74, photograph 7 (a hibernating chipmunk), figs. 1-2 (diagrams of
burrows); 10) Zverev M., Vrednye gryzuny Sibiri (Harmful Rodents of Siberia), 1930, p. 24, fig. 6 (entire
animal); 11) Velizhanin G. A. , Zoo-ekologicheskoe obsledovanie khor'kovogo zakaznika Sibirskogo
otdelniya instituta zashchity rastenii (A Zoological-Ecological Survey of the Polecat Reserve of the Siberian
Institute for Plant Protection), Trudy po zashchit rastenii, 2 (8): 61-62, Novosibirsk, 1931, pl. III
(photograph of entire animal), pl. 1V (photograph - poorly reproduced); 12) Zverev M. D., Instruktsiya dlya
nablyudeniya za burundukom (Instructions on the Control of Chipmunks), Vsesoyuznoi gosudarstvennoe
ob'edinenie po bor'be s vreditelyam, 1932, p.1, fig. (entire animal); 13) Plyater-Plokhotskii K., Vrednye i
poleznye zhivotnye v sel'skom khozyaistve DVK (Harmful and Useful Animals for Agriculture of the Far
Eastern Territory), Moskva-Khabarovsk, 1934, pp. 40-47, fig. 18, p, 41 (entire animal); 14) Zalesskii I. M.
M.D. Zverev, Burunduk (The Chipmunk). —In the Collection published by Vsesoyuznot Kooperativnoe
Izdatel'stvo Moskve, 1935, pp. 80-102, fig. 33, p. 81 (entire animal); fig. 34 (map of geographical
distribution); fig. 35, p. 86 (hibernating chipmunk); fig. 37, p. 88 (chipmunk on tree); fig. 38, p. 89
(burrows); fig. 39, p. 91 (chipmunk with full cheek pouches),
MATERIAL EXAMINED: 531 specimens (pelts with skulls).
DIAGNOSIS. Slender animal with rounded-oblong head, without a
markedly thinner neck.
459 Ears of moderate length, about one third longer than wide. Tip of ears
without tufts of hair.
Tail without terminal hair about 2/3 to 3/4 of length of body. Hair of
tail moderately long. Ventral side of tail without lateral ''combing"'.
Hind feet relatively long. Short and dense hair covers the soles from
the heel to the calluses. Four calluses at the base of the toes. The two
lateral calluses are of about equal size, larger than, and posterior to, the
inner calluses.
Anterior sole with five calluses. The inner posterior callus is the
largest. There is an additional small callus on the innter side of the foot,
at the base of the fifth toe.
Vibrissae arranged as follows: 1) the longest, dark vibrissae between
eye and nose; 2) 3-4 vibrissae above the eyes; 3) cheeks with a small
group of 2-3 vibrissae; 4) on chin and throat; 5) outer sides of forelegs,
posteriorly and laterally to the wrist, on the elbow. Vibrissae are also
present in the genital region in males.
Meats: p.1=-1;,.a.2-2; i, 1-1; total 8.
* [Obviously an error, since moryumki is not a Chinese name, ]
399
461
Five longitudinal black-chestnut stripes on the back. The middle stripe
begins at the occiput, between the ears. The two stripes next to the middle
stripe nearly reachthe ears and continue posteriorly, almost to the base of
the tail. The dark stripes alternate with light whitish yellow stripes which
give an ocher-rust color to the lower part of the back. Belly white.
Frontal processes of premaxillaries widened. Width of each frontal
process in its posterior part about equal to width of both nasals in their
basal part.
Coronal suture sometimes nearly rectangular. Posterior part of coronal
suture sometimes with two sharp lateral processes directed ventrally and
laterally. Interorbital width moderate, about 26-28% of condylobasal length.
Bullae osseae always longer than wide.
The skull of the chipmunk differs from that of most ground squirrels
as follows: 1) broader nasal processes of premaxillaries; 2) middle of
frontal area not flattened or depressed but convex* and slanting; 3)
braincase very elongate, rounded-oval; 4) anterior part of zygomatic
arch at level of zygomatic process of maxilla flattened horizontally; 5)
diastema very long, almost twice as long as upper tooth row.
The skull of the chipmunk differs from that of the squirrel in some
important characters: 1) the interorbital space is much narrower; 2)
braincase narrower and more elongate; 3) infraorbital foramina larger,
without angular, bony process as the base of the foramina; 4) anterior
part of zygomatic arch flattened horizontally (see above), not forming an
elevated, anterior plate as in true squirrels.
DESCRIPTION. Rostrum moderately elongate. Zygomatic arches
tapering anteriorly, widest in their posterior third. Braincase rounded-
oval, moderately broad.
Nasals widened anteriorly and narrower at the base. Frontal processes
of premaxillaries markedly widened, but not forming wing-shaped plates
obliquely pointing downwards as in squirrels. Posterior part of frontal
process as broad as basal part of both nasals.
Zygomatic arches moderately tapering anteriorly, widest in their
posterior third.
Postorbital processes thin, sharp, and shorter than in squirrels and
ground squirrels.
Posterior part of coronal suture almost rectangular, straight, with
sharp lateral processes (Figures 212-215).
Diastema almost twice as long as upper tooth row.
Bullae osseae always longer than broad.
Molars with transverse enamel folds as in squirrels, alternating with
grooves, and enamel tubercles at the inner margin of all teeth in form of
small, rounded arches.
First upper premolar very small, inclined backwards against the anterior
side of the second premolar. Crown of second premolar about five times
as large as that of the first premolar. Crown of first premolar trapezoidal,
with three transverse enamel ridges. The distal, anterior ridge is poorly
developed. The other upper molars have two transverse enamel ridges
alternating with three grooves. Two of these grooves lie along the outer
margin of the ridges and one of them lies between the ridges. All teeth with
* Such a convexity is present also in the American species Citellus tridecemlineatus.
400
well-marked inner enamel arch or crest. These crests form a chain along
the inner side of the tooth row.
FIGURES 212-215, Skull of Eutamias sibiricus Laxm,, from a specimen from the Minusinsk Territory
Drawing by Yu. A. Kostylev
Each lower molar with two rounded denticles on the outer side of the
crown. The anterior denticle is larger than the posterior. Each molar
with one high anterior denticle at the inner side of the tooth row, followed
by the raised inner margin and the lower posterior, inner marginal
denticle. A large depression in the middle of the crown which is
particularly large on the last molar. Anterior margin of crowns with a
small transverse enamel ridge which connects the outer and inner anterior
denticles.
Color. Top of head varying from gray-buff-chestnut to more
intense rust-brown with buff mottling in different subspecies. Eye
surrounded by a whitish ring. A whitish stripe with a buff tone at the
margins from the ring to the nose. Another stripe from the posterior
ventral corner of the ring to the base of the ear. A chestnut-black-brown
stripe from the posterior margin of the ring to the ear, ending in the
middle of the inner side of the ear. A dark-chestnut stripe bordered by
401
462
chestnut of varying intensity from the anterior part of the muzzle, beneath
the ring around the eye, and widening posteriorly. Lower part of cheeks
dirty whitish buff. Posterior parts of cheeks sometimes more yellowish
rust colored. Occiput behind the ears ash-gray with a rust tinge. Five
chestnut-black longitudinal stripes running from the occiput to the middle
of the back. Middle stripe from occiput to base of tail. The stripes next
to the middle stripe begin on the shoulders and continue to the posterior
part of the middle of the back. The lateral stripes begin anterior to the
base of the forelegs and continue to the sides of the buttocks. Between
the five dark stripes there are four light stripes. The anterior part of the
light stripes is whitish buff, and the posterior part more reddish rust
colored. The intensity of the rust shade varies in different subspecies
between cinnamon-buff and cinnamon, sayal-brown and orange-cinnamon
(Ridgway, pl. XXIX). The whole posterior part of the back and the haunches
are reddish rust of varying intensity. Yellowish rust tones are also present
on the flanks. This color is sharply delimited from the dirty whitish tone
of the belly. Tail chestnut-gray. Tips of hair on tail white. Color of hair
on tail follows: Roots usually pale buff-rust, followed by broad black bands
and white tips. There are also many almost completely black hairs with
whitish tips. There are also rust-buff hairs, especially on the ventral side
of the tail.
FIGURE 216, A chipmunk
Drawing by A.N. Formozov
402
463
464
Dorsal side of feet gray-chestnut-buff (drab).
Vibrissae gray. Claws dull gray-rust colored.
Color of chipmunks in the middle of summer dull without rich yellowish
rust and reddish tones. This results from the loss of the rust-colored hair
tips, which causes a grayer and duller tone.
The chipmunks molt only once a year. Variations in the texture and
color of the fur are caused by wear of the hair tips.
The color of young chipmunks is duller and grayer than that of adults,
and their hair cover is thinner, shorter and softer.
SYSTEMATIC NOTES. Nine subspecies of Eu. sibiricus are
known at present. Their distribution is shown in Map Ix.
GEOGRAPHICAL DISTRIBUTION. (Map IX). The northwestern boundary
of the range of the chipmunk runs from Vologda in the north to Kholmogory,
reaching the lower reaches of the Mezen river. Chipmunks possibly occur
northwest of this boundary as forests of the eastern type with Siberian tree
species form an extension northwest of Kargopol. The chipmunk is
probably found in the upper reaches of the Onega*.
East of the Mezen the northern boundary of the range runs to the Pechora,
in the northern coniferous forests mixed with Siberian trees. In the
northern Urals chipmunks have been reported by good authority from about
64°N. Farther east the boundary crosses the southern part of the Ob and
touches the West Siberian tundra, entering deep into the forest tundra.
Chipmunks are rare near Yanov Stan (66°N) and near Sidorovskaya on the
Taz (66°30'N). According to A. Middendorf, they occur in the Turukhansk
Territory near Igarka (68°N). Inthe north Siberian taiga they occur
near the southern part of Norilskoe Lake. Farther east, the boundary
crosses the middle reaches of the Olenek river and passes to the Lena
river (at about 67°N). Onthe Yana river the boundary turns sharply to
69°N). Farther east it crosses the Indigirka, but the distribution of
chipmunks here is incompletely known. It then continues to the estuary of
the Kolyma. Inthe Anadyr Territory chipmunks occur in the valley of the
Anadyr river. The boundary then passes to the Apuka river, reaching the
Bering Sea and the Olyutorski Gulf. It then turns to Penzhina Bay.
The southern boundary of the range of the chipmunks runs from Vologda
through the central part of the former Kostroma Province (Varnavin)
to the Kerzhenets river, where chipmunks are rare and to the Vetluga river,
where the animal becomes more common (A.N. Formozov, in litt.) I
recently received information on the occurrence of chipmunks in the
Temnikov District (the northern part of the former Tambov Province). In
this region chipmunks are hardly hunted by fur companies, Only three
animals were caught in 1934. V.A. Kraft saw one pelt in 1935, but he
could not obtain it from the fur company of the town Temnikov. Chipmunks
have been found in the vicinity of Sarov, among fallen trees and in groves
of European mountain ash near millet fields. This should be further
investigated. If it is confirmed that chipmunks occur constantly in this
area, the boundary will have to be moved considerably to the southwest.
G.K. Gol'tsmaier (1934) reports that in early summer 1910, P.P.
Yal'tsev saw a striped animal which he identified as a chipmunk, ina
mixed forest near the village Lutskoe in the Kotelva Subdistrict (Chuvash
* A.A. Byalynitskii-Birulya recently wrote to A.N. Formozov that he read in a Finnish publication that
chipmunks have been found in Finland, This is highly interesting,
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465
ASSR). The forests of the vicinity of Lutskoe are close to the Trans-Sura
forest massif on the right bank of the Volga.
If the occurrence of the chipmunk in the Trans-Sura forests is confirmed,
the range of the animal in the former Temnikov County is not separated
from its main range. All this should be further investigated.
In the Volga region the southern boundary reaches the Kama river, but
the animal apparently does not occur on the left bank of this river.
Farther east the boundary turns sharply south along the Belaya river. The
chipmunk is occasionally common in forests on the right bank of this river.
In the southern Urals chipmunks occur to 52°30'N. in the Troitsk District
and on the upper reaches of the Ik and Sakmara rivers. The distribution of
chipmunks in the Baraba Region is not clear. The animal was not found
in forests south of Kokchetav (the Borovskoe forestry). The southern
boundary of the range apparently-turns sharply north in this area, reaching
the Tara. It then passes near Omsk near the steppe. However, chipmunks
occur on the islands of Lake Chany. The boundary thus includes the
forests of the right bank of the Irtysh which are also inhabited by the flying
squirrel and the Teleut squirrel. Farther south chipmunks occur
in ribbon pine forests in the Semipalatinsk steppes, e.g., in the Srostki
pine forest. To the southeast the boundary continues to Lake Zaisan and to
the Mongolian Tarbagatai Mountains (the Malyi Oi Range), including a part
of the Greater Altai, the Tannu-Ola Range, Urianghai Territory [Tuvinian
ASSR] Hébségél Lake, Hentey Range. The boundary then turns sharply
north and skirts the steppes of Dauria. It crosses the Argun and runs east
through the Hsingan Range and Manchuria. The exact boundary in Manchuria
is unknown. I once obtained a specimen from the eastern part of the Kirin
Province.
The boundary then passes through northern and central China, not
including the Mongolian Desert and the Tibetan highland. It is not known
exactly how far south chipmunks occur in China. However they do not
occur in the Hunan and Kwangsi provinces, where they are replaced by
Tamiops macclellandi*.
Chipmunks are found on Sakhalin, Hokkaido and Iturup islands.
The distribution of the chipmunk in the Vologda Territory is incompletely
known. I.F. Brandt (1856) recorded it from the former Vologda Province,
where it apparently reaches the Vologda. A specimen from the former
Totma County is in the ZMAN collection. Evg. Debyuk (1920) did not find
the chipmunk in the Kostroma Territory. However, he mentions
geographical names which indicate that the chipmunk was known in this
area inthe past. For example, Debyuk mentions the Burunduchikha
{Chipmunk's] Brook in the Vetluga floodland. According to Zarubin (1856),
chipmunks occurred in the former Varnavino County. K.A. Vorob'ev
(1927) writes that it is found in the northern part of the former Kostroma
Province. B. Bogachev recently found it near the Katromskoe Lake, west of
the Vologda and north of the Kostroma. According to S. V. Lobachev
(1932), chipmunks are numerous along the Vychegda. According to old data
by Georgi they occur along the Northern Dvina, the Vyatka and the Kama.
I. F. Brandt (1856) writes that the range of the animal reaches the Mezen.
* See McAmicus Shih, Preliminary Report of the Mammals from Jaoshan, Kwangsi collected by the
Jaoshan Expedition, Bulletin of the Department of Biology, San Jatsen University, Canton, No, 4, 1930,
p. 6. Same author, Note on the Mammals from Southwest Hunan, Loc, cit, No, 9, 1930, p, 3.
404
Chipmunks, Upper figure — Eutamias orientalis Bonhote, Lower figure — E, s. sibiricus Laxm.
Drawing by V. A. Vatagin
466
In the Academy of Sciences are specimens caught in the vicinity of Mezen.
A specimen was recently caught in this locality by A.M. Sergeev (1935).
According to him, chipmunks are rare between Mezen and Kuloi, they
apparently do not pass to the western bank of the Kuloi. Chipmunks occur in
the basin of Vashka. In my collection is a skin from the Kholmogory
District (obtained by V. Ya. Parovshchikov). The chipmunk is certainly
widely distributed in the Pechora and Perm Territories. I found it east
of the town Cherdyn on the Polyudov Kamen Mountain.
Distribution of the chipmunk in the Gorki Region and in the Tatar
Republic. A.N. Formozov provided detailed information on the distribution
of the chipmunk in the former Nizhni Novgorod province. It occurs in the
northeastern part of this province, along the Vetluga river, and is well
known by the local hunters. Chipmunks are much rarer along the Kerzhenets,
but they occur on the right bank of this river. The unique discovery of the
animal in the Temnikov District of the Mordvinian ASSR has been mentioned
above.
The distribution of chipmunks in the Tatar Republic (former Kazan
Province) has been described by Prof. M.N. Bogdanov (1871). According
to him the animal is common in spruce forests. It is not found on the
right bank of the Volga. It is very common in old forests in mountain
passes along the Kazanka and Mesha, in the northern parts of the former
Kazan and Laishevo Counties. It was also found by Bogdanov along the
right bank of the Mesha river, in an oak forest near the village Devlezer'.
M.N. Bogdanov (1873) mentions that chipmunks occur in the former
Kozmodemyansk and Tsarevokokshaisk [Krasnokokshaisk, or Ioshkar Ola]
counties. According to new data by I. Bashkirov and 1I.D. Grigor'ev (1931),
it is found along the right bank of the Kama. However, the animal does
not reach the Kama Region in the south (near the western boundary of the
Tatar Republic). Large numbers of chipmunks were collected by the staff
of the Volga-Kama Commercial Biological Station of the Mozhga District
(former Yelabuga and Malmyzh counties of the Vyatka Province). According
to A.M. Korsakov (1933), chipmunks are common in the Mozhga District of
Udmurt (Votish) Region, in the former Yelabuga County and the southern
part of the Malmyzh County (former Vyatka Province). Only small numbers
occur. in the Mamadysh Canton (former Kazan Province), along the left
bank of the Vyatka, in the Lubyany forestry (on the boundary of the former
Vyatka Province). Chipmunks are common in the Storozhevsk District of
the Komi Region (former Ust-Sysalsk [Syktyvkar] County of the Vologda
Province) along the Vishera river, a right tributary of the Vychegda. The
chipmunk is very common in the Ugorskii District, in the upper reaches of
the Mezen river.
Farther east the chipmunk reaches the right bank of the middle reaches
of the Belaya. We found it in this region in the Angasyak forestry of the
former Birsk County (Ufa Province) and in large numbers in the deciduous
forest near the Dubrovka Village. It was also found in the middle reaches of
the Chusovaya river by N.E. Schultz (summer 1936). L.P. Sabaneev
(1874) wrote that the chipmunk occurs in the Bogoslovskii [Karpinsk] Urals,
in the Pavda woods, where it is more common than in the former
Yekaterinburg (Sverdlovsk) County. It is very rare in the southwestern
corner of the former Perm Province, and absent in the Shadrinsk steppe.
It is common in the former Perm Province. N.M. Dukel'skaya (1928) found
it in the Ilmenskii state reserve near Miass.
1254 406
Prof. Eversmann (1850) foundthat the chipmunk occurs in forests in the
Urals, between the Ik and the Samara. It is particularly numerous near
the village Miass. According to N.A. Zarudnyi (1897), it inhabits forests
of Orenburg Bashkiria, especially in mixed and conifer forests along the
upper reaches of the Ik, Sakmara, Belaya, and between Kaginskii Zavod
and Verkhne-Uralsk. The chipmunk is also common near Tastuba (former
Ufa Province).
A.I. Argyropulo (1930) writes that the southernmost record of the
chipmunk east of the Urals is from near the village Vikilovo, 150km south
of Ishim. To the west the boundary turns to Miass. It was found 80km
south of Zlatoust. In 1926 it was found in birch coppices near the village
Novinskoe (Troitsk District) near 52°30'N. This locality and the record
of N.A. Zarudnyi at the Sakmara river are the southernmost localities
in which the chipmunk occurs in Transural.
K.K. Flerov (1933) states that Eutamias is common in the northern
Tobalsk Ural, in the basin of the Sosva and the Lyapin, Chipmunks are
found here both in the taiga and in forests on the Urals. They are much
rarer in the basin of the Synya.
I. Ya. Slovtsov (1892) writes that chipmunks are common in the Tyumen
District of the Tobolsk Territory. They are rarer in the southern part of
this district than in its northern part. They are common in the former
Turinsk, Tobolsk, Berezovo and Obdorsk [Salekhard] districts. They are
rare along the Yurga river of the former Yalutorovsk District and do not
occur in the former Ishim and Kurgan districts. The range of the chipmunks
reaches the Tara District on the right bank of the Irtysh. According to
P. Stepanov (1886), it occurs in the forest-marsh between the Om, Tara
and Irtysh rivers along the Tartas, especially along the Tara. Chipmunks
have been recorded from the Surgut District, from the taiga near the
Barsovaya Mountain (S.M. Chugunov, 1915). The expedition of the
ZoologicallInstitute of Moscow University collected many specimens (1932-
1933) in the Surgut forests. Farther north chipmunks were observed by
K.M. Deryugin in the valley of the Ob river, near the Bolshe-Atlymskie
camps and the village Kushevat.
467 The chipmunk is sometimes common in the Tomsk Territory. It was
found near Narym (V.P. Anikin, 1902) and is very common along the
Chulym river (G.E. Iohansen, 1923). According to B.S. Vinogradov
(1924), it occurs in the vicinity of Barnaul and Ust-Kan.
Farther southwest it is foundnorthofIshim and possibly in forests near
Pavlodar and on islands in Lake Chany. Chipmunks in this region occur
together with species of the forest fauna (forest mice, etc.) which
survived the huge fire which separated Lake Chany from the northern
forests. These forests extended almost to Lake Chany in the last century
(M.D. Zverev andI.M. Zaleskii, 1935). Chipmunks occur in the Barnaul,
Srostki, Kasmalininskii ribbon pine forests, etc. East of the Ob the range
of Eutamias includes all forests along the tributaries of this river,
the Inya, Berd', Chulysh, etc.).
In the Altai Mountains the chipmunk is common along the Chulyshman,
near the Teletskoe Lake, in the Bele area, near Uimon, in the Seminskii
mountain pass, in mountains near Lake Tengi, and in the basin of the Biya
river, etc. (see N. Kashchenko, 1902). It was found near Lake Zaisan,
in forests near the village Maitabai and lake Marka-Kul (G.I. Polyakov,
1914). It occurs in the Sayan highland and Urianghai Territory [Tuva
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468
ASSR] (vide B.S. Vinogradov, 1927). It was observed near Lake Hébsdgél
along the Bilutyi, a left tributary of the Hara Gol river, in Mongolia (vide
N.F. Kashchenko, 1910). According to M.D. Zverev, the chipmunk occurs
rarely in the Kuznetsk Steppe and in the valley of the Inya river to the
forested slopes of the Salair Ridge, where it is common, In my collection
is a good series from near the village Monok, 150km southwest of Minusinsk
(by N.M. Dukel'skaya). It was recorded by I.N. Shukhov (1925) from the
Krasnoyarsk, Achinsk, Minusinsk and Yeniseisk Districts. It is widely
distributed in the Khakassian District (M.D. Zverev, 1930). In the Tungus
District the chipmunk is very common.(N.P. Naumov, 1934). According to
S.I. Orlov (1930), it occurs in the Verkhne-Imbatskoye, Turukhansk and
Karasino districts. In the basin of the Taz river it enters the forest tundra
to 66°30'N, near the Sidorovskaya Pristan [Harbor] (S.P. Naumov, 1931).
On the Turukhan it was found up to 66°N near Yanov Stan (S. P. Naumov,
1931). It was recorded by A. Middendorf (1867) from the vicinity of Igarka
(68°N). Farther east it occurs along Kureika river, in the region of the
Chirinda and Nekonda lakes, along the upper reaches of the Kureika, and
near Lake Ochata. The records of E.O. Yakovlev (1930) of the chipmunk
along the Medvezh'ya river as far as its lower reaches and along the Ayakli
and Kheta rivers to their estuary are incorrect. He informed me that
in the forest tundra region east of the Yenisei river only isolated specimens
occur at the southern slope of Lake Norilskoe. This is apparently the
northernmost record of chipmunks in this part of Siberia.
The chipmunk is not found farther east, along the Kheta and Khatanga.
Chipmunks are numerous in many localities in the Irkutsk Territory.
They were observed in the Ust Uda District, in the northern part of the
Irkutsk District, along the right bank of the Angara and near the estuary
of the Bol'shaya Yanda river (M.G. Bakutin, 1930). They are widely
distributed in forests of Transbaikalia, very numerous in the Barguzin
forests (G.G. Doppel'maier, 1926). They are common in the
Troitskosavsk [Kyakhta] District and occur in eastern Transbaikalia, in
the taiga, near the villages Monastyrskoe, Zyul'zikan and Kyker (B.A.
Kuznetsov, 1929). Farther south they were found near Urga [Ulan Bator],
Uliassutai [Jibhalanta], (Gl. Allen, 1925), Sain Noyan Khan and on the
eastern slopes of Great Khingan (a specimen collected by A.S. Lukashkin).
Chipmunks are widely distributed in Manchuria (A. Sowerby, 1923), in the
Ussuri Territory, Amur Region, on Sakhalin and the Shantar Islands. In
Yakutia they are numerous along the Lena and Vilyui rivers (R. Maak,
1859, 1886, and according to recent collections). They occur in the
Olekma-Vitim highlands (I. Polyakov, 1873), in the valley of the Yana to
its estuary, at 69°N (A. Bunge, 1887). They are found along the Indigirka,
the Kolyma to its estuary, the Anadyr and farther south along the Penzhina.
They have not been found in Kamchatka.
A. Samorodov recently informed me (in litt., 1936) that the chipmunk is
rare near the Olyutorski Gulf, according to Chukchis, Lamuts and Koryaks.
The animal is called ''pitan'’ in the Chukchi language.
The chipmunk occurs along the upper reaches of the Apuka and Pakhacha
and their tributaries, both in deciduous forests with alder, willow and
poplar and more frequently in dwarf cedar forests on mountain slopes. The
chipmunk is unknown to the Russian population. A. Samorodov put at my
disposal two skins from these areas. They belong to the Yakut race
according to their color.
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470
The record of the chipmunk in this region moves the boundary
considerably towards the east.
MODE OF LIFE. The habitats of the chipmunk are closely connected
with the forest. At the southern boundary of its range it is found in isolated
forests near the steppe, e.g., in birch coppices in the Baraba steppe. At
the northern boundary of its range it enters the dwarf forest along the tundra,
but it is very rare there.
In Bashkiria and the former Birsk County of the Ufa Province I found it
in deciduous undergrowth of linden, oak, maple, birch mixed with hazelnut,
rowan and European dewberry, on the border of millet fields. It could be
found in old windfall at the borders of millet fields. According to
unpublished data by S.S. Donaurov and V.P. Teplov (''Material on the
Biology and Commercial Exploitation of the Chipmunk" (Materialy po
biologii i promyslovomu ispol'zovaniyu burunduka, 1936)), the chipmunk
is restricted to fir and larch forests. It is common in spruce-fir
forests, but definitely avoids pine forests.
According to 1.D. Kiris, chipmunks occur sporadically in West Siberia
in the Surgut District in urmans* (cedar forests). In fir groves and
spruce forests the chipmunk is much rarer, apparently inhabiting areas
near cedar forests, as wellas dry ridges. It is even rarer in pine forests.
The chipmunk is numerous in birch forests, especially in forests with
much old windfall.
According to G.A. Velizhanin (1931), the chipmunk regularly occurs in
pine forests of the Baraba steppe, in the Kiprinskii District of the Kamenka
Region. Its prefered habitat in this region is shrubs among birch and
aspen. It is very fond of European bird cherry in river valleys. Chipmunks
occur frequently on low ridges and dry steppe coppices.
According to M.G. Bakutin (1930), the favorite habitat of the chipmunks
in the Irkutsk District are taiga windfall areas, mountain brooks and
plantations of cedar, larch and spruce. The animals are much rarer on
mountain ridges than in the foothills. They occur near fields of different
crops, in coppices, and among isolated shrubs of European bird cherry.
They are apparently attracted to these habitats by the cereal crops. They
sometimes occur in small forests and enter valleys with shrubs and trees.
Chipmunks are found everywhere in the Irkutsk Territory, changing their
habitat according to the food available.
According to B.A. Kuznetsov (1929), chipmunks in Transbaikalia inhabit
both coniferous and deciduous forests. Near the village Kyker they occur
in the mountain and lowland taiga, most commonly in thickets near streams
and brooks. Near the village Zyul'zikan, chipmunks occur in young birch
forests along the Zyul'zya and Zyul'zikan rivers. They are especially
numerous in birch-grown ravines on the northern slope of the Undin Range.
According to K.A. Plyater-Plokhotskii (1932), the typical habitat of the
chipmunk in uninhabited localities in the Maritime Territory is thickets of
European bird cherry, currants (Actinidia kolomienta Max.), Amur
grape (Vitis amurensis Rupr.) and other trees and shrubs growing at
the banks of forest streams and brooks. In large forests chipmunks mainly
occur among windfall, and their burrows are under fallentrees. They
occasionally settle in screes and in fields among forests. The chipmunks
are certainly attracted by plentiful food.
* [Translator's note: 'urmans''— dense, marshy coniferous forests in the taiga].
409
FIGURE 217. Postures of Ural chipmunks
(Sketches from nature by K. K. Flerov)
The chipmunk is an elegant animal. It is timid, but not too cautious, and
it is very curious. It runs rapidly, but is more at home among windfall
branches. It climbs rapidly on trees, running from one branch to another.
S.S. Donaurov and V.P. Teplov (in litt., 1936) write: 'The chipmunk
climbs very well with its head up or down. It easily jumps to the lower
branches of trees 5-7m apart and changes direction of its leap in midair
with the help of its tail. We know of instances in which chipmunks jumped
from the top of spruce trees 30m high tothe ground. They stretched
their legs in the air and resembled small flying squirrels. In autumn, in
the season of storing food, we saw chipmunks on the thinnest ends of
branches of maples, lindens and rowan trees. The animals often held their
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471
heads down and clung to the branch with their hind feet only.'' When man
appears, the animal gives a feeble ''gurgling'' cry, and then loud jerky
"chirping'’*. When the danger is remote (e.g., a man approaches from
afar) the chipmunk whistles jerkily and then keeps quiet for a long time.
During this silence it rises on its hind legs and carefully contemplates
the approaching danger. If one walks quietly and without sudden movements,
one may approach the animal to within 15-20 steps. It then rushes away
repeatedly giving its whistle. If one hears the whistle of a chipmunk in the
forest, it is easy to decide whether it is sitting quietly in one place or
running away. Frightened chipmunks do not always run to their burrow.
They usually hide in windfall or brushwood, or run on the branches of a
tree to hide among the dense foliage.
The chipmunk is a diurnal animal. In BashkiriaI saw it most frequently
in the morning when it became warm and in the evening, before sunset.
The same was observed by B.A. Kuznetsov (1929) in Transbaikalia. K.A.
Plyater-Plokhotskii (1932) gives the following description of the daily life
of chipmunks in the Maritime Territory: ''The daily life of the
chipmunks varies according to the season. In spring they come out of the
burrow only when the sun has warmed the ground sufficiently. Their main
activity lasts from 10 a.m. to 3.30-4 p.m., i.e., the hottest hours of the
day. No chipmunks remain outside by sunset. They also remain in their
burrows in windy and rainy weather.
"In summer they come out before daybreak as soon as the first signs
of light appear. They then jump from one branch to another, sit on
their hind legs and wash themselves with their forefeet, descend along
branches and upturned trees to a river or stream to drink and feed.
Between 12-2 p.m. the animals disappear and their whistling is not heard.
They take shelter in the burrows during the hot hours, come out again
when the heat subsides, and turn in at sunset. In the dense, shady and
cool Ussuri taiga the chipmunks are active throughout the day, without an
interval at noon.
"Inautumnandon overcast days in summer chipmunks stay outside from
sunrise to sunset. Rainy weather usually keeps them in their burrows,
but in the Ussuri Territory they are seen even during rain. This may be
due to the great number of rainy days. The sounds made by them before
rain differ sharply from their ordinary whistling. This is so well known in
the Ussuri Territory that 'root hunters' (collecting ginseng roots) predict
rain from these sounds."
Burrows and nests. The chipmunks build their burrows between roots of
trees, under pieces of rock or similar shelters. All burrows found by
S.S. Donaurov and V.P. Teplov (in litt., 1936) in the Volga-Kama
Territory were on dry hillocks. The entrance to the burrows was usually
at the base of tree stumps or near small mounds. The burrow usually
descends at an angle of 45° immediately after the entrance. It then
continues horizontally and ends in a round chamber which contains the nest.
* Chipmunks make their peculiar "gurgling" sounds warning others of danger already during their first months
of life, S.S. Donaurov and V.P. Teplov (in litt. , 1936) studiedthe daily life of young (2-2,5-month-old)
chipmunks in open-air cages in the forest. They repeatedly observed that the chipmunks have a peculiar
system of signals. All five specimens in the cage were usually active for many hours after feeding,
climbing on trees and running among fallen branches in the cage. However, as soon as one of the chipmunks
saw a large bird, even at a distance, or a passing man, it emitted a quiet "gurgling". Then, all chipmunks
stopped and remained motionless for two to three minutes,
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Side galleries are rare and there are never more than three. The burrows
usually have one exit. The galleries are 60-400cm long, and the chamber
is 20-35cm wide and 40-95cm underground. Some burrows have small
corridors filled with excrement. Food reserves are kept in chambers,
rarely in side galleries. The nest is also in a chamber and it consists of
dry leaves of linden, maple, fern, raspberry and aspen. Some nests are
made of moss (Hypnum) and scales of pine bark. The nest is round and
fills the whole space of the chamber in which there are no food reserves.
M.G. Bakutin (1930) gives the following description of chipmunk
burrows in the former Irkutsk District: ''There is usually one exit which
is covered by vegetation. From the exit runs a gallery, 5-6cm wide.
At a distance of 40-50cm from the entrance is a round depression not
wider than 30-35cm, which is the storeroom. This is connected by a
straight, 40-cm-long gallery to a similar depression which contains the
nest."'
M.G. Bakutin observed one or two blind side galleries from the main
gallery onthe same level. They were 20-30cm long and were full of
excrement.
According to Bakutin, the nest itself is lined with leaves of birch,
aspen and grass. In burrows in fields the nest is lined with broken stalks
of rye and wheat, and empty ears of cereals.
B.A. Kuznetsov (1929) gives the following description of summer
burrows of chipmunks: ''The structure of winter burrows differs from that
of summer burrows which are very simple. They begin with a round
opening, 4-5cm wide. The main gallery descends into the ground and ends
in a round chamber usually without any padding. The burrow is 50-75cm
long. The beginning of the main gallery sometimes has a small side
gallery."
K.A. Plyater-Plokhotskii (1932) gives a similar description: ''The
chipmunks usually dig their burrows among roots or under fallen trees.
Sometimes there are whole chipmunk ''towns'' in screes. Some burrows
begin in a rotten tree stump and continue deep under ground. It is difficult
to find burrows, as they are usually concealed by overhanging stones, or
roots, or are inthe middle of vegetation. Burrows are rarely made in
hollows of live trees, but sometimes in those of fallen trees. The entrance
of the burrow is round or oval, not much more than 4cm wide. The first
part of the burrow descends from the opening at an angle of 45° to 70°.
After two or three bends the burrow ends in the nest chamber. Two or three
side galleries branch from the main gallery. Some of them contain only
excrement. Others are widened at the end into large storerooms. Usually
there are no other side galleries between those ending in storerooms and the
nest chamber. The main gallery is sometimes widened into a storeroom
before the nest chamber, and then narrows again until it reaches the nest.
In this case there is only one side gallery which serves as a latrine. On
plots among isolated trees and shrubs there are burrows which are
apparently temporary. They are usually 60-70cm below the ground, short
and without storerooms. The main gallery bends once or twice and ends in
a nest chamber. We found some permanent burrows with two storerooms.
"The depth and length of the burrow depend on the soil. Burrows in sand
usually have a gradual slope, are 70-80cm deep and 2.5-3m long. Burrows
in loamy soil have a steeper slope (70°), and are 1.5m deep and 2-2.5m
long."
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FIGURE 218, Chipmunk caught in the valley of the Belaya in Bashkiria. The cheek pouches are filled
Food. The chipmunk is mainly vegetarian. Animal food is a less
important part of its diet. The main information on this subject is given
by S.S. Donaurov and V.P. Teplov (1936) who studied the food of
chipmunks in the Mozhginsk District of the Votish Region (former
Yelabuga County and southern part of the Malmyzh County, former Vyatka
Province).
They examined the content of the cheek pouches of 324 specimens
caught in the field. The food consisted of vegetable matter, in 97.8% and of
animal matter in 30.9%.
Only 0.3% of the vegetable food consisted of seeds of cultivated plants.
This is strange because the cheek pouches of specimens caught in autumn
near the Belaya river (former Birsk County) were stuffed with millet seeds.
According to S.S. Donaurov and V.P. Teplov, wild plants eaten by the
chipmunk belong to two groups. The first group includes the following
trees and shrubs: rowan, elm, smooth-leaved elm, hazel, maple, linden,
European bird cherry, raspberry and honeysuckle. The seeds of these
plants are found constantly in the food stores of the chipmunk. The
second group consists of seasonal plants, berries or seeds of forest plants:
strawberry, bilberry, Paris, bramble, sedge, asarum, Oxalis, Melica
nutans, tall buttercup (Ranunculus acris), crowberry, aconite and
Spindle tree.
Mushrooms are not eaten by chipmunks.
The main animal foods of the chipmunk are invertebrates (30.3%).
Vertebrates constitute only 1.5%. Intwo cases remains of the bank vole
(Evotomys glareolus) were found in its stomach. Remains of
Lacerta agilis were found twice, and those of a small bird, apparently
a nestling of Fringillidae were found once. The chipmunk rarely eats eggs
of birds, e.g., of thrushes.
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FIGURE 219. Head of Bashkirian chipmunk, cheek pouches filled with millet.
The most common invertebrate remains are those of insects (18.2%),
followed by those of snails (12%). Helix and slugs (Arion) are also eaten.
Of insects, mostly ants are eaten.
S.S. Donaurov and V.P. Teplov estimated the weight of the average
daily food intake. They found that the animal consumes 7.5 to 15g of seeds
and 55-94 g of berries.
According to M.G. Bakutin (1930) chipmunks in the Irkutsk District feed
on cedar nuts soon after coming out of hibernation. In the middle of June
this author caught several chipmunks with cheek pouches filled with seeds
of Geranium pseudosibiricum, wild rose and seeds of sedge,
crowfoot, etc. Seeds of Carex pediformis were commonly found
in cheek pouches. This is due to the fact that this plant grows on slopes
in thin forests, windfall, etc. The favorite grains are wheat and rye.
Oats and hemp are taken less readily by the chipmunk.
In Transbaikalia B.A. Kuznetsov (1929) found larch seeds and seeds of
various forest herbs in the cheek pouches. He rarely found remnants of
insects, usually of beetles.
In autumn chipmunks are found in large numbers near fields at the edge
of the forest, because of the ripening grain crops.
According toI.M. Zalesskii and M.D. Zverev (1935), chipmunks feed
on buds of birch, larch and other trees in spring. When the snow melts,
they begin feeding on old cedar cones, young shoots of grass and seeds of
forest sedge. The diet becomes more varied in summer and the animal
eats rowan berries, cranberries, red bilberries, European bird cherries
and raspberries. When mushrooms grow, the chipmunk, like the squirrel,
fixes them on branches, The chipmunk is fond of seeds of wild forest
grass, Umbelliferae, seeds of sedge, wild millet, species of Polygonum,
crowfoot, Thymus serpyllum, tufted vetch, etc. In Siberia, pine
nuts are the favorite food of the chipmunk in autumn. In the Far East the
animal is fond of pine nuts, acorns and linden seeds. According to K.A.
Plyater-Plokhotskii (1932), the chipmunk eats the following cultivated
plants: wheat, rye, barley, buckwheat, oats, corn, flax, millet,
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sunflower, cucumber, squash and eggplant, watermelons, melons and
gourds. In orchards it destroys plums, cherries, apricots, pears and
apples.
European chipmunks occasionally eat animal food. They eat insects and
their larvae, they may attack frogs (according to Korsh) and even lizards
(according to observations of A.M. Nikol'skii on Sakhalin).
In late July or early August, according to the locality the chipmunks
begin to collect stores for the winter. The food is carried into the burrow
in the cheek pouches. The rodent bites off a stalk, holds it between its
forefeet, rapidly extracts the seeds and puts them into the cheek pouches.
As the cheek pouches become stuffed, the whole head becomes round and
large (Figures 218-219).
The chipmunk stores in its burrow various plants in the area. According
to S.S. Donaurov and V.P. Teplov (in litt., 1936), food reserves of the
chipmunk include seeds of the following plants: 1) maple; 2) linden; 3)
rowan tree; 4) hazelnut; 5) European bird cherry; 6) raspberry; 7) elm;
8)Ulmus scabra. Some chipmunks also store seeds of sedge,
Actaea, and honeysuckle.
According to Donaurov and Teplov, the stores are small, at most 1,480g.
In the Siberian taiga the chipmunks hoard pine nuts, seeds of larch and
grass, as well as buds of trees (e.g., birch buds). K.A. Plyater-
Plokhotskii (1932) observed in the Far East that burrows rarely contain
food reserves of one chipmunk. The burrows usually contain a large
variety of food. He found corn, wheat, barley, buckwheat, sunflower
seeds, hazelnuts, acorns and dry mushrooms. Each kind of food was
stored separately.
The size of the food reserves varies. K.A. Plyater-Plokhotskii writes:
"The largest food reserve weighed 6 kg and consisted of 3,650 ghazelnuts,
1.5kg of wheat, several sunflower seeds, acorns, dried crab apples and
corn seeds.
"Reserves of 1.5-2kg are common. In some villages of the former
Amur District peasant children dig up chipmunk burrows to get the nuts.
In fairs, nuts collected by chipmunks are more expensive than those
collected by man, as chipmunks never collect spoiled nuts."' M.G.
Bakutin (1930) excavated chipmunk burrows and found up to 1kg of rye and
wheat.
The observations of K.A. Plyater-Plokhotskii prove that the chipmunks
make long trips to collect food, sometimes a kilometer or more from the
burrow.
In one burrow K.A. Plyater-Plokhotskii (1932) found 1kg of rye, 500g
of buckwheat, 500g of corn and some sunflower seeds. The nearest corn-
fields were more than 1km from the burrow.
REPRODUCTION. According toI.M. Zalesskii and M.D. Zverev (1935)
chipmunks breed once a year, and a litter consists of 4 or 5 young. The
young are born naked and blind. According to 8.8. Donaurov and V.P.
Teplov (1936), gestation in the Volga-Kama Territory lasts 35-40 days.
There are 3 to 10 embryos (5.57 on the average). Unfavorable years cause
mortality of embryos through resorption. This was also established for
squirrels and ground squirrels. Only indirect data were obtained for
chipmunks. A marked retardation of growth of some embryos was observed.
The reproduction rate of chipmunks fluctuates widely, due to variations
in the number of embryos. Sterile females are rare.
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S.S. Donaurov and V.P. Teplov (in litt., 1936), studied the sexual life
of chipmunks. They found that burrows are occupied by hibernating
couples in autumn. In the Volga-Kama Territory chipmunks mate in late
April. They possibly mate before coming out of the burrows. The mating
period ends in early May. The authors state that the chipmunks do not
show great excitement during copulation, and males do not chase each
other or fight. According to Donaurov and Teplov the suckling period lasts
28 to 30 days, and there isonly one litter in spring. After coming
out of the burrows the young grow rapidly and reach their full size by the
end of July. Maturity is reached during the first year, in about the eleventh
month.
G.A. Velizhanin (1931) gives a description of life in spring and mating
of Eutamias in West Siberia. The animals come out of hibernation on
the 1st of April. For several days they show little activity, climb trees and
sit there for hours, basking in the sun. Several days later, with the
beginning of spring, they become much more active. One frequently hears
their cries at this time and sees males fight and chase the females.
Copulation usually takes place on atree. Afterwards the animals remain
motionless for 30 to 60minutes, only making writhing movements with
their tails. The young appear in West Siberia in May, and in July they
are already independent.
According to M.G. Bakutin (1930), estrus begins in late April in the
Irkutsk Territory. The young are born in late May or early June. They are
suckled during June. In early July they come out of the burrows and feed
on red currants and seeds of sedge (Carex pediformis). Young born
near fields feed on rye and wheat seeds. The female guards the young
during the first days; at the sight of danger she emits a peculiar chirping
cry which is answered from all sides by the young rushing to hide.
Bakutin thinks that chipmunks have only one litter per year.
According to K.A. Plyater-Plokhotskii (1932) chipmunks have estrus
in late April and early May. During estrus the males fight and chase each
other, emitting shrill whistles. The young of chipmunks in the Maritime
Territory are born in late May or early June, a litter consisting of 4 to 6
young (10 to 12, according to some information). K.A. Plyater-
Plokhotskii observed only one litter in spring. According to information
from local people, the chipmunk has 2 or 3 litters a year, but this seems
doubtful. G. Radde (1862-1863) writes that there are two litters, in May
and August. B.A. Kuznetsov (1929) states that his observations confirm
Radde's statement, but he does not explain this. B.S. Vinogradov and
S.1. Obolenskii (1927) also write that there are two litters, in June and
August.
To judge from our data and from observations on ground squirrels, the
chipmunks have only one litter in spring. *
Sex ratio. S.S. Donaurov and V.P. Teplov (1936) studied a large
amount of material. They found that there are more males than females,
i.e., 60% males to 40% females. Males predominate among the newborn,
constituting 80%. Natural mortality gradually levels this off. Most of the
* AN. Formozov informed me that he heard cries of females in the Ussuri Territory in late June, the same
as those made by rutting animals in spring. In A.N, Formozov's opinion, the chipmunks have estrus
only once, and the sounds mentioned may only be manifestations of a ''false estrus".
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young hibernate in couples. Most of the single males die because they
remain active in the bare forest.
MOLT. Chipmunks molt apparently only once a year. The molt begins
in June and ends between July and the middle of September. The molt
apparently takes place in the following order. Molt begins in the middle of
the back, where the skin shows an irregular dark patch. Darkly pigmented
stripes often appear exactly beneath the dark stripes of the fur. In some
specimens molt begins in the lower posterior part of the back like in
squirrels in autumn. Sometimes the whole back molts at the same time.
According to S.S. Donaurov and V.P. Teplov (in litt., 1936), adult
males in the Volga-Kama Territory begin molting at the end of the breeding
period, i.e., in the last ten days of April. The most intensive molt is
observed between the end of May to the
end of June. Some males which had not
yet begun molting were caught in the
middle and end of May. A specimen in
the initial stage of molt was collected
in the first days of August.
Molt may last 50 days in adult males.
The females begin molting almost at
the same time as the males, i.e., in
the last days of April. Some specimens
begin molting in early May. Molt is
slow and becomes more intense only after
lactation (from the beginning of July).
Females finish their molt early in
August. Molt thus lasts 90 days.
The young begin molting before
coming out of the burrows. Molt in most
intensive in the last days of July and
ends in the middle of August. Skins with
signs of molt are found until the end of
September. In Siberia the beginning of
molt is much delayed in females, and
ends in late September. Young molting
specimens are found until September-
October.
MIGRATIONS. The chipmunks may
make long migrations, usually in
search of better food. Hunters report
that chipmunks may be observed in late
April migrating over the frozen snow
from the taiga to the forest bordering
cultivated areas.
M.G. Bakutin (1930) describes
migrations in the forests of the Irkutsk
FIGURE 220. The inside of the skin of a molting belies /From ey toyeeide eune
PSPs ratics roheswiStrec a ite Seen OMe pr chipmunks lived on slopes of mountains
1932) near the Bolshoi Tangat river. In the
Drawing by A. N. Formozov second half of June the chipmunks began
to leave these places to reach the forested
valleys and feed on red currants. They remained there until the middle of
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August and then returned to their original habitat, except for a few which
apparently had decided to remain in the forested valleys during the winter.
They collect food, e.g., berries of red and black currant and seeds of
grasses.
In 1925 migration of the chipmunks was large-scale and caused great
damage. The animals left the taiga, migrated to the fields and destroyed
the crops. Wheat and rye-were severely damaged, oats and hemp less.
This migration was apparently caused bya severe failure of the crop of pine
nuts."
K.A. Plyater-Plokhotskii describes a migration of chipmunks: ''In
August 1927 there was a great flood in the Maritime Territory. Small
forests near cultivated land in the valley of the Iman were invaded by large
numbers of chipmunks in the first days of August. All chipmunks
disappeared suddenly, and moved to the mountains. On 12 August, the
valley was flooded. If the chipmunks had not left at this time, most of them
would have been drowned. The same valley was inhabited by large numbers
of other small rodents. When the river overflowed and carried away the
sheaves of wheat, many of these were literally covered with field mice,
rats and meadow mice, but no chipmunks were among them. "'
HIBERNATION. In West Siberia and the northern part of the European
U.S.S.R. chipmunks hibernate from late September to early October.
East European chipmunks apparently hibernate at the same time. K.A.
Plyater-Plokhotskii (1932) writes on the hibernation of chipmunks in the
Far East: ''From the beginning of October, chipmunks become less
common. Inthe middle of the month they go into hibernation. In the
southern part of the Maritime Territory chipmunks may sometimes begin
to hibernate only in November. In mild, warm winters chipmunks may be
found outside their burrows, even in December."’
S.S. Donaurov and V.P. Teplov (in litt., 1936) observed in the Volga-
Kama Territory that burrows are occupied by couples in autumn (see above).
In their opinion, chipmunks are monogamous, and the male apparently
leaves the female only during suckling, rejoining her afterwards. These
habits, especially the hibernation of couples, should be further studied.
DAMAGE. The chipmunk may cause great damage to grain crops,
especially in years with crop failures of cedar nuts or other food plants
of the taiga.
I.M. Zalesskii and M.D. Zverev (1935) write: ''The chipmunks settle
near the fields and begin their harmful activity immediately after sowing.
The animals dig out the seeds atthe edge of the field, and occasionally
cause great damage. When the grain begins to grow the animal bites off
the young stalks. However, the greatest harm is caused when the grain
ripens and the animal begins to store food for the winter. It spends all
day travelling from the burrow to the field and back. It comes out of the
forest early in the morning. If there is no danger, it runs to the wheat
fields where it may hide from enemies. It leaves spoiled or small ears
intact. If it finds a good, full ear, it climbs the stem, takes it between
its teeth and bends it to the ground. It cuts the stalk near the ear, takes
this into its paws and picks out the grain which it stuffs into its cheek
pouches. It then runs to the burrow and puts the grain into the storeroom.
The chipmunk is intelligent. When the crop is very dense and it cannot
bend the stalks to the ground, it adopts a different method; it cuts the stalk
into parts until it reaches the ear. It may sometimes cover 1km or more
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from the burrow to the field. The cheek pouches usually contain only 8-
9 grains. It has thus to run repeatedly to the field in order to obtain
sufficient grain for its stores."
Millet fields were severely damaged in the middle reaches of the Belaya
in Bashkiria (former Birsk County of the Ufa Province). The cheek pouches
of specimens caught were stuffed with millet grains.
The Birobidzhan District (Far Eastern Territory) suffered an invasion
of chipmunks in autumn 1926, in which 70-80%, sometimes 100%, of the
crops were destroyed. The animals settled in the fields in stacks of wheat.
K.A. Plyater-Plokhotskii (1932) reports that a blow with a stick drove 10-
15 chipmunks out of the shocks. In 1927 a mass invasion of fields on the
border of the forest in the Iman District, Far Eastern Territory, forced
the peasants to abandon their fields. In 1925 there was a mass migration
of chipmunks from the mountain taiga of the Ust Uda District (Far
Eastern Territory) to the grain crops in the valleys. Wheat and rye fields
suffered most, oat fields and hemp plantations less.
I.M. Zalesskii and M.D. Zverev (1935) write: 'In central and western
Siberia chipmunks cause damage, though not very severe. Damage to
grain crops was caused in the Narym Territory, the northern part of the
Baraba Steppe, the northern part of the Tara District, the Achinsk District
and adjacent districts, and to some extent in the Minusinsk Territory.
Concentrations of chipmunks were observed in birch coppices near oat
fields. Their burrows were found to contain grain stores, mainly oats but
also wheat and rye. In some places stalks of oats were cut 1 2 yvershok*
above ground, and the grain had been taken. In another place, oat fields
on the border of the taiga were severely damaged...Chipmunks were also
found to cause great damage to shocks of rye, andtheir food stores contained
seeds of hemp and flax. "'
Chipmunks also damage gardens and orchards. Cucumbers, eggplants,
marrows, corn, poppy, peas, beets, carrots and potatoes suffer from the
raids of chipmunks. The last three vegetables are readily eaten by
chipmunks in captivity. Chipmunks climb the stalks of sunflowers and pick
out the seeds, fill their cheek pouches and carry them to the nest. They
also damage melons, gourds and watermelons. They have been reported
to have damaged cherries, apples and pears.
Methods of control are little developed. Chemical methods may be used,
whereby sheaves of wheat are treated with natrium arsenite. K.A.
Plyater-Plokhotskii (1932) reports that peasants control chipmunks by
placing cats into their habitats, especially into those of gravid females.
Kittens born in the forest quickly learn to hunt chipmunks and kill large
numbers, but this method gives only temporary results.
ENEMIES. According to S.S. Donaurov and V.P. Teplov (in litt., 1936),
buzzards and sparrow hawks are enemies of the chipmunk inthe Volga-Kama
Territory. The chipmunk is an important part of the diet of the buzzard
during the breeding period, and 30% of its food consists of chipmunks. The
following mammals hunt the chipmunk: hamster, weasel, ermine, marten
fox, wolf, rat, polecat and badger. The black hamster which is widely
distributed in forests of the Gorki Region and Bashkiria is a great enemy of
the chipmunk. It not only robs the stores of the chipmunk, but also eats the
chipmunk if it does not succeed in hiding.
* [Vershok — 4.445 cm. ]
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According to M.G. Bakutin, who studied the chipmunk of the Irkutsk
Territory, its main enemies are the ermine, the ferret and the bear.
During the autumn hunt of ermine and Siberian ferret, hunters often find
them in the burrows of chipmunks which have often already been eaten.
Where chipmunks are numerous bears hunt them and excavate their
burrows. The bear eats the food stores of the chipmunk, and also eats
the chipmunk. In the Far East the chipmunk is also hunted by weasels,
raccoon dogs, foxes, wolves, and occasionally by snakes (Elaphe
schrenckii). It is also hunted by birds of prey like the goshawk.
COMMERCIAL VALUE AND TRADE. Chipmunk pelts have only recently
appeared on the fur market. It has a beautiful pelt which can be used for
fur coats.
The chipmunk is caught by three main methods: 1) by shooting with the
help of dogs (especially eskimo dogs), 2) with decoys; 3) with traps.
V.V. Gubar' (1930) describes hunting with a lure in the Upper Ob
forest (former Biisk County). ''Chipmunks are hunted with decoys in April.
In clear weather the hunter takes a special whistle and a long rod witha
loop of horse hair. He settles in a glade and, with the whistle made of a
cartridge, makes sounds imitating those of a rutting female.
"The cry of the rutting female is a characteristic sound quite different
from the ordinary ‘chirping. '*
"Allmalesinthe vicinity are attracted by this mating call and run towards
the hunter, ignoring even cold water. The excited males swim across
streams of icy water and even in the presence of man continue trying to
find the female. The male waves its tail, chirps, jumps on trees and into
the grass, approaching the waiting hunter.
"The animal approaches to within two or three steps from the source of
the call. It pauses and sits on its hind legs. At this moment the hunter
slips the loop over the head of the animal, pulls the rod and the chipmunk
is captured."
Hunters may catch 39-40, and even 100 specimens in a day, by this
method.
The Yenisei Tunguses and the Ob Ostyaks and Shorians use bows. The
metal arrowheads are replaced by blunt heads, because sharp arrowheads
spoil the fur. The bow is large, as tall as a man, and the string is made
of a strong, twined packthread. The hunters shoot with great accuracy;
the blunt arrow, if shot from a short distance stuns the animal, or kills it.
Dogs are necessary in this primitive hunting, to drive the animals on to
tie trees.
West Siberian hunters have recently introduced a special trap for
chipmunks which they call "lobster pot'' or ''muzzle''. The principle of this
trap is that of fishing creels used in the north of the Narym Territory, in
the Achinsk taiga and on the Altai. These traps are used in spring, during
the rut period. A female is placed in a small cage which is placed inside
the creel. The trap is arranged on a fallen tree, and the hunter hides
nearby, calling the males with a whistle, imitating the voice of the female.
The males enter the creel and on a good day as many as 50 specimens can
be caught.
* I,M, Zalesskii and M.D. Zverev describe this cry as a short whistling differing sharply from the cry of the
male which consists of repetition, at intervals of 10-15 seconds, of "'tyurlyu-tyurlyu".
420
The fur of the chipmunks is of high quality in spring as the molt begins
in summer, in females later than in males.
ECTOPARASITES. According to K.A. Plyater-Plokhotskii (1932) the
chipmunk is parasitized by mites (species not recorded) and fleas
(Ceratophyllus tamias) (identified by Prof. Yu. N. Wagner).
Subspecies of Eutamias sibiricus
The subspecies of Hutamias sibiricus are not well differentiated,
because they differ mainly in the variable color characters. Cranial
characters give no reliable means of distinguishing the subspecies.
However, the following races can be differentiated in series.
377. Eutamias sibiricus sibiricus Laxmann (1769). Central
Siberian chipmunk
1769. Sciurus sibiricus Laxmann ME , Sibirische Briefe, Géttingen und Gotha, S. 69-71.
1859, Tamias pallasi BairdS.F., Mammals of North America, Philadelphia, p, 295.
1867. Tamias striatus Middendorf A. Th,, Sibirische Reise, Sdugethiere, S. 83.
1880. Tamias pallasi Tell in Nordenskjold, 'Ekspeditsiya k ust'yu Eniseya'"' (An Expedition to the
Estuary of the Yenisei), p. 178.
1884. Tamias striatus Nikol'skii A.M., Puteshestvie v Altaiskie gory (A Journey to the Altai
Mountains), Trudy Sankt Peterburgskogo obshchestva estestvoispytatelei, XI, p. 174.
1899. Tamias pallasii Kashchenko N.F., Rezul'taty Altaiskoi ekspeditsii (Results of an Expedition
to the Altai), p. 60.
1900. Tamias asiaticus Kashchenko N.F., Opredelitel' mlekopitayushchikh Tomskogo kraya (A Key
to the Mammals of the Tomsk Territory), pl. 32.
1912, Eutamias asiaticus altaicus Hollister N., Five New Mammals from Asia, Proceedings of
the Biological Society of Washington, p. 182; Mammals Collected by the Smithsonian, Harvard
Expedition, Proceedings of the U.S. Natural Museum, 1913, p, 522 (collected near Tapuchi).
1930, Eutamias asiaticus asiaticus Zverev M.D., Obzor gryzunov yuzhnykh chastei Minusinskogo
i Khakasskogo okrugov (A Survey of Rodents of the Southern parts of the Minusinsk and Khakassian
Districts), Sbornik Srednesibirskogo geograficheskogo obshchestva, p, 83.
TYPE LOCALITY AND DEPOSITION. Vicinity of Barnaul. Holotype collected by Laxmann, apparently
lost,
MATERIAL EXAMINED: 180 skins, mostly with skulls.
DIAGNOSIS. Color saturated, without the yellow tones typical of the
West Siberian subspecies (see below). Anterior part of back rust-yellow-
gray. Black, relatively broad stripes on the back. Buttocks with an
intense rust-red-yellow tinge (between pinkish cinnamon and cinnamon).
Top of head more rust-colored than in West Siberian chipmunks, e.g., in
the Surgut form (see below). Rust color of head mixed with many
chestnut-black hairs which give a darker color.
Length of skull 37.6-41mm (M.39.1mm); condylobasal length 33.9-
38.5mm (M.35.8mm); interorbital space 9.3-11.1mm (M. 10.2 mm);
zygomatic width 20.8-23.1mm (M.22.6mm); length of upper tooth row
6-7mm (M.6.4mm); Length of body 127-168mm (M.149.2mm); tail 79-
115mm (M.106.8mm); hind feet 31-37mm (M. 34.8mm).
SYSTEMATIC NOTES. The Siberian chipmunk is named Eutamias
asiaticus Gmel. (1788) in most publications. Its exact range is given
by I. F. Gmelin as Northern Dvina and Kama. The name asiaticus is
therefore applicable to the European-West Siberian race of Eutamias.
Gmelin's description had been preceded by a forgotton description in
421
485
M.E. Laxmann's ''Sibirische Briefe'', under the name Sciurus
sibiricus. Laxmann gives the following diagnosis:"... cinereo-
rufescens, striis quinque nigris longitudinalibus, palmis tetradactylis,
plantis pentadactylis" (p.69). It is called ''Burunduk"' and the first specimen
was caught in the vicinity of Barnaul. The valid name of the chipmunk
is therefore Eutamias sibiricus, according to the law of priority.
S.F. Baird (loc. cit., 1859) gives the following short descniption:
''Finding the Siberian animal thus without a name, I have called it Tamias
pallasii''. Ina footnote he says: ''In the Siberian species the color of
the upper parts is a dirty ocher yellow: in the American, brownish rust,
with much gray intermixed on the back..."
The name Tamias pallasii Baird together with the general terra
typica, Siberia, has to be considered as a synonym of Eutamias
sibirnicus sibiricus, Laxm.
This form differs clearly in series from the lighter, more yellowish
West Siberian and Ural-European forms. One may find specimens in the
range of the western form which practically do not differ from central
Siberian specimens. There are also intermediate forms, e.g., in the
vicinity of Tobolsk and in Bashkiria. However, the differences between
the western and central Siberian specimens are clearly shown in series.
GEOGRAPHICAL DISTRIBUTION. Central Bashkiria, the south of the
Tobolsk Territory, throughout the Irtysh area, the Semipalatinsk ribbon
pine forests, the Tomsk and Yeniseisk territories, the south of the
Turukhansk Territory, central and southern Yakutia and Transbaikalia to
the upper reaches ofthe Amur, andtothe Great Khingan range and the Kirin
Province of Manchuria (details on Map IX).
378. Eutamias sibiricus striatus Pallas (1778). West Siberian
or European chipmunk
1778. Sciurus striatus Pallas P,S, (nec Sciurus striatus Linnaeus, 1758). Novae species
quadrupedum e Glirium Ordine, pp. 378-384; Auct. cit., Zoographia Rosso-Asiatica, 1811, pp. 187-189,
1788. Sciurus striatus asiaticus GmelinJ.F,, in Caroli Linné, Systema Naturae, Vol.1, pp. 150-
151 ("Habitat: in omni Asia boreali as fluvio europaeus Dwina et Kama usque"’).
1856. Tamias striatus Brandt 1.F., Pozvonochnye zhivotnye Severno-evropeiskoi Rossii i v osobennosti
Severnogo Urala (Vertebrates of North European Russia, especially of the Northern Urals), in Hoffman,
‘Northern Urals" (Severnyi Ural), Sankt Peterburg, pp. 30-31. 3
1873. Tamias striatus Bogdanov M.N., Okhotnich'i i promyslovye zveri i ptitsy Evropeiskoi Rossii (Game
and Commercial Mammals and Birds of European Russia), 'Okhota i konnozavodstvo", No.4, p. 96.
1877, Tamias asiaticus var. borealis, CouesandAllen, Monographs of North American Rodentia,
Report of the U.S. Geological Survey of the Territory, Vol. XI, Washington, pp. 793-794 (Habitat:
"North Europe, North Asia and North America"),
TYPE LOCALITY AND DEPOSITION. In "Novae Species Glirium", 1778, p. 378, Pallas writes: "per
omnem Kamae fluvii tractum et ad Dwinam fere explicavit, quemadmodum per omnes Uralensium montium
frigidiores sylvas copiose passim occurrit, "'
Forests near the Kama river may be considered as the type locality of this race. Type lost.
MATERIAL EXAMINED: 239 skins and skulls.
DIAGNOSIS. Color light yellowish, without well marked reddish rust
tones. Anterior part of back buff-yellow-gray. (cinnamon-buff) with a
well marked gray tone. Black stripes narrow. Buttocks with dull rust-
red-yellow tinge (pinkish cinnamon). Top of head light rust-yellow with
marked admixture of chestnut-black hair.
Length of skull 38.2-40.2mm (M.39.2mm); condylobasal length 34.3-
37.3mm (M.35.8mm); interorbital space 9.2-11.mm (M.10.4mm);
422
486
zygomatic width 21-22.1mm (M.21.6mm); length of upper tooth row 6-
6.9mm (M.6.3mm); length of body and head 130.8-160mm (M. 150.1 mm);
tail 105-118mm (M.110.7mm); hind feet (without claws) 30.5-38 mm
(M.35.2mm).
SYSTEMATIC NOTES. Large series of this form differ clearly from
the central Siberian form in the more yellowish color and narrower stripes
on the back. Buttocks with less marked rust tones thaninE. sibiricus
Sibiricqus, Laxm.
Specimens from the vicinity of Tobolsk are intermediate between E.
Sibrrrews! "Ss rbrri cus Waxm. and Ss -strratu’s Pall”ain their
color. The same applies to specimens from the middle reaches of the
Belaya river in Bashkiria.
Specimen No. M. 4437 of my collection was caught by V. Ya.
Parovshchikov near the Tundra station of the Northern Railway in the
Khalmogory District (former Arkhangelsk Province). It has a saturated
color and a well marked rust tone, like in E.s.lineatus Siebold from
Sakhalin.
The name ''striatus'' proposed by Pallas is not used in modern
literature.
The name Sciurus striatus Linnaeus (Systema Naturae, Ed. 10,
1758, Vol.I, p.64) applies to an American species of a different genus,
Tamias striatus L.
The description of S. striatus in Pallas's ''Novae Species
Quadrupedum"' applies to Russian chipmunks. Pallas states (p. 381) that:
"Amerikanos Sciuros striatos ipse nunquam vidi..."
According to the law of priority this subspecies should be called E. s.
striatus Pallas.
GEOGRAPHICAL DISTRIBUTION. The boundary of its range runs
from Vologda and Kholmogory in the west to the southeast, reaching the
lower reaches of the Kama, but not the Volga. The boundary crosses the
northern districts of the former Kostroma and Nizhni Novgorod provinces.
It does not pass to the right bank of the Kama but runs from this river to
the Urals (see map) and continues towards Tobolsk. It includes the basins
of the Ob and Yenisei and the lower and middle reaches of the Stony
Tunguska. The boundary turns north alongthe Yenisei, including the region of
the Kureika river in the north to 68°N (see above).
379. Eutamias sibiricus jacutensis Ognev (1936). Yakut
chipmunk
1886. Tamias striatus Maak R. , Vilyuiskii okrug (The: Vilyuisk District), Vol. II, Sankt Peterburg,
p. 143.
1887. Tamias striatus Bunge A., Berichte tiber die Expedition nach den Neusibirischen Ins. , S. 38.
1903, Eutamias asiaticus Allen J, A., Report on the Mammals Collected in Northeastern Siberia,
Bulletin of the American Museum of Natural History, Vol. XIX, pp. 137-139.
1926. Eutamias asiaticus OgnevS.I., Mlekopitayushchie severovostochnoi Sibiri (Mammals of
Northeastern Siberia), pp. 86-89.
1928. Eutamias asiaticus Dukelski N.M., Materialien tiber die Sdugetier-Fauna des Jakutien
Gebietes, Zoolog. Anzeiger, Bd. 78, H.5-8, S. 107.
1934, Eutamias asiaticus asiaticus Tugarinov A. Ya., Smirnov N.A. and A.N. Ivanov, Ptitsy i
mlekopitayushchie Yakutii (Birds and Mammals of Yakutia), pp. 28, 39, 47, 56.
1936, Eutamias sibiricus jacutensis Ognev S.1., Sistematicheskii obzor burundukov (A Systematic
Survey of Chipmunks), Uchenye zapiski Moskovskogo universiteta, Issue IV, 1935 (1936), p. 93.
TYPE LOCALITY AND DEPOSITION. Vicinity of Yakutsk, Type No. M. 2290 male, 21 September
1927 (collected by K.E. Vorob'eva), in S.I. Ognev's collection.
MATERIAL EXAMINED: 55 skins and skulls,
423
488
DIAGNOSIS. Lighter whitish yellow color of anterior part of back.
Reddish rust tones of buttocks pale, and with admixture of white. Stripes
on back narrow. Top of head pale rust-yellow with some chestnut-black
hair.
Length of skull 38.8-41mm (M.39.5mm); condylobasal length 35-
36.5mm (M.35.6mm); interorbital space 10.3-11.6mm (M.11mm);
zygomatic width 21.2-22.6mm (M.22.mm); length of upper tooth row
6.1-7mm (M.6.5mm).
SYSTEMATIC NOTES. One of the forms with the lightest color and the
narrowest stripes . Its color closely resembles that of the West Siberian
subspecies (e.g., the Surgut and Ural chipmunks), but it lacks the rich
yellow color of this subspecies. In series the Yakut chipmunk clearly
appears more whitish.
GEOGRAPHICAL DISTRIBUTION. The range includes a wide area from
Khatanga, Olenek and Vilyui to the Anadyr in the west and the Penzhina in
the east. In East Siberia the southern boundary passes near 60°N, to about
120°E. Farther west the boundary turns north (see Map IX).
380. Eutamias sibiricus orientalis Bonhote* (1898). Ussuri
chipmunk
1858. Tamias striatus Schrenck L.v., Reisen und Forachungen in Amur-Lande, I,S, 124-125 (partim/).
1870, Tamias striatus Przheval'skii N. M., Puteshestvie v Ussuriiskom krae (A Journey to the Ussuri
Territory), Sankt Peterburg, pp. 256-257,
1899, Tamias orientalis Bonhote J.L., On a NewSpeciesofTamias. Annals and Magazine of
Natural History, 4(7):385-386.
1912, Eutamias asiaticus orientalis OgnevS.I., O kollektsii mlekopitayushchikh iz Ussuriiskogo
kraya (A Collection of Mammals from the Ussuri Territory), Ezhegodnik Zoologicheskogo muzeya
Akademii Nauk, XVI, pp. 497-500.
1923, Eutamias asiaticus orientalis Sowerby A., The Naturalist in Manchuria, II-III, pp. 148-149,
TYPE LOCALITY AND DEPOSITION. The Sungacha river, upper reaches of the Ussuri, Holotype in the
British Museum,
MATERIAL EXAMINED: 20 skins and skulls,
DIAGNOSIS. Color rich, with well marked saturated red-rust shades
(orange-cinnamon), especially on the buttocks. Top of head with well
marked reddish rust tones. Infraorbital stripe and middle of cheeks of
the same rich color.
Skull larger than in other forms. Length of skull 39.2-41.8mm
(M.40.2mm); condylobasal length 36.1-38mm (M.37.1mm); interorbital
space 10.2-11.3mm (M.10.8mm); zygomatic width 22-23.8 mm (M. 22.8 mm),
length of upper tooth row 6.1-7.1mm (M.6.6mm); length of body and head
148mm; tail 114.2mm; hind foot 36mm.
SYSTEMATIC NOTES. This form differs clearly in series from others
in its rich, saturated rust-red color of the buttocks. Skull larger than in
the other subspecies.
* In Chaworth-Musters's opinion (A Note on the Nomenclature of Two Sciurids from Eastern Siberia. Ann.
Mag. Nat. History, 13(10):555-557, 1934), orientalis Pall, is synonymous with uthensis Pall.
(see below),
This is not correct. I have shown that the name Sciurus uthensis applies to a melanistic
specimen from the village Udskoi, i.e. , from the range of the central Amur race Eutamias sibiricus
lineatus. As this is a melanistic specimen it is correct to use the name proposed by Siebold (see below).
424
GEOGRAPHICAL DISTRIBUTION. Southern part of the Maritime
Territory, the Ussuri Territory, Manchuria. Specimens from the eastern
side of the Great Khingan and Kirin Province of Manchuria are more
closely related to the nominate form than toorientalis. The range of
E. sibiricus sibiricus inthe Kirin Region is wedged into the range
oforientalis.
381. Eutamias sibiricus lineatus Siebold (1826). Amur
chipmunk
1811, Sciurus uthensis Pallas P,, Zoographia Rosso-Asiatica, pp. 189-190 (description of a melanistic
specimen),
1826. Myoxus lineatus Siebold Ph, F,, De historia naturalis in Japonia Statu, etc. , Wirgerburgi,
pp. 16, 17.
1851, Tamias uthensis Simashko Yu,, Russkaya fauna. II, (Russian Fauna. II), p. 603.
1858. Tamias uthensis SchrencklL., v., Reisen und Forschungen im Amur-Lande, I, p. 123.
1867. Tamias (Sciurus) uthensis Middendorf A, Th., Sibirische Reise, S. 83.
1889, Tamias asiaticus borealis Nikol'skii A. M., Ostrov Sakhalin i ego fauna pozvonochnykh
zhivotnykh (The Vertebrate Fauna of Sakhalin Island), pp. 125-126.
1890. Tamias pallasii Suprunenko, Pozvonochnye Sakhalina, (The Vertebrates of Sakhalin), p. 39.
1922, Tamias uthensis ochotensis Dybowski Ben., Arch., Tow. Naukow. Lwowie, III, p. 8.
1936, Eutamias sibiricus lineatus OgnevS.I., Sistematicheskii obzor burundukov (A Systematic
Survey of Chipmunks), Uchenye Zapiski Moskovskogo universiteta, Issue IV, 1935 (1936), p. 94.
TYPE LOCALITY AND DEPOSITION. Yezo Island (Hokkaido). Holotype not known.
MATERIAL EXAMINED: 37 skins (most of them with skulls),
DIAGNOSIS. Closely relatedtoE.s.orientalis but differing from
it in its paler color, especially on the buttocks where the rust-red tones are
not as saturated as inorientalis. Top of head also lighter and yellower
than inorientalis.
This form is intermediate between orientalis andjacutensis.
Length of skull 38.2-39.3mm (M.38.9mm); condylobasal length 35.2-
37.2mm (M.36.1mm); interorbital space 10.3-11.1mm (M. 10.7 mm);
zygomatic width 20.3-22.4mm (M.21.5mm); length of upper tooth row
6.2-6.9mm (M.6.5mm ); length of body and head 139-147 mm; length of
tail 112-124mm; hind feet 33-37 mm.
489 SYSTEMATIC NOTES. Pallas (''Zoographia Rosso-Asiatica", p.189-
190, 1811) describes 'Sciurus uthensis" from a melanistic specimen
caught on the Uda river as follows: ''tota pellis nitidissime atra, paulo
minor Sc. striato; auriculae paulo minores et cauda proportione brevior,
quam in eodem. Pedum structura, digitis pollice subunguiculato
verruciformi palmarum, et parte nuda simillimi Strige dorsi item similes.
Sub collo tractus longitudinalis albus, a labio inferiore ad sternum
continuus; caeterum pars prona tota nigra. Cauda tereti — pilosa, sed
minus late villosa, pilis magis uniformibus, strictius laevigatis ubique
Bterrimis. ...
As Pallas described a melanistic specimen, it would be correct to
apply the subspecific name "lineatus'' to the Sakhalin-Amur form.
Ph. F. Siebold 1826, loc. cit.) gave the following description of Myoxus
lineatus: "M.lineatus tergo lineis quinque longitudinalis
nigrescentibus notato, habitat in insula Yezo. Longitudo corporis 6 12 poll. ,
caudae 41/2 poll.".
According to Nagamichi Kuroda, who published a survey of chipmunks in
1932, the islands of Yezo [Hokkaido] and Sakhalin are inhabited by the same
subspecies, namely lineatus. AsI have not seen any specimens from the
425
Yezo Island, I have to accept the opinion of Kuroda. His conclusion is
reasonable as the fauna of Yezo and Sakhalin are closely related.
GEOGRAPHICAL DISTRIBUTION. E.s.lineatus occurs on the
islands of Yezoand Sakhalin, on the Shantar Islands, in the middle and lower
reaches of the Amur and along its tributaries Khor, Goryun, Khungari,
ere.
426
490
SUPPLEMENT TO VOLUME THREE
Felis (Eremaelurus) thinobius Ognev
The sand or barchan cat.
Several publications on the systematic position of the barchan cat, a
peculiar representative of the Russian fauna, have appeared recently.
W. Heptner and G. Dementiev published an article ''Sur les relations
mutuelles et la position systématique des chats desertiques
Eremaelurus thinobius Ognev et Felis margarita Loche'
("Mammalia", Vol.I, 1937, Paris, pp. 217-242).
V.G. Heptner and G.P. Dementiev became interested in this question
through an article of H. Heim de Balzac (''Biogéographie des Mammiferes
et des Oiseaux de l'Afrique du Nord", supplément XXI au Bulletin Biologique
de France et de Belgique, Paris 1936, pp.87, 171-172, 239, 258, 393).
H. Heim de Balzac suggests that Eremaelurus thinobius is
closely related to the Sahara sand cat which was described by Loche (1858)
but its systematic position was not clear because of lack of material.
Heptner and Dementiev obtained a specimen of Felis margarita
male ad., caught 29 May 1930 in Algeria (Lagouat) from the Musée
_ National d'Histoire Naturelle de Paris.
491
A comparison of the skin and skull of this specimen with Eremaelurus
thinobius showed that these two species are closely related. In their
opinion, thinobius is merely a subspecies of margarita. After
study of the skull, although they had only limited material, they came to
the conclusion that the genus Eremaelurus should be considered as a
synonym of the genus Otocolobus (to which Felis manul belongs).
The paper of Heptner and Dementiev attempts to solve two problems:
1) whether the genus Eremaelurus should beconsidered as a synonym
of Otocolobus; 2) what is the relationship between margarita and
thinobius.
The authors give a positive answer to the first question because the skull
of Felis manul and Eremaelurus resemble each other in a number
of characters: the ratio of the length of the zygomatic arches to the greatest
length of the skull, the ratio of the zygomatic width to the condylobasal
length, and the ratio of the length of the bullae osseae to the condylobasal
length.
The two last ratios are of little value. The skull of Otocolobus is
much shorter than that of Eremaelurus, and the condylobasal length
is small. Although the bullae osseae of Otocolobus are much smaller
than those of Eremaelurus, the ratio of the length of the bullae to the
small condylobasal length give a similar ratio. The authors also indicate
that the chambers of the bullae osseae of Eremaelurus and Otocolobus
are of similar structure. However, in addition to these similarities there
are a number of differences. The authors state that the two genera differ
427
492
in twelve important characters (pp. 234-235). If differences not mentioned
by Heptner and Demetriev are added to these twelve, it appears that
Otocolobus andEremaelurus differ sharply.
We shall mention here only the most important differences; 1) the
orbits of Otocolobus are smaller than those of Eremaelurus, more
rounded and directed more forwards; 2) the bullae osseae are much
smaller; 3) the limit between the anterior and posterior chambers of the
bullae osseae is much sharper in Otocolobus thaninEremaelurus,
and the anterior chamber is generally smaller; 4) talon of pm* much
smaller; 5) rostrum much shorter; 6) jaws and tooth rows shortened;
7) coronoid process shorter and wider, or a quite different form from that
of Eremaelurus. The form of the coronoid process in Felis manul
is very peculiar, and differs from that of Felis silvestris and
Felis ornata.
Felis manul andEremaelurus do not differ only in characters
of the skull. Felis manul differs sharply from Eremaelurus inits
short ears, its luxuriant fur and the unique pattern of the fur of the tail. The range
of Felis manul is restricted mainly to the highlands of Central Asia.
Eremaelurus has large ears, its feet are covered with luxuriant black
fur and the fur on the back is pale and not smooth. It is a typical
inhabitant of the desert. The shagginess of the soles resembles that of
hares or other rodents of sand deserts (e.g., jerboas) and are apparently
an adaptation to life in the barchans.
For the sake of ''simplification'' generic or subgeneric classifications
may be ignored, and subgenera may be placed into a generally accepted
genus, as it has been done by R. Pocock.
However, to place two species which differ morphologically and
ecologically like Eremaelurus and Otocolobus, into the same genus
as Felis manul is shortsighted and difficult to understand ina
mammalogist or ornithologist. It would cause only confusion. R. Pocock
(1938) made some critical remarks on the work of Heptner and Dementiev,
but he ignored their suggestion to place thinobius and margarita in
the genus Otocolobus. This question seemed apparently of little interest
to Pocock.
This discussion attempts to clarify the problem for those readers who
may be misled by the article of Heptner and Dementiev.
It seems impossible, in our opinion, to unite Eremaelurus with
Otocolobus. AsEremaelurus shows some characters intermediate
between Felis and Otocolobus, and as Otocolobus is better
differentiated than Eremaelurus, we consider it correct to regard
Eremaelurus as a subgenus of Felis.
Heptner and Dementiev also gave a positive answer to the second
question and named the Transcaspian sand cat 'Otocolobus margarita
thinebra.
Let us now examine whether this problem is so simple.
R. Pocock recently described two subspecies of Felis margarita™*.
The first subspecies was collected near Niogus [?] on the border between
Tripoli and Algeria, and was named Felis margarita
* R. Pocock, The Algerian Sana Cat (Felis margarita Loche), Proceedings of the Zoological Society of
London, Series B, Vol. 108, pp. 41-46, 1938; A New Race of the Sand Cat, Annals and Magazine of
Natural History, Series 11, Vol.I, pp. 472-476.
428
meinertzhageni Pocock. The other subspecies was caught in In-
Abbangarit, west of Air (southwestern Sudan) and named Felis
margarita airensis Pocock.
R. Pocock describes the differences between F. thinobius and
F. margarita and comes to the conclusion that thinobius isa
separate species. In his opinion, margarita andthinobius
developed from F.ocreata. He also considers Felis ornata and
Felis caudata as subspecies of ocreata. In Pocock's opinion,
thinobius andmargarita evolved from ocreata in geographically
isolated regions, as their range was interrupted. He therefore interprets
the common characters of these species as convergences, which evolved
in adaptation to desert life.
Without discussing the arguments of Pocock, the problem certainly
seems more complicated than Heptner and Dementiev thought.
I cannot solve this problem for lack of material (we have only one skin
and skull of F. margarita and several specimens of F. thinobius).
However, if the careful descriptions of R. Popcock (who is a renowned
expert on carnivora) are compared with our specimens, some differences
between F. margarita and F. thinobius can be pointed out.
Adult males of the Transcaspian sand cat differ from F. margarita
as follows: 1) much larger size; 2) larger bullae osseae; 3) the
occipital region is lower in relation to the length of the skull (this difference
was also noted by R. Pocock); 4) relatively shorter nasals; 5) large
orbits; 6) rostrum of thinobius generally longer than that of
margarita in spite of the relatively short nasals; the long nasals of
F, margarita project further backwards; 7) the color of F.
margarita is paler than that of F.thinobius.
493 All our data prove that Felis (Eremaelurus)thinobius Ogn.
should be considered as a good species which belongs to the same
subgenus as F. (E.) margarita Loche.
The ranges of these species are adjacent. F. margarita occurs
in the western Sahara desert and reaches Algeria in the north. Heptner
and Dementiev confidently populate Egypt with Eremaelurus (loc. cit.,
p. 240), referring to an article by S. Flower (''The Recent Mammals of
Egypt", Proceedings of the Zoological Society of London, 1932, p. 249).
However, Flower only assumes that F. margarita occurs inthe
western Egyptian desert. Eremaelurus has not been found so far in
the wide area including eastern Tripoli, Egypt, Arabia, Syria, Palestine
and Iran, although this was explored by many zoologists. It is not probable
that it will be found in this region.
To conclude, it is very interesting that an African and a Turkmenian
species of the same highly specialized subgenus are so similar and closely
related.
429
| 28
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