0 'j^A.
0
THE FAMILIES AND GENERA OF
LIVING RODENTS
BRITISH MUSEUM
(NATURAL HISTORY)
THE FAMILIES AND GENERA
OF
LIVING RODENTS
^(^^'^ BY
a.OJ^'^ ^^^ |. R. ELLERMAN '
^v' WITH A LIST OF N.\]\IED FORMS (1758 -1936)
BY
R. W. HAYMAN and G. W. C. HOLT
VOLUME L RODENTS OTHER THAN MURIDAE
LONDON:
PRINTED BY ORDER OF THE TRUSTEES
OF THE BRITISH MUSEUM
Issued 8th yune, 1940] [Prke One Pound Fifteen Shillings
Sold at The British Museum (Nait'r.\l History), Cromw-ell Road. S.W.y,
and by
B. QuARiTCH, Ltd.; Dulau & Co., Ltd.;
and the Oxford University Press
made and printed in great BRITAIN
BY JARROLD AND SONS LTD. NORWICH
THIS WORK IS
DEDICATED
TO THE MEMORY OF
MY FRIEND
WILLIAM COX
PREFACE
As the result of several years devoted to a careful study of the rich recent
collections belonging to this museum, the author has prepared the following
review of the families and genera of living Rodents; he has also supervised the
preparation by Mr. R. W. Ilayman and Mr. G. W. C. Holt, members of the
Museum staff, of a list of all the species and subspecies of rodents, described
from the loth edition of Linnaeus's Systema Natura, I'j^d, down to the end of
1936. Such a review and such a list have long been two of the most needed
desiderata of zoologists.
The author has endeavoured in each case to find out not only what characters
have been assigned to a given genus, but what characters it in fact possesses,
and to test their value and constancy. Genera are recognized by their intrinsic
characters ; a mere geographical differentiation of genera does not exist. When-
ever the author has been able to study a genus for himself he has included it
in his " Keys" ; when not seen personally he merely gives a note of the ascribed
characters. The carrying through of this great examination has led to a con-
siderable reduction in the number of genera here recognized ; thus, of 440 forms
or groups given, at some time or another, full generic rank, only 343 (151
non-murine, 192 murine) are now regarded as valid genera. Indeed, had
the Museum collections of North American Rodents been more extensive, and
had it been possible to make a really detailed survey of the South American
Muridae, a further much-needed reduction of genera would doubtless have
taken place.
With regard to major classification, the author excludes the Duplicidentata
from the Order Rodentia. He reviews the more recent classifications of the
Order (so restricted), to wit, those of Oldfield Thomas, 1896; TuUberg, 1899;
Weber, 1904 and 1928; Miller & Gidley, 1918; and Winge, 1934. He then
proposes a new classification of the families and genera, which, while necessarily
sharing some features with one or other of the older systems, on the whole
appears to be a great improvement upon any of the systems previously proposed.
Of great interest is the chapter on distribution and the conclusion reached
by the author that, as regards its peculiar Muridae, Australasia may be claimed
as an evolutionary' centre. That view, in face of the characters of the group in
question and the very high antiquity of the Order, appears to me to be perfectly
sound.
To conclude these general remarks the author is to be congratulated on
having performed a colossal task, and we are indebted to him for providing an
honest one-man view which cannot fail to be of service to all who wish to study
this great and complex Order in future.
With regard to the list of named forms, every endeavour has been made to
make it complete and accurate, but it is too much to hope that nothing has
escaped the compilers.
viii PREFACE
The beautiful drawings of skulls and teeth prepared by Mr. A. J. Engel
Terzi greatlv enhance the value of this work; with the exception of Figs. 62, 63,
90. 96, 112, 115, iiS, 119, 146, 164, 167, 170, 173 and 184, bv the same artist,
hut originally published in Miller's Catalogue of the Mammah of Western Europe,
IQ12, all the figures have been specially drawn tor this volume.
Owing to the heavy work undertaken by the artist and to the prevalent war
conditions, considerable delay has occurred in the publication of Vol. I, which
deals with the general matters discussed above and with the families and genera
of non-murine rodents.
Vol. II, dealing with the murine rodents, is already in type; in order to
avoid further delay it is proposed to issue it immediately without waiting for
the preparation of its full complement of figures.
\'ol. Ill, to be published later, will be an "Atlas" containing all the figures
of Vol. I repeated; the full complement of figures for Vol. II; to these it is
hoped to add drawings of some of the more interesting and important external
characters and of dissections ot jaw muscles.
MARTIN A. C. HINTON,
Keeper of the Department of Zoology.
British Museum (Natural History)
25//; March, 1940
AUTHOR'S FOREWORD
In 1896 Oldfield Thomas proposed a classification of Simplicidentate
Rodents in which he recognized 156 genera.
In 1904, Trouessart in his Catalogue of Mammals listed 205 genera of this
section of mammals.
At the present day, more than four hundred and forty forms have been
given, at some time or another, full generic rank, so that in the last thirt\--five
years there has been an increase of approximatelv two hundred and forty genera.
The object of this work is primarily to inquire into the status of these
named genera, and to give, in each family and subfamily, a key which will
indicate as reasonably, clearly and briefly as possible, the differences between
such genera as are supported on characters which appear constant through the
various groups, and worthy of generic recognition.
This has led on the one hand to a careful studv of the classification of the
families and superfamilies or major groups of all authors who have attempted
an arrangement of the Order based on adequate material and including all the
principal leading genera of all families as recognized to-day (Thomas, 1896;
Tullberg, 1899; Weber, 1904, 1928; Miller & Gidley, 1918; and Winge,
1 924), and on the other hand to the collection of a list of all named forms (species
and subspecies) which have been named in the order since Linnaeus (i7:;8), up
to and including the year 1936.
It is not my intention to enter into a detailed description of the skeleton,
soft parts, etc., of each genus; this has been done alreadv in a far more com-
petent manner than I could attempt (Tullberg, 1899). Ninety-nine out of a
hundred genera are based on cranial, dental, and external characters, or to put
it more crudely, "skull and skin" characters. This work is based almost
entirely on these characters. It must be noticed that in cranial, dental and
external characters, ver\' many specimens of a species or genus may usually be
at hand for examination, so that whether such a character is constant or not
can generally be checked easily ; generic names based on skeleton, soft parts,
or characters such as the baculum, which has been used for generic names
(here not retained) in the Sciuridae, can as a rule only be examined for one
species and often one specimen of a genus ; therefore it is not possible to give
full notes on such a character throughout a whole genus or, if it is so, the notes
refer to a restricted number of specimens only. Under these circumstances, it
seems wiser not to pay too much attention to names which have been based
solely on such characters.
This work is based entirely on the collection of the British Museum, to the
authorities of which I am much in debt for their kindness and consideration
throughout the compilation of this work. No genus which is not represented
in that collection is included in my keys; as it is difficult to include in a key
any form which has not been examined, and in the case of certain Muridae,
X AUTHOR'S FOREWORD
impossible. However, there are only five non-Alurine genera at present un-
represented in London, and about sixteen (out of nearly two hundred) Muridae;
notes on these will be included, but generally speaking no comments. Of the
genera which are included, I have endeavoured to give in each case the approxi-
mate range, the number of forms at present recognized (to 193(1), a list of these
forms, and a short description of the main cranial, dental and external characters.
Concerning the dental characters, in this Order, great difference of opinion
exists regarding the homologies of the various parts of each tooth. American
authors use in the main a series of names for each cusp which are figured and
explained by Goldman, North Anicr. Fauna, 43, p. 11, 191S; Miller & Gidley,
iqiS, divide the greater part of the Order into "tritubercular" and "quadri-
tubercular" series; Winge apparently uses quite a different theory which he
takes throughout Mammalia; and Hinton uses still a different notation. With
the exception of the Murinae, in which a series of eight or sometimes nine
main cusps go through the entire subfamily, no attempt has been made in this
work to use a definite formula; I have endeavoured to describe the dentition of
each genus as I see it, and am content to leave the working out of cusp homolo-
gies to those with more experience than I have.
The view that the dental pattern of modern Rodentia is generally speaking
derived from a much more complex pattern than is now present, expressed by
Hinton, Monograph of Voles and Lemmings, 1926, Evolution of Molars, pp.
102-124, is here accepted. However, it is not the purpose of this paper to enter
into an argument as to whether this view is correct, or the view frequently held
that a complex dentition in a living Rodent is a secondarily acquired one. Take
for instance two cases, Hapalomys, a very complex-toothed Rat (Murinae) as
compared with Rattiis (normally a simple-toothed Rat); in this work, Hapalomys
is considered the primitive type, Rattiis the specialized one; but turn these
views round, and the two genera will still be at the opposite end of two extremes,
which is broadly speaking what I set out to prove in each case.
So far as the list of named forms of each genus is concerned, I list those
which are named, making no attempt to guarantee the validity of any subspecies
or species. How many of these names will ultimately be reduced to synonymy
is not clear; but I believe that very many of them will prove invalid with more
material available. I have attempted in each case in which a genus has not been
revised, where possible, to divide the genera into "specific groups" as is now
done by American authors. These groups indicate certain characters within
the different species of each genus, but must be regarded as provisional. The
attitude, however, is held that a list of names in some semblance of order, no
matter how provisional, is better than a string of meaningless alphabetical
names. At least I hope it gives a start to those who are interested in the
characters w^hich run through the species of the various genera. I expect how-
ever that a large number of South .\merican Mice (Cricetinae) will have for the
time being to be abandoned, and listed alphabetically. It is perhaps not too
much to hope that these lists will act as a deterrent to authors who rush to give
names to new forms before consulting all the literature on the genus in question.
Though great care has been taken I can give no guarantee that the list of
AUTHOR'S FOREWORD xi
named forms is complete, particularly in the case of some of the older synonyms,
to which less attention has been paid than to names described more recently.
The list was originally based on that of Trouessart (Cat. Mamm. Viv. et Foss,
1904); names which appear in synonymy in this work have in most cases been
included here as they appear in Trouessart's list, and their position has not been
verified. I have listed si.xty-four hundred forms which are supposed to be valid at
the present day. Subspecies, except in cases of a genus which is definitely revised,
are listed as far as possible geographically. Each named form is listed under its
present accepted name, or the name which appears to be correct; in many cases
not under the generic name under which it was described, for instance " Sminthus
longer," Nathusius, 1840, is now listed as " Sicista subtilis loriger," etc., etc.
The Order is absolutely dominated by one family, the Muridae, both in
number of genera and named forms, as proved by the following figures, which
must be taken as approximate.
Twenty-two families of Rodents other than Muridae:
151 valid genera containing 2,773 named forms.
Family Muridae :
192 valid genera containing 3,600 named forms.
I have had therefore to divide the work into two volumes, the first of which
contains all Rodents not belonging to the family Muridae, the second devoted
entirely to that family.
This work is based solely on Rodents which are living, or assumed to be
living, though I have added short notes on the fossil history of each family,
chiefly from a distributional point of view.
My sincere thanks are due to the officials of the British Museum for their
kind help and consideration throughout the time I have been preparing this
work; especially I must mention Captain Guy Dollman, who originally made it
possible for me to undertake this review; Mr. M. A. C. Hinton, who has under-
taken the task of editing the work; Mr. Hayman and Mr. Holt, who have
between them got together the references and type localities of more than
sixty-four hundred names in the Order, and the former for much assistance in
dealing with some of the species of the more unwieldy genera; Dr. Tchernavin,
who has translated several papers from Russian, enabling me to give some
details concerning the distribution of the various groups of Rodents occurring
in the U.S.S.R.; Mr. A. J. Engel Terzi, who has made drawings of the more
important genera; Mr. J. L. Chaworth-Musters, to whom I am indebted for
nearly all my knowledge of Palaearctic Rodents; also I must thank Mr. E. R.
Newman, who has given me much help throughout the compilation of the work,
and Miss R. Blizard and Mr. F. C. Hitch, who have assisted during the later
stages. Lastly I would thank my wife for numerous working drawings of
specimens and continuous help in other ways. The indulgence of readers is
sought for any typographical or other small errors in this work. The writing
of the book itself was finished in June last, but the revision of the final prools
had not been completed when war broke out. Since then it has been possible
for me to give onlv the most cursorv and intermittent attention to such revision.
b
CONTENTS
Preface
Author's Foreword
Limits of the Order Rodextia
Variation in Rodents .
Previous Classifications of the Order
Oldfield Thomas, 1896
Tullberg, 1899
Weber, 1904, 1928
Miller & Gidley, 1918
Winge, 1924 ....
Outline of Cl.\ssification here adopted
Zygomasseteric Structure
Distribution .....
Lists of Genera and Principal Species dealt
Palaearctic ....
Nearctic . . . . .
Indo-Malayan ....
African .....
Neotropical ....
Order RODENTIA
Key to Superfamilies
Superfamily BATHYERGOIDAE
Family Bathyergidae
Genus Bathyergus, Illiger
Genus Heliophubius, Peters
Genus Georychus, Illiger
Genus Cryptomys, Gray
Genus Heterocephalus, Rijppell
Superfamily IIYSTRICOID.AE .
Family Echimvidae .
Subfamily Echimyin.\e .
Genus Echimys, Cuvier
Genus Isothrix, Wagner
Genus Diplomys, Thomas
IN this V'
olume
PAGE
vii
i.\
I
3
5
5
8
II
13
18
22
42
47
57
57
60
64
66
69
77
77
79
79
83
84
86
86
94
96
lOI
106
108
"3
115
CONTENTS
Genus Piorchinivs. Allen
Genus Hoploinvs, Allen
Genus Cercoinys, Cu\ier
Genus Euryzygomatornys, Goeldi
Genus Clyomvs, Thomas
Genus Carterodon, Waterhouse
Genus Mesomvs, Wagner
Genus Loncliothrix, Thomas
Subfamily Capromyinae
Genus Capromvs, Desmarest
Genus Geocapromys, Chapman
Genus Procapromxs, Chapman
Subfamily Plagiodontinae .
Genus Plagiodontia, Cuvier
Subfamily Dactylomyinae
Genus Thrinacodus, Giinther
Genus Dactvloinys, Geoffroy
Genus Kaimahateomvs, Jentink
Subfamily Myocastorinae
Genus Mvocasior, Kerr
Subfamily Thryonomyinae .
Genus Tluyonomys, Fitzinger
Subfamily Petromyinae
Genus Petromus, Smith
Subfamily Abrocominae
Genus Ahrocoma, Waterhouse
Subfamily Octodontinae
Genus Octoiiiys, Thomas
Genus Aconaemys, Ameghino
Genus Octodon, Bennett
Genus Octodoiitoinys, Palmer
Genus Spalacopiis, Wagler
Genus Ctenomys, Blainville
Family Dinomyidae .
Genus Dinomys, Peters
Family Frethizontidae
Subfamily Chaetomyinae
Genus Cliaetomys, Gray
PAGE
"5
122
123
124
125
126
127
128
128
132
133
133
134
13s
136
137
139
140
144
144
149
149
152
156'
160
161
170
171
173
174
175
CONTENTS
Subfamily Erethizontinae
Genus Erethizon, Cuvier
Genus Echinoprocia, Gray
Genus Coendou, Lacepede
Family Dasyproctidae
Genus Dasyprocta, Illiger
Genus Myoprocta, Thomas
Family Hystricidae .
Genus Trichys, Giinther
Genus Atherurus, Cuvier
Genus Thecurus, Lyon
Genus Hystrix, Linnaeus
Family Cuniculidae
Genus Ctinicultis, Brisson
Family Chinchillidae
Genus Chinchilla, Bennett
Genus Lagidium, Meyen
Genus Lagostomus, Brooks
Superfamily CAVIOIDAE
Family Caviidae
Subfamily Caviinae
Genus Cavia, Pallas
Genus Galea, Meyen
Genus Caviella, Osgood
Genus Kerodon, Cuvier
Genus Dolichotis, Desmarest
Subfamily Hydrochoerinae .
Genus Hxdrochoerus, Brisson
Superfamily APLODONTOIDAE
Family Aplodontiidae
Genus Aplodontia, Richardson
Superfamily SCIUROIDAE
Family Sciuridae
Outline of previous classification of the family
Classification here adopted .
Genus Belomys, Thomas
Genus Trogopterus, Heude .
Genus Pteromyscus, Thomas
XV
PAGE
•77
178
181
182
189
190
196
197
203
205
209
212
220
221
226
227
229
233
237
237
238
240
242
243
246
247
249
250
253
253
255
259
259
2fil
269
276
279
280
CONTENTS
Genus Petaurista, Link
Genus Aeromys, Robinson & Kloss
Genus Pteromys, Cuvier
Genus Glaucomys, Thomas .
Genus Eoglaucomys, Howell
Genus Hylopetes, Thomas .
Genus Petinomxs, Thomas .
Genus Petaurillus, Thomas .
Genus lomvs, Thomas
Genus Eupetaunis, Thomas
Genus Mvosciuriis, Thomas
Genus Nannosciurus, Trouessart
Genus Sciurillus, Thomas
Genus Microsciuriis, Allen .
Genus Svntheosciuriis, Bangs
Genus Sciurus, Linnaeus
Genus Tamiasciunis, Trouessart
Genus CaUosciurus. Gray-
Genus Fuuamhulus, Lesson .
Genus Dremomxs, Heude
Genus Ratufa, Gray .
Genus Menetes, Thomas
Genus Lariscus, Thomas & Wroughton
Genus Glyphotes, Thomas .
Genus Wieithrosciurtis, Gray
Genus Rhinosciurus, Gray
Genus Hvosciuiiis, Tate & Archbold
Genus Heliosciurus, Trouessart
Genus Paraxerus, Forsyth Major
Genus Funisciurus, Trouessart
Genus Protoxenis, Forsyth Major
Genus Mvrsilus, Thomas
Genus Epixenis, Thomas
Genus Xenis, llemprich &: Khrenberg
Genus Atlantoxeriis, Forsyth Major
Genus SpeiinuphHopsis, Blasius
Genus Sciurotamias, Miller
Genus Tamicn, Illiger
PAGE
281
290
291
294
297
29S
300
302
3°3
304
312
313
317
319
321
321
345
348
376
3S0
383
390
391
393
393
395
39S
399
405
410
415
416
417
41S
422
423
425
426
CONTENTS
Genus Citellus, Oken .
Genus Marmota, Blumcnbach
Genus Cynomys, Rafinesque
Superfamily CASTOROIDAE
Family Castoridae
Genus Castor, Linnaeus
Superfamily GEOMYOIDAE
Family Heteromyidae
Subfamily Heteromyinae
Genus Heteromys, Desmarest
Genus Liotnys, Merriam
Subfamily Dipodomyinae
Genus Perognathus, Wied
Genus Microdipodops, Merriam
Genus Dipodomys, Gray
Family Geomyidae .
Genus Thomomys, Wied
Genus Geomys, Rafinesque .
Genus Pappogeomys, Merriam
Genus Cratogeomvs, Merriam
Genus Platvgeomvs, Merriam
Genus Ortliogeomys, Merriam
Genus Heterogeomys, Merriam
Genus Macrogeomvs, Merriam
Genus Zygogeomys, Merriam
Superfamily ANOMALUROID.\E
Family Anomaluridae
Subfamily AiNOMALURINAE
Genus Anomalurus, Waterhouse
Genus Anomalurops, Matschie
Subfamily Idilrinae
Genus Idiunis, Matschie
Genus Zenkerella, Matschie
Superfamily PEDETOIDAE
Family Pedetidae
Genus Pedetes, llliger
Superfamily CTENODACTYLOIDAE
Family Ctenodactylidae
437
454
461
464
464
465
468
470
471
472
476
479
480
492
494
505
507
524
527
528
530
531
532
533
534
535
535
537
537
541
542
543
546
547
547
549
553
553
CONTENTS
Genus Pectinalor, Blvth
Genus Ctetwdactyhis, Gray ■
Genus Massoutiera, Lataste
Genus Felm-ia, Lataste
Superfamily DIPODOIDAE
Family Dipodidae .
Subfamily Sicistinae
Genus Sicista, Gray .
Subfamily Zapodinaf .
Genus Eozapus, Preble
Genus Zapiis, Coues .
Genus Xapaeozapiis, Preble .
Subfamily Cardiocraminae .
Genus Salpingotm, Vinogradov
Genus Cardiocraiiiiis, Satunin
Subfamily EfCHOREUTiNAi-:
Genus Euclioietites, Sclater .
Subfamily Difodinae .
Genus Allactaga, Cuvier
Genus Alactagidus, Xehring
Genus Pygeretmiis, Gloger .
Genus Parmiipiis, Vinogradov
Genus Dipiis, Zimmermann .
Genus Scirtopoda, Brandt
Genus jfaciilus, Erxleben
Genus Ereiiiodipiis, \'inogradoy
Superfamily MUROIDAE ,
Family Muscardixidae
Subfamily Graphiurinae
Genus Graphiurus, Smuts
Subfamily Musc.ardinixae
Genus Eliomxs, \\'agner
Genus Dvromys, Thomas
Genus Gliruhis, Thomas
Genus Glis, Brisson
Genus Muscardinus, Kaup
Genus Mvoiniiniis, Ognev
Subfamily Platacaxthomyinae
CONTENTS
xix
PAGE
Genus Platacanthomys, Blyth ...... 627
Genus Typhlomys, Milne-Edwards
. 629
Family Lophiomyidae
. 632
Genus Lophiomys, Milne-Edwards
. 632
Family Spalacidae ....
• 636
Genus Spalax, Guldenstaedt
■ 638
Family Rhizomyidae
• 644
Genus Rhizomys, Gray
. 646
Genus Cannomys, Thomas .
. . 651
LIST OF TEXT-FIGURES
FIG
PAGE
I
Bathyergiis suillus suillus, Schreber .
. Skull X I
82
2
do.
do.
83
3
do.
Mandible x i
83
4
Cryptomys damarensis, Ogilby .
. Skull X 2
88
S
do.
do.
89
6
Heleroccphalus glaber, Riippell .
. Skull X3i
94
7
do.
do.
95
8
Echimys armatus armatus, Geoffrey .
. Skull X 4
110
9
do.
do.
no
10
do.
Cheekteeth X 7
III
1 1
Proechimys cayennensis, Desmarest .
. Skull xij
116
12
do.
do.
116
13
do.
Cheekteeth x 8
"7
H
Geocapromys brownii, Fischer .
. Skull x I
130
15
do.
do.
130
16
do.
Cheekteeth X 7
131
17
Kannabateomxs amblyonyx, Wagner .
. Skull xij
138
18
do.
do.
138
19
do.
Cheekteeth X 5
139
20
Myocastor coypiis santaecruzae, Hollister
. Skull X*
141
21
do.
do.
142
22
do.
Mandible X f
142
23
do. ,
Skull X i
143
24
do.
Cheekteeth x 2
'43
25
Thryonomys gregorianus, Thomas
. Skull X I
146
26
do.
do.
146
27-
do.
do.
147
28
do.
Cheekteeth X 4
147
29.
Petrumus typicus. Smith ....
. Skull X 2
150
30-
' do.
do.
ISO
31-
do.
Mandible X2
Cheekteeth X 7
151
32.
Abrocoma bennetti, Waterhouse
. Skull XI*
152
LIST OF TEXT-FIGURES
34-
35-
3b.
37-
38.
39-
40.
41-
4^-
43-
44-
45-
46.
47-
48.
49-
50.
51-
52-
53-
54-
55-
56,
57-
58.
59-
60.
61.
62.
^V
64.
^5-
66.
67.
68.
69.
Ahrocorna hennetti, Waterhuuse
do.
OctoniYS iiiiniax, Thomas
Spalacopus cyanus, Molina
Ctenomys tuconax, Thomas
do.
do.
Dinomvs hriviickii, Peters
do.
Chaeiomvs siibspinosiis, Kulil
do.
do.
Ercthizon ipixaiitliuin myops, Merriam
do.
do.
do.
Coendou prehensilis holiviensis. Gray .
do.
do.
do.
Dasyprocta punctata isthmica, Alston
do.
do.
Trichxs macrotis. Miller .
do.
At/wniiiis tiirni'ii, St. Legcr
do.
T/ieciiriis ciassispinis, Giinther .
do.
Hvstrix cristata, Linnaeus
do.
Cuniculus paca, Linnaeus
do.
Chinchjlla laniger, Molina
do.
Lagostomus maxinius, Desmarest
do.
PAGE
Skull ■ lA
153
Cheekteeth X7
153
Cheekteeth x 8
157
Skull xzA
160
Skull xii
162
do.
162
Mandible x 1 J
163
Cheekteeth x 6
Skull xf
172
do.
172
Skull X I
17s
do.
176
Cheekteeth X 3
176
Skull A I
179
do.
179
do.
180
Cheekteeth x 2 J-
180
Skull X I
183
do.
183
do.
184
Cheekteeth X3^
184
Skull X I
191
do.
192
Cheekteeth X4
192
Skull ■, I
204
do.
204
Skull X I
207
do.
207
Skull X I
210
do.
210
Skull X I
215
Cheekteeth xi\
216
Skull xr
222
Cheekteeth xzl
223
Skull X i\
228
do.
228
Skull X \
233
do.
234
LIST OF TEXT-FIGURES
FIG.
PAGE
70.
Lagostomus maxitmis, Desmarest
. Skull X 4
235
71-
do.
Cheekteeth X2j
235
72.
Caviella atistralis joannia, Thomas .
. Skull X 2
244
73-
do.
do.
24s
74-
do.
. Cheekteeth X 6
24s
75-
Hydrochoerus hydrochaeris, Linnaeus
. Skull xi
249
76.
do.
do.
250
77-
do.
do.
251
78.
do.
Cheekteeth x i \
252
79-
Aplodontia rufa, Rafinesque .
. Skull X I
256
80.
do.
do.
257
81.
do.
Mandible x i
Cheekteeth X4
257
82.
Belomys pearsoni trichotis, Thomas .
. Skull X 2
278
83-
do.
do.
278
84.
Trogopterus xanthipes mordax, Thomas
. Skull X 5
280
85.
Petaurista philippensis, Elhot .
. Skull X I
282
86.
do.
do.
283
87.
do.
Cheekteeth X 4 J,
283
88.
Pteromys volans, Linnaeus
. Skull X 2
292
89.
do.
do.
292
90.
do.
Cheekteeth X 5
293
91.
Myosciurus pumilio, Le Conte .
. Skull X 3^
314
92
do.
do.
314
93a. Myosciuriis pumilio, Le Conte .
. Skull X 2
315
b. Sciurillus pusilhis, Desmarest .
do.
c. ,, rnurimis, MiJller & Schlegel
do.
d. Callosciurus tenuis mrdus, Miller
do.
94
Sciurus vulgaris, Linnaeus
. Skull xi|
328
95
do.
do.
329
96
do.
Cheekteeth x 5
329
97
Rheithrosciurus macrotis. Gray
. Skull X I
394
98
do.
do.
394
99
Rhinosciurus laticaudatus tupaioides, Gray
. Skull X 2
397
100
do.
do.
397
101
Funisciurus pyrrhopus leonis, Thomas
. Skull X 2
410
102
do.
do.
411
103
do.
Cheekteeth X 8
411
XXIV
Liai UF Ih.Vl-FlUUKt
H
FIG,
PAGE
104.
Xcnis niti/iis, C'rctzchmar
. Skull XX J
419
105.
do.
do.
419
106.
Sptrniopliilopsis leptodact vliis. Lichtenstein
. Skull -I A
424
107.
do.
do.
424
loS.
Tcvnias dorsalis, Baird ....
. Skull ■,2i
427
log.
do.
do.
428
1 10.
CitcHiis citfUus, I.iiinaeus
. Skull . 2
43S
III.
do.
do.
438
112.
do.
Cheekteeth
■5
439
113-
Marmota marnuitii. Linnaeus .
. Skull • I
456
114.
do.
do.
456
115.
do.
Cheekteeth
. t
457
I If).
Cxnomvs ludoTiciamis. Ord
. Skull ■ I
4^"
117.
do.
Cheekteeth
■3
464rt
118.
Castor fiber, Linnaeus ....
. Cheekteeth
■ H
4*>4«
119.
do.
Skull X i
465(7
120.
Iletcriimys aiioimilus, Thompson
. Skull X 2*
473
121.
do.
do.
473
122.
do.
Cheekteeth
9
474
123-
Pcrognathiis hispidus, Baird
. Skull X3A
481
124.
do.
do.
48 1
125.
do.
Cheekteeth ,
17
4S2
126.
Dipodomvs merriaini iiieliinunis, Mcrriam .
. Skull X 2*
495
127.
do.
do.
495
12S.
do.
Cheekteeth
13
496
129.
T/ioiiiniiiys pcrpidlidiis, yi^vridm
. Skull X 2*
508
130-
do.
do.
509
131-
do.
Cheekteeth ;
,6
509
132-
Anomalurus fraseri jacksoni. de Winton
. Skull xii
53S
133-
do.
do.
53S
134-
do.
Cheekteeth
5
539
135-
Idiurus macriitis. Miller ....
. Skull ■ 3i
544
136.
do.
do."
545
'37-
do.
Cheekteeth
IS
545
138.
Pi'detcs surddstcr larval/s, lloWistev .
. Skull ■ I
55°
■39-
do.
do.
550
140.
do.
Cheekteeth
4
551
141.
C'tciiodiictyhis rtiiiidi, Rothman
. Skull xii
557
LIST OF TEXT-FIGURES
. Ctenodactylus gundi, Rothman
. Skull xij
557
do.
Mandible x i J
Cheekteeth x-j
558
. Sicista siiblilis loriger, Nathusius
. Skull X 4
565
do.
do.
565
do.
Cheekteeth X 9
566
. Zaptis hudsonius, Zimmermann
. Skull X3I
570
do.
do.
570
do.
Cheekteeth X 15
571
. Eiichoreutes naso, Sclater
. Skull X 3
578
do.
do.
578
do.
Cheekteeth x 1 1
579
. Allactaga euphratica, Thomas
. Skull X2i
58.
do.
do.
581
do.
Cheekteeth X 8
582
. Jacultis jaculus, Linnaeus
. Skull X2i
594
do.
do.
594
do.
Cheekteeth x 1 1
595
. Graphiurus hueti, Rochebrune
. Skull X2i
605
do.
do.
605
do.
Cheekteeth x 10
606
. Eliomys quercinus, Linnaeus .
. Skull X2i
614
do.
do.
614
do.
Cheekteeth x 10
615
. Dyrotnys nitedula, Pallas
. Skull X 3^
617
do.
do.
617
do.
Cheekteeth x 10
618
. Glis glis, Linnaeus . . . .
. Skull X 2
621
do.
do.
621
do.
Cheekteeth x 10
622
. Miiscardinus avellanarius, Linnaeus
. Skull X3 J
624
do.
do.
624
do.
Cheekteeth x 10
625
. Platacanthomys lasiurus, Blyth
. Skull X2.\
628
do.
do.
628
do.
Cheekteeth xio
629
. Typhlomys cinereiis, Milne-Edwards
. Skull X4
630
do.
do.
630
XXVI
LIST OK TEXI-FIUU
IRES
FIG.
PAGE
179.
Typhlomys cinereus, Milne-Edwards
. Cheekteeth X 14
63.
180.
Lophiom\s imhausi, Milne-Edwards .
. Skull:-: a
('33
181.
do.
do.
634
182.
do.
Cheekteeth x 5
634
183.
Spalax monticola dolbrogcac, Miller .
. Skull xii
639
184.
do.
Cheekteeth x 5
640
185.
Rhizomvs pruinosus, Blyth
. Skull xii
647
186.
do.
do.
647
187.
do.
do.
648
188.
do.
do.
64S
189.
do.
Cheekteeth x 5
649
(N.B. — In the figures of cheekteeth the left-hand figure (in a few cases
lettered a) shows the upper tooth row of the right side seen from below; the
right-hand figure (occasionally lettered b) shows the lower tooth row of the
left side seen from above.)
LIMITS OF THE ORDER RODENTIA
In 1912 (Science, New York, n.s. XXXVI, p. 285), Gidley proposed a
separate Order for the Rodentia Duplicidentata (Leporidae and Ochotonidae),
and restricted the Order Rodentia to the great mass of animals usually known
as Rodentia Simplicidentata. This division is currently accepted by American
authors, usually not so elsewhere (excepting Flower, 1927, Vertebrate List,
Zool. Soc. London, 1828-1927, Mammals, p. 239).
Whether it is a classification that is likely to be universally followed seems
open to question. Gregory' in his excellent work The Orders of Mammals, Bull.
Amer. Mus. X.H. XXVII, 1910, inclined to the contrary opinion, though
stating that no fossil forms have yet been discovered that will connect the
Duplicidentata with the Simplicidentata.
I can only say that in my opinion the Lagomorpha mav reasonably be
regarded as an Order distinct, and that for the purposes of the present work
they are regarded as such. The fundamental ditTerences in the appearance of
those parts of the skull to which jaw-muscles are attached may surely at once
be stated in the Lagomorpha to be a much more important character than the
retention of the functionless second upper incisor which seems to be quoted
always as the main difference between the two groups.
To those of the contrarv' opinion I must plead guilt)' of saying: notes on the
characters of three hundred and forty-five genera containing sixt\--four hundred
forms proved enough work tor one; I do not look for extra forms to include in
an Order as vast as this; and I have no intention of including in this Order
forms which may very well not belong there. Before including the Lagomorpha
in the Order, let us wait until an intermediate family is discovered fossil between
the two groups. Because the fact that both Rodents and Lagomorphs are
adapted for gnawing does not seem to prove conclusively that they must of
necessity be so nearly related as to be included in the same Order.
The Order Rodentia, therefore, as here understood, has been defined by
Miller & Gidley as follows:
"Terrestrial and fossorial, occasionally arboreal or semi-aquatic placental
mammals with both brain and placentation generalized in type; feet unguicu-
late; elbow joint always permitting free rotary motion of forearm; fibula never
articulating with calcaneum; masseter muscle highly specialized, divided into
three or more distinct portions, having slightly different functions; caecum
without spiral fold; dental formula not knowTi to exceed i. j, c. y, p. t,
m. 5=22 permanent teeth; incisors scalpriform growing from persistent pulp,
the enamel of upper tooth not extending to posterior surface ; distance between
mandibular and maxillar\- toothrows approximately equal, both pairs of rows
capable of partial or complete opposition at the same time, the primarj- motion
of the lower jaw in mastication longitudinal or oblique."
For further notes on the essential characters of the Order see TuUberg,
1 — Livine Rodents — I I
2 LIMITS OF THE ORDER RODENTIA
Uehcr das System der Nagetiere, 1899; Flower & Lydekker, .Mammals Lhing
and Extinct, 1891, pp. 443-448; Gregory, Orders of Mammals, 1910, p. 323;
Weber, Die Saugetiere, II, p. 238, 1928; Gidley, Science, n.s. XXXVI, p. 285
(the separation of the Order Lagomorpha from Rodentia, and differences between
the two Orders); Parsons, 1894, Proc. Zool. Soc. London, p. 251, Myology of
Sciuromorphine and Hystricomorphine Rodents, etc.
VARIATION
The great variation or diversity in structure within the Order is one of the
features that makes it such an interesting study; it is perfectly safe to say that
nowhere among all Orders of mammals is such div'ersity found in a single
Order with the possible exception of the Marsupials.
Great specialization has been attained independently again and again within
this order for various modes of life. Among these may be mentioned extreme
modification for aquatic life, most developed in Hvdromys and immediate allies
(Muridae, Hvdromvinae); Ichtlivomvs and immediate allies (Muridae, Crice-
tinae) ; to a lesser degree in Ondatra (Muridae, Microtinae), Myocastor (Echimy-
idae), and Castor (Castoridae); modification attaining a high degree for arboreal
life; two families contain "flying" genera, with a flying membrane which
enables them to take flying leaps from tree to tree (Sciuridae, Pteromys group),
and all genera of Anomaluridae except Zenkerella) ; in non-flying arboreal types,
the Erethizontidae show great specialization of the feet, the hallux sometimes
being replaced by a broad movable pad [Coendou, Echinoprocta, Chaetomys);
and many Muridae (Murinae) of the Indo-Malayan region have a fully opposable
hallux, as Hapalomys, Vandeleuria, Chiropodomys, Chiroinyscus, etc. -Many
genera are fully modified for underground life, with Mole-like appearance,
and either immensely developed incisors, or immensely developed claws
for digging. In the Muridae, examples are Ellobiiis and Prometheomys (Micro-
tinae), Xotiomys (Cricetinae), and Myospalax (Mvospalacinae). The Spalacidae
(Spalax) go a step further than any in that even the eyes are suppressed. The
Geomvidae, quite unrelated to the above, are all as highly specialized for
underground life as any Murine; as are the African Bathyergidae, another
isolated group. And in South America, certain Hystricoid genera, as Ctenomvs
and Spalacopus, have taken to this form of life and become just as modified in
external form. High specialization towards bipedal saltatorial life is developed
in three or possibly four unrelated families, with Kangaroo-like form and
elongated hindlimbs and tail; the Dipodidae (all genera except Sicista), the
Pedetidae, the Heteromyidae (Dipodomys and Microdipodops), and probably
one member of the Muridae, Xotomys (Murinae), from .\ustralia. Also perhaps
some Gerbils.
Cursorial specialization, with form like that of a primitive Ungulate, and
reduction of the digits of the hindfoot to three has taken place twice in South
-America, in the Dasyproctidae, and in the Caviidae (Dolkliotis).
Great specialization of the covering of the body into spines is shown in the
Hystricidae, being in this family at extreme development superior to that of
any other Rodent and perhaps any other Mammal. Other families which have
a more or less specialized spiny covering are the Erethizontidae (most effective!),
certain genera of Echimyidae (not highly developed), and certain Muridae, as
in some species of Rattus, and in Acomys; also to a lesser degree in the Cricetine
3
4 VARIATION
genus Neacomvs. One genus of Muscardinidae, Platacantliomys, is spiny. And
in two families of Rodentia extraordinarj' and abnormal development in the
skull has taken place, the Cuniculidae, with their enormous bony cheekplates,
and the Lophiomyidae, in which the temporal fossae are roofed in by bony
plates.
PREVIOUS CLASSIFICATIONS
OF ORDER
OLDFIELD THOMAS, 1896
(Proc. Zool. Soc. London, 1896, p. 1012)
(Suborder SIMPLICIDENTATA)
A. ANOMALURI
Family Anomaluridae
Anomahirus, Idiurus.
B. SCIURO.MORPHA
Family Sciuridae
Subfamily Sciurinae
(a) Rheithrosciurus, Xerus, Sciurus, Tamias, " Spermophilus"
( = Citellus), Cynomys, " Arctomys" ( = Marmota)
(b) Eupetaurus, Petaurista, " Sciuropterus" { = Pteromys)
Subfamily Nannosciurinae
Nannosciuriis
Family Castoridae
Castor
C. APLODONTIAE
Family Aplodontiidae
Aplodontia
D. MYOMORPHA
Family "Gliridae"
Subfamily "Glirinae"
Glis, Eliomys, Muscardinus, Graphiurus
Subfamily Platacanthomyinae
Phitacanthomys, Tvphlom \s
Family IMlridae
Subfamily Hydromyinae
Hydromys, Xeromys, Chrotomys
Subfamily Rhynchomyinae
Rhytichumys
Subfamily Phloeomyinae
Phloeomxs
Subfamily Gerbillinae
Gerbilliis, Pachyuromys, Psammomys, Meriones, Rhombomys
Subfamily Otomyinae
Otomys, " Oreinomys" { = Oiomys)
PREVIOUS CLASSIFICATIONS: THOMAS
Subfamily Dendromyinae
Deomvs, Dendromus, Stciitomxs, Mahiculhrix, l.cimiuomys
Subt'aniilv Miirinae
Miis, Xesukiti, Cricetomys, Malacoinys, Luplntromys, Saccostoiinis,
Acoinys, Ariicantliis, Cjohiiida, lande/eiiiiii, C/iiropudoinvs. liat-
omys, Carpomys, " Chiruromys" ( = Pogoiiomys), Hapalomvs,
Pitheclieir, Cniteromys, " Craiiiot/nlx" { — Echiot/irix). Miistiic-
onivs, I'romvs, Conihirus
Subfamily Lophiomyinae
Lup/lWI?!VS
Subfamily " Sigmodontinae"
"Hamster" { = Cricetiis), Mystromys, Bicuiivtcusoinvs, A'esomvs,
" Ihilloiiivs" { = .\eso?iiys), Braclivuromxs, Hvpoi;eomvs, Eliurus,
Gymnurumys, Onychomys, Peroinysctis, Rhipidomys, Tyloiiivs,
Holuchihis, Sigmodou, Oryzomys, Reithrudontomys, Eligmodontia,
Neoiomvs, Reithrotion, PhxUotis, Scapteroinys, Iclithyomys, Akodon,
Oxvmvctenis, Bhiiiiwinys, Xotiomxs
Subfamily Neotominae
Neotomii, Xenomys, Hodomys
Subfamily iXIicrotinae
Plienacoinys, " Erotoinys" (--C/ethilunuinys), Mkiutiis, Syiuipt-
oinys, Lemmus, Dicrostonyx, Ellohius
Subfamily " Siphneinae"
"Siphiieus" ( = Myospalax)
Family Sfalacidah
Subfamily Rhizomyinae
Rliizoinys, Tacliyoryctes
Subfamily Spalacinae
Spalax
Family Geomyidae
Geomys, Thomomys
Family IIeteromyidae
Subfamily Dipodomyinae
Dipodomys, " Perodipiis" { = Dlpodoiiiys), Microdipudops
Subfamily Heteromyinae
Perogniithus, Heteromxs
Family Bathvergiuae
Biitliyergus, Gcoiycliiis, " Myoscolops" (-^ Heliophohius), Hetero-
cepludus)
Family Dipodidae
Subfamily "Sminthinae"
" Smiiithus" ( = Sicista)
Subfamily Zapodinae
Zapiis
Subfamily Dipodinae
Dipiis, A/lactaga, " Platycercomys" (== Pygeietinus), Eiuhoreiites
PREVIOUS CLASSIFICATIONS: THOMAS 7
E. HYSTRICOMORPHA
Family Pedetidae
Pedetes
Family Octodontidae
Subfamily Ctenodactylinae
Ctrnodactylus, Massoiiliera, Pectinator, Petromvs
Subfamily Octodontinac
Ctenumys, Aconaemys, Spalacopus, Octudon, Ahrocuma
Subfamily "Loncherinae"
(a) Dactylomys, Thrinacodus, Kannabateomys, " Lonclwres"
( ^ Echimys)
[b) " Thrichomys" { = Cercomys), Cercomys, Carterodon, Mesomys,
" Echinomys" ( = Proechimys)
Subfamily Capromvinae
Myocastor, Capromys, Plagiodontia, Thryonomys
Family Hystricidae
Hystrix, Atherura, Tricky s
Family Erethizontidae
Subfamily Erethizontinac
Erethizon, Coendou
Subfamily Chaetomyinae
Chaetomys
Family Chinchillidae
Chinchilla, Lagidium, Lagostomus
Family Dasyproctidae
Dasyprocta, " Coelogenys" { = Cuniculus)
Family Dinomyidae
Dinomvs
Family Caviidae
Cai'ia, Dolichotis, Hydruchoerus
This classification is admittedly nothing more than a rearrangement and
bringing up to date of an earlier classification of Alston. It may at once be
discarded as unnatural, as being based mainly on the character of the fusion or
separation of tibia and fibula. In the "Myomorpha" of Alston these bones are
fused; in the " Sciuromorpha" and "llystricomorpha" they are separate.
Alston states that "in the few cases in which the cranial differences fail us in
separating the Sciurine Rodents from the Murine, and the latter from the I lystri-
cine, the complete ankylosis of the lower part of the tibia and fibula in the
second group comes to our aid." As Bathyergidae (which have Hystricine
mandible formation) are placed in " Myomorpha " on account of the fibula
structure, presumably this was considered the chief character in placing a Rodent
systematically.
But a Rodent did not become a Rodent because its fibula fused or remained
separate. If Flower and Lydekker's book Mammals Living and Extinct is
looked through with relation to the classification of other Orders, it may be seen
S PREVIOUS CLASSIFICATIONS: TULLBERG
that in one family of Insectivora, the Centetidae (now I believe known as
Tenrecidae), in one branch of the family the tibia and fibula are described as
fused (Oryzorictinae, p. 638), in the other branch they are distinct (Centetinae,
p. 637). It therefore these two conditions may exist in the same family of
Insectivora, the character is surelv one which can scarcely be used for super-
familv arrangement in another Order.
A Rodent becomes a Rodent because it gnaws, and its gnawing is done with
its incisors (which do not vary throughout the genera of modern Rodentia
sufficiently for any supertamily grouping to be arranged on this account),
and with its jaw-muscles; and the jaw-muscles have modified those portions
of the skull, to which they are attached, in various ways throughout the larger
groups, as recognized by all other authors who have comparatively recently
attempted a classification of the Order. I venture to suggest that if Raitiis,
for example, had never taken to gnawing, whatever the condition of its hindleg
bones it would not be classed in the present Order to-day; or if Orvsorictes had
bv chance taken to this form of life and developed the characteristic Rodent
skull, dentition, and jaw-muscle arrangement, whether or not its tibia and fibula
were fused, it would automaticallv have come under the heading of "Rodent"
as understood to-day. If the Order is to be dumped into three superfamilies
or "waste-paper baskets" the " Sciuromorpha," "Myomorpha," and "Hvstrico-
morpha" (into which as I see it the families will not naturally go), it should be on
a verv different basis from that of Alston and Thomas, and more like that of
Tullberg (below), with a separate group " Bathyergomorpha" for the Bathyer-
gidae, and probably one also for the Aplodontiidae, the Sciuromorph branch
to include Geomvidae, the Myomorph branch to include Ctenodactvlidae,
Anomaluridae and Pedetidae, etc.
TULLBERG, 1899
(Utber das .System der XaKcthierc. Nova .Acta Reg. Soc. Sci. Upsaliensis,
vol. XVIII, No. I)
(Suborder SIMPLICIDENT.'\TI)
Tribus I. HYSTRICOGN.Vnil
Subtribus i. BATHYERGOMORPHI
Family B.\thverg!d.\e
( Georychiis, Biithyergns)
Subtribus 2. HYSTRICOMORPHI
Family Hystricidae
{Hystiix, Athirura)
Family C.wiidae
(" Coelogenvs" { = CunkuIin), Dasypiocta, Ciivia,
Dolichotis. Hydiochoena)
Family Erethizontidaf.
{Erethizun, Coendoii, Chaetumys)
PREVIOUS CLASSIFICATIONS: TULLBERG 9
Family Ciiinchili.idae
(Chinchilla, Lagidium, Lagostomus)
Family " Ailacodidae"
("A ulacodus " ( = Thryonomys))
Family Echinomvidae
Subfamily " .Myopotamini "
(" .1 lyopotamus " ( = Myocastor))
Subfamily Echinomyini
Echinomyes
{" Echinomys" ( = Proechimys), " Nelomys" (based
on a species of Cercomys), Kaimabateomys)
Octodontes
("Habrocoma" { = Abrocoma), Octodon, Spalacopus,
Ctenomys)
F"amily Petromyidae
(Petromys)
Tribus2. SCIUROGNATHI
Subtribus i. MYOMORPHI
Sectio I. Ctenodactyloidei
Family Ctenodactylidae
{Ctenodactylus)
Sectio 2. Atwmaluroidei
Family Ano.maluridae
(Anoimilurus)
Family Pedetidae
(Pedetes)
Sectio 3. Myoidei
Subsectio i. IVIyoxiformes
Family "Myoxidae"
(Graphiurus, " Myoxus" { = Glis), Eliomys, Muscar-
dtnus)
Subsectio 2. Dipodiformes
Family Dipodidae
("Sminthus" { = Sicista), Zapus, Dipus (based on
Jaculus), Allactaga)
Subsectio 3. INIuriformes
Family Spalacidae
("Siphneus" [^ Myospalax), Spalax, Rhisomvs,
Tachyoryctes)
Family Nesomyidae
(Gymniiromys, Nesomys, Eliiirus, Brachyuromvs,
Brachytarsomys)
Family Cricetidae
[Cricetus)
lo PREVIOUS CLASSIFICATIONS: TULLBERG
Family LoPHiOMYiDAn
(iMphiovivs)
Family Arvicolidae
(Ellohius, Anicola, Neofiber, "Fiber" ( = Ondalra),
" Ciiiiiculus" (^- Dicrostonyx), " Mvodes" ( = Lem-
III lis))
Family Hesperomyidak
(Hespeioiiivs (based on Peioiiivsciis), Neotoma,
Siginodoii, Nectomvs, Oxvmvctenis)
Family Muridae
Subfamily Alurini
(Miis, Nesokia, Cliiropodomys, " Hapalotis" (based
on Notomvs), Hvdroiiivs, Dendioiiiiis, Steaiomvs,
Saccostoiiius, Cricetomvs, Deoiiivs, Lopliiiromvs)
Subfamily Phloeomyini
(Phloeomxs)
Subfamily Otomyini
(Otomvs)
Family Gerbillidae
(Gerbillus, Psammomvs)
Subtribus 2. SCILROMORPHI
Sectio 1. Sciiiroidei
Family ' ' H aplodontidae ' '
{" Haplodoii" ( = Aplodoiitia))
Family Sciuridae
(Sciurus, " Sciuropteiiis" {^ Pteromvs), " Pteromvs"
( = Petaurista), " Arctoinvs" ( = Marmota), Cynomvs,
" Spernioplii/us" { = Citelliis), Tainias)
Sectio 2. Castoroidel
Family Castoridae
(Castor).
Sectio 3. Geomyoidei
Family Geomyidae
Subfamily Dipodomyini
{" Perodipiis" (- Dipodoinvs), Dipodonivs, Perogna-
tlius, Heteromvs)
Subfamily Geomyini
(Geoiiivs, Tlwmoiiixs)
This is in my mind perhaps the best classification of the Order that has been
done. The only points which seem unnatural are the too close association of
Aplodontiidae with Sciuridae, the lumping together of all the Old World
Murine " borrowers " as a family Spalacidae (R/iizoiiiys, Myospahix, Tachy-
oryctes, Spalax), and the lumping together of the Dasyproctidae and Cuniculidae
with the Caviidae.
PREVIOUS CLASSIFICATIONS: WEBER ii
MAX WEBER, 1904, 2nd ed., 1928
(Die Saugetiere, 1928, II, p. 238)
(Living forms only)
(Suborder SIMPLICIDENTATA)
Tribus 1. HAPLODONTOIDEA
Family "Haplodontidae"
{Aplodontia)
Tribus 2. SCIUROIDEA
Family Sciuridae
(Sciurus, Neosciunis, Rheithrosciurus, Cullosciurus, Tamiasciurus,
Ratufa, Heliosciurus, Funisciuriis, Nannosciurus, Funambiihis)
Family Pteromyidae
(Eupetaurus, Pteromys {—Petaurista), " Sciuropterus" (=Pter-
omys), Glaucomys)
Family Xeridae
{Xerus, Geosciuriis)
Family Tamiidae
(Tamias, Eutaniias, Citellus)
Family Marmotidae
(Cynomys, Marmota)
Tribus 3. CASTOROIDEA
Family Castoridae
{Castor)
Tribus 4. GEO.MYOIDEA
Family Heteromyidae
{Dipodomys, Heteromys, Perognathus)
Family Geomyidae
{Geomys, Thomomys)
Tribus 5. ANOMALUROIDEA
Family Anomaluridae
(Anomaluriis, Idiurus, Zenkerella)
Family Pedetidae
(Pedeies)
Tribus 6. "AIYOXOIDEA"
Family "Myoxidae"
{Graphiurus, Muscardimis, Glis, " Dryoniys" {=Dyromys),
Eliomvs)
Family Platacanthomyidae
(Platacanthotnys, Txphlomys)
Tribus 7. DIPODOIDEA
Family Sicistidae
{Sicista)
12 PREVIOUS CLASSIFICATIONS: WEBER
Family Dipodidae
(Zapiis, Allactaga, " Scartiirus" {=AUactaga), Dipiis, Jacidus,
Euchoreules, Pygerettmis)
TribusS. MYOIDEA
Family Spalacidae
(Spalax, Rhizomys, Tachyoryctes, " Myotalpa" { = Myospalax))
Family Nesomyidae
(Brachxuroinxs, Nesomys, Braclivtarsoinys, Eliuriis)
Family Mlridae
Subfamily Cricetinae
(Cricetus, Mesocricetus, Crketulus, Mystromys, Hesperomys, Pero-
mvsciis, Oryzotuys, Reithrodoiitomys, Sigmodon, Tylomys, Holo-
cliilus, Xectomys, Eligmodontia, Ichthyomys)
Subfamily Lophiomyinae
(Lophiotnys)
Subfamily Microtinae
{Lemmus, Myopiis, Dicrostonyx (group Lemmi) ; EUobius (group
EUobii); " Erotomys" (=Clethrionomys), "Fiber" (=Ondatra),
Microtus, Pitymys, Arvicola (group Microti))
Subfamily Murinae
(" Epimxs" (--Rattiis), Mm, Apodemus, Mkivmys, Nesokia,
Pliloeomys, Pithecheir, Crketomys, Saccostoinus, Otomys,
" Oreomvs" (=Otomvs), Dendromus, Deomvs, Mastacomys, Lepo-
rillus, Uroiiivs, Mallomys, Conilurus, " Chiruromys" {^Pogo-
nomxs))
Subfamily Cit-rbillinae
(Gerbilbis, Pacliviiiomvs, Meriones, Rhombomys, Psammomys)
Subfamily Hvdromyinae
(Hydromys, Leptomys, Xeromys, Celaenomys. Chrntomys. Cru-
noiiivs, Rliyncliomys, " Craurothrix" (=Echiothrix))
Tribus 9. BATHYERGOIDEA
Family Bathyergidae
{Buthyeigits, Georychus, " Myoscalops" { --IJe/iophohiiis), Hetero-
ccpluilus)
Tribus 10. HYSTRICOIDEA
Family Hystricidae
{Ilystrix, Atherwa, Trichys)
Family Erethizontidae
(Eretliizvn, Coendoii, Cliaetoinys)
Family Cayiidae
Subfamily Dinomyinae
(Dinomys)
Subfamily Dasyproctinac
{"Coelogenys" (=Cuiiiciilus), Dasyprocta, Myoprocta)
PREVIOUS CLASSIFICATIONS: MILLER & GIDLEY 13
Subfamily Caviinae
(Cai-ia)
Subfamily Hydrochoerinae
{Ilydrochoenis, Doliclwtis)
Family Chinchillidae
(Chinchilla, Lagidium, " Vizcacia" {=Lagostomus))
Family Capromyidae
(Myocastor, Capromys, Plagiodontia)
Family Octodontidae
{Echiniys, Octodon, Ctenomys)
Family Ctenodactylidae
(Ctenodactylus, Pectinator, Petromys)
Family Thryonomyidae
( Thryonomys)
So far as Superfamily grouping is concerned, this classification is followed
in the present book, with some modifications, as for instance the separation
of Pedetidae from Anomaluridae; also following Tullberg I cannot credit that
the Ctenodactylidae should be referred to the Hystricoid branch [Petromiis is
here transferred to the Echimyidae), and also on account of intermediate forms
I am unable to find characters to keep the IMuscardinidae separate as a super-
family from the Muroidae.
Several of Weber's divisions into families appear unnecessary or unnatural,
as the dividing up of the Sciuroidea into five "families"; the retention of the
old family Spalacidae; the retention of the "family Nesomyidae"; the lumping
together of Dinomvs, Cuniculus and Dasvprocta with the Caviidae; and the
formation of a subfamily Hydrochoerinae including Dolichotis (obviouslv very
nearly allied to Cavia) against the Caviinae with Cavia only.
MILLER & GIDLEY, 1918
(Joum. Washington Acad. Sci., VIII, No. 13. p. 431, 1918)
The Order, including Rodentia as here understood only (the Lagomorpha
not included), is divided into five superfamilies based on zvgomasseteric
structure.
Superfamily SCIUROIDAE
Three-cusped series
Family Sciuridae
Subfamily Sciurinae (the entire familv except the two follow-
ing groups)
Subfamily Nannosciurinae
(Xannosciurus. Myosciurus, Sciurilliis)
Subfamily Pteromyinae (Flying-Squirrels)
Family Geo.myidae
Subfamily Entoptychinae (fossil)
[Entoptychiis)
14 PREVIOUS CLASSIFICATIONS; MILLER & GIDLEY
Subfamily Geomyinae (North American Pocket-Gophers)
Family Hetfro.myidae (North American Pocket-Mice and Kan-
garoo-Rats, Oligocene (HcUscomys) to Recent)
Four-cusped series
Family Ahjidacmidae (fossil)
{Adjidauino)
Family Eutypomyidae (fossil)
(Eutvpoviys)
Family Chalicomvidae (fossil)
(Chalicomvs (Steneofiber) and related genera; Tros^otio-
therium, Palaeocastor, Eiicastor and related genera)
F'amily Castoridae
(Castor)
Family Castoroididae (fossil)
(Castoroides)
Superfamily MUROIDAE
Three-cusped series
Family Muscardimdae
(Eliomvs, Dvromvs, Glis, Muscaidiniis, also Leitliia (fossil))
Four-cusped series
Family Ischyromvidae (fossil)
{Ischvioinys)
Family Cricetidae
Subfamily Cricetinae (Cricctinae, Sigmodontinae, Neotominae
and Nesomvinae of authors)
Subfamily Gerbillinae (Gerbillinae of authors)
Subfamily Microtinae (Microtinae of authors)
Subfamily Lophiomvinae
(Lopliiornvs)
Family PLATACANTHOMYinAE
(PldtacantJioinxs, Txphlomvs)
Family Rhizomyidae
Subfamily Tachyoryctinae
( Tachyoryctes)
Subfamily Rhizomyinae
{Rliizomvs and related genera)
Subfamily Braminae (fossil)
(Braiiiiis (-- EUohiui), a Microtine; see Hinton, Monogr.
\'olfs & Lemmings, I, p. S7, 1926)
Family Spalacidae
Subfamily Myospalacinae
(Myospahix)
Subfamily Spalacinae
(Spalax, Recent; Prospalax (fossil))
PREVIOUS CLASSIFICATIONS: MILLER & GIDLEY ,5
Family Muridae
Subfamily Dendromyinae (Dendromvinae of authors)
Subfamily Murinae (Murinae of authors)
Subfamily Phloeomyinae
(Phloeomys)
Subfamily (jtomyinae
(Otomvs)
Subfamily Hydromvinae (Hydromvinae of authors!
Superfamily DIPODOIDAE ' ' '
Three-cusped series
Group A
Family Pammyidae (fossil)
{Paramys, Mysops, Prosciiirus and related genera)
Group B o /
Family Graphiuridae
(Graphiuriis)
Group C
Family Allomyidae (fossil)
(Allomys, Haplomys, Meniscomys, Mylagaiilodon)
Family Aplodontiidae
{Aplodontia, Recent; Liodontia, fossil)
Family Cylindrodontidae (fossil)
(Cylindrodoii)
(Position of group doubtful)
Four-cusped series
Group A
Family Pseudosciuridae (fossil)
{Psendosciurus)
Group B
Family Mylagaulidae (fossil)
(Mylagaulus, Ceratogaulus, Epigaulus)
Group C
Family Anomallridae
[Auomalurus)
Family Idiuridae
Subfamily Idiurinae
(Idiurus)
Subfamily Zenkerellinae
(Zenkerella)
Group D
Family Scilravidae (fossil)
{Sciuravus)
Family Zapodid.\e
Subfamily Thcridomyinae (fossil) (Theridomyidae of authors)
Subfamily Sicistinae
{Sicista, Recent; }Eomys, fossil)
i6 PREVIOUS CLASSIFICATIONS: MILLER & GIDLEY
Subfamily Zapodinac
(Eozdpiis, Zapiis, Sapaeozapus)
Family Dipodidap,
Subfamily Protiiptvchinae (fossil)
(PriitoptycliHs)
Subfamily Dipodinac (Dipodidae of authors, who recognize
Zapodidae as a distinct family)
Family Ctknodactylidae
(Ctenodactylus and related genera)
Family Pedktidak
" (Pedctes)
Superfamily BA'l'I lYERGOIDAE
I'amily BATiiYnRciDAE (Bathyergidae of authors)
Superfamily HYSTRICOIDAE
Lateralis series.
Group A.
Family Hystricidae
Subfamily Hystricinae
(Hvs/n'x, A Clint /lion, Tlieciiriis)
Subfamily Atherurinae
(Atlicrurus, Trichys)
Family Erethizontidae
(New World Porcupines except Chaetomys; fossil genera,
Asteromvs. Eosteiromvs, PmasteironiYs, Steiroinvs; ?PliiomYS,
}iM('/llp/lioil!YS)
Family Echimyidae
Subfamily Echimyinae
(Spiny Rats, provisionally including Chaetoniys; Hutias
(Ciipromvs, Plagiodoiitiii, etc.); many extinct genera, among
them AciireiiiYS, Boromvs, BrotoiiiYS, Colpostciiiiui, Eociirdia,
Eoctodoii, GraphiiiiYS, Gyngiiophiis, Haplustroplia, Hetei-
opsoiiiYS, Homopsomys, Isobolodon, Prospaiiiomys, Prottidel-
plioiiiYS, Protacaremys, Sciiiinvs, Scleromys, Spaniomys,
StichiniYS, Strophostephamis, Tribodon)
Subfamily Octodontinae
(CteiioiiiYs, Octodoii, OctodoiitomYs. Spahicopus. Amongst
fossil genera are CeplinloiiiYS, Dicoe/ophoiiis, Eiicoe/oplionis,
LitoduiituiiiYS, NeopIuinmiiYS, Palaeoctodoii, Plilorumys, Pitli-
tiiioliiiiiYS, PlatoeoiiiYS, ScotuiiiYs)
Family Petromyidae
(PctrnmYs)
Family Myocastoridae
(MYociistor and related fossil genera)
Family Tiiryonomyidae
( Thryonomys)
PREVIOUS CLASSIFICATIONS: MILLER & GIDLEY 17
Family Dinomyidae
(Includes the living Dwomji and extinct genera Amblyrhiza,
Briaromys, Discolomys, Elasmodontomys, Gyriabrus, Me-
gamys, .Xeoepiblema, Olenopsis, Potamarchus, Tetrastylus)
Family Cunil(;lid.\e
[Cuniculus)
Family Hept.wodontidae (fossil)
{Heptaxodon; JMurenia)
Group B
Family Dasyproctidae (Dasyproctidae of authors with Cuniculus
removed, and Neoreomys (fossil) added)
Family Chinchillidae
(Chinchilla, Lagostomus, " Viscaccia" ( = Lagidium) ; extinct
genera: Euphilus, Perimys, Pliolagostomus, Prolagostomus,
Scotaeumys , Sphaeromys)
Family Abrocomidae
(Abrocoma)
Medialis series
Family Caviidae (Caviidac of authors, with Hydrochoerus and
allies removed ; extinct genera : Anchimys, \eoprocavia,
Orthoniyctera, Palaeocaiia, Phugatherium, Procardio-
therium)
Family Hydrochoeridae
{Hydrochoerus and extinct allies, Plexochoeriis, Prohydro-
choerus, Protohydrochoertis; perhaps Cardiomys, Cariodon
and Cardial herium)
Great attention has been paid to this classification, on which the present
work was originally based. It attends much more strictly to detail characters
than either those of Weber or Winge. But it seems to break down where the
"Dipodoidae" (for instance, Graphiurus) and Muroidae are compared; it
does not appear good classification to separate Graphiurus from the Muscar-
dinidae so far that it is placed in another superfamily; moreover, it appears
that the Murine genus Deomys has the "Dipodoid" zygomasseteric structure
as defined by Miller & Gidley, and should be referred to that superfamilv if this
classification was maintained. As already noted by Wood in his monograph of
Heteromyidae, there is a very wide distinction between the zygomasseteric
structure of Paraniys, and the Mylagaulidae and that of the Dipodidae, both
referred to "Dipodoidae" of Miller iSc Gidley; from descriptions Paramvs and
Mylagaulidae have similar zygomasseteric structure to that of Aplodontia, here
considered the most primitive living Rodent as regards this arrangement;
Wood states, "The character of zygomasseteric structure as given by Miller
& Gidley . . . includes widely different types, which appear to have reached
their present condition in widely different manners. In Paramys and the
Mylagaulidae . . . the zygomatic plate is nearly horizontal because that is the
primitive condition for Rodents, and the growing masseter has not as yet
2 — Living Rodents — I
i8 PREVIOUS CLASSIFICATIONS: WINGE
effected any great change. In the Dipodidae . . . the zygomatic plate is hori-
zontal because the masseter has passed through the infraorbital fenestra and,
on expanding, has forced the zygoma down until it becomes even lower than
in the primitive forms."
In Miller & Gidley, the families appear in some cases to be over-split. For
the undesirahility of separating "Cricetidae" from Muridae, see Hinton,
Monogr. \'oles & Lemmings, 1926, p. 121.
WINGE, 1924
(Pattcdyr Slaegter, vol. II, Rodcntia, p. i)
The Order is divided into nine families, one of which, the Leporidae,
corresponds to the Suborder Duplicidentata or Order Lagomorpha.
1. Leporid.M'
(Leporini: Palacolagus (fossil), Lepus ; "Lagomyini": " Lagomys" =
Ocliotoiia)
2. "H.'\plodontidae"
AUomyini (fossil)
AUomys
Ischyromvini (fossil)
Paramys, Sciuravus, Ischyromys
Mylagaulini (fossil)
Mxlagaidus, Ceratogaulus
"Haplodontini"
" Hapludoii" = Aplodontia (sole living genus)
3. Anomaluridae
Pseudosciurini (fossil)
Pseiidosciurus, Sciuroides
Trechomyini (fossil)
Trecliovivs
Anomalurini
Aiwmalunis, " Aet/innis" = Zenkerelhi, Idiurus
Theridomyini (fossil)
Tlieridomys, Issiodoromys, Archaeomys
Pedetini
Pedetes
4. Dipodidae
Eomyini (fossil)
Eomys
Dipodini
" Sminthi"
" Simiithus" = Sicista ; "Jactilus" based on Zapus
Euchoreutae
Etichoreiites
Dipodes
" Scirtetes" = Allactagu, Dipii^
PREVIOUS CLASSIFICATIONS: WINGE ,9
Spalacini
Spalax
5. "Myoxidae"
Graphiurini
Graphiurus
"Myoxini"
"Myoxi"
Eliomys, Leitbia (fossil), Hypnomys (fossil), " Myoxus" = Glis
Muscurdiniis
Platacanthomyes
Platacantliomys
6. MURIDAE
Rhizomyini
Cricetodontes (fossil)
Cncetodon, Eumys
Rhizomyes
Nesomys, Brac/iyfarsomys, Gvmnuromys, Eliurus, Brachvuromvs
lachyoryctes, Rhizomys ' '
Cricetini
Criceti
Cricetus, Calomyscus, Lophiomys, " Siphneiis" = Myospdax
Hesperomyes
Hesperomys, Sigmodon, Neotoma, '' Habrothrix" (apparently
based on_ Akodon), Oxymycterus, Ichthvomys, Scapteromys,
^alomys (species or genera referred to this group not clear)
Hhtptdomys, Nectomys
Arvicolae
''Hypudaeus" (=Cletlmonomys plus Dolomys plus Phenacomys),
htber =Ondatra, Ellobiiis, Arvkola (including Microtus)
Dtcrostonyx, " Myodes" = Lemmus
Murini
Mures
Mus, " Spalacomys" =. Xesokia, PIdoeomvs, Crateromvs, Carpnmys
tchiotmx, Rhynchomys, Leiwmys, Vandekuria, Chiropodomys
Hapalomys; C/uiuromys, Notomvs, Mastacomvs, Zvzomvs, - Hapa-
lotis" = Comhtrus ; Acomys, Cricetomys, Saccosto,nus,\Steatomys,
Dendromiis, Deomys, Oenomys, Otomys
Gerbilli
Gerbillus, Rliombonns
Hydromyes
Xeromys, Crunomys, Hydromys, Chrotomys, Celaenomys
7. Hystricidae
Bathyergini
Bathyergiis, Heterocephalus, Georychus, Heliophobius
20 PREVIOUS CLASSIFICATIONS; WINGE
Hystricini
Hystrices
Triciiys, Athenira, llvstiix
" Sphinguri"
Steiromys (fossil), Eietliisoii, " Sphtnguriis" = Coendou, Chaetomys
Capromyini
" Aulacodus" = Thryonomys, " Myopotamus" = Myocastor, Cap-
romys, " Plagiodon" — Plagiodontia ; hobolodon (fossil)
Ctenodactylini
Petro)iivs, Pectinator, Ctenodactylus
Dasvproctini
Dinomyes
Dinomys ; Elasmodontomys (fossil)
Dasyproctac
Dasypiocta ; Xeoicomys (fossil); "Coelogeiiys" =Ciinicii!iis
Caviae
Sclu'stomvs, Eocardia (both fossil), Cavia, Doliclio/is, Hvdio-
clwerus
"Eriomyini" = Chinchillini
"Eriomves"
" Erionivs" =Cliiiic/iil/a, LagidiiDii
Lagostomi
Scotaeumxs, Peiimvs (both fossil), Lagostomus.
Octodontini
Echinomyes
Cercomys ( = what?, not Cercomys as here understood), Dacty-
lomvs, Thrinacodus, " Lasitiiomys" — Isotlirix, " Loiicheres" =
Echimxs, " Echinomxs" = Proecliimxs, " Nelomvs" =Cercomys,
Mesowys (based evidently on Clyomys and Eioyzygomatomys);
Carterodvn
Octodontes
Acareinxs, Sciainys (both fossil); Dicolpviiivs (fossil); " Hiihro-
coina" = Abrocoiiiti, " Sclihodon" = Acontiemys, Spalncopus, Octo-
don, Ctenomys
8. SCIURIDAE
Castorini
Eutxpomys, Steneofiher, Euliapsis (all fossil), Castor; Trogon-
therium, Ccistoioides (fossil)
Sciurini
Sciuri
Tamias, Otospennopliilus, Sciurus, Pteromys, Eiipctminis, Xeriis
" Arctomyes"
" Arctomys" — Maniiota, " Spennoplulus" =Citellus, Cynomys
9. " Saccomyidae" = Heteromyidae
Gymnoptychini
GymnoplychiiS { = Adjidniimo? fossil)
PREVIOUS CLASSIFICATIONS: WINGE 21
"Saccomyini"
Heliscomys (fossil), " Saccomys" = Heterotnys, Perognathus, Dipo-
domys
Geomyini
Pleurolichus, Entoptychus (both fossil), Thomomys, Geomys.
"I Form af Stamtrae" (phylogenetic tree)
Myoxidae Mujidae Saccopyidae
Dipodidae Hystricidae Sciiirjdae
Anomaluridae
Haplod Dntidae
Leporidae
The theory behind this work is brilliant, but the detailed characters are
incorrect many times, and the nomenclature is deplorable. The classification
of the "Hystricidae" seems in particular unnatural, with the inclusion of the
Bathyergidae, the lumping together of Hystricidae and Erethizontidae, the
whole family based more or less on the formation of the paroccipital process
which in the British Museum material seems at any rate in Dactylomys and
Capromys a variable character even within a genus. Further, if I understand
the work rightly, the infraorbital foramen is supposed to transmit muscle (or
to have transmitted muscle, which cannot be proved), in many groups in which
it does not do so (Sciuridae, Castoridae, Geomyidae, Heteromyidae, Bathyer-
gidae). Pedeta is regarded as with "fibula free," which is not so; M.i is re-
garded as larger than M.2 in Cricetini and Murini, which is not constant
{Anisomys agreeing with Winge's Rhizomyini in this character, as do some
Neotropical Cricetines), etc., etc.
OUTLINE OF CLASSIFICATION
HERE ADOPTED
The present classification is a combination as tar as possible of points which
appear to he correct from the five classifications Hstcd above. It is based on
characters which are constant throughout all the various genera referred to each
familv, and has been the subject of great care and observation.
The Order is here divided into twelve superfamilies. In case it may be
thought that too man)' of these are retained, and that forms which share charac-
ters have been too widely separated, I would quote a passage from Wood,
who in his review of fossil and recent Heteromyidae, states, "The most important
point to be emphasized is that ' parallelism, parallelism, more parallelism and
still more parallelism' is the evolutionary motto of the Rodents in general."
I have already stated that the Order is dominated to such a degree by one
family, the Aluridae, that it is necessary to devote a separate volume to that
family alone. It is necessary, therefore, for me to place this group last on the
list (certainly not because it is considered the most highly specialized!); I,
therefore, take first the group called Hystricognathi hv Tullberg, in which the
mandible is more highly specialized, and last the Sciurognathi of Tullberg,
which includes the Muridae, and in which the mandible is comparatively
unmodified.
With regard to the structure of the tibia and fibula, the following skeletons
have been examined :
"Fibula reduced, fully fused with the tibia in lower portion"
Pedetidae : Pedetcs
Bathyergidae: Batln'i'igiis, Georychus
Geomyidae : Geomys
Spalacidae : Spala.x
Dipodidae: Alactagulus, Allactaga, Jaculus, Sicista
Lophiomyidae : Lophiomys
Aiuscardinidae: Glis, Eliomys, Muscaidiiiiis
Muridae. (As this group is not dealt with in the present volume, less
attention has been paid to them, as they are clearly separable
from other families without this character. However, it has
been checked in a very small number: Mus and Apademiis
representing Murinae; Mesociicetus and PJiodopus representing
Cricetinae; Gcrbilliis and Pacliyurumys representing Gerbillinae;
and Tachyoryctes and Brachyiiromys representing Tachyoryctinae.)
All examined are perfectly clear in this character, with the exception of Glis,
in which the bones are widely separate, and only fused at extreme base.
No Heteromyidae are available for examination, but Dipodoinys as figured
by Howell agrees with the other Rodents.
OUTLINE OF CLASSIFICATION HERE ADOPTED 23
" Fibula not reduced, not fully fused in lower portion w ith the tibia "
Ctenodactylidae : Clenodactylus, Pectinator
Anomaluridae : Anomahtrus
Aplodontiidae : Aploduntia
Castoridae: Castor
Sciuridae : Petaiirista, Citellus, Spermophilopsis, Sciurus, Xenis, Tamias
and Marmota
Echimyidae : Dactylomys, Octodon, Aconaemys, Ctenomys, Capromys,
Sivocastor, Tlir\onom\s, Isothrix, Petromus
Erethizontidae : Erethizon, Coendou, Chaetomys
Dasyproctidae : Dasyprocta
Cuniculidae : Cunicithis
Hystricidae: Hystrix, Tlieciirus, Atherurits
Caviidae: Caiia, Dolichotis, Hydrochoerus
Chinchillidae: Chinchilla, Lagidium, Lagostomus
No skeletons of Dinomys nor of the Idiurinae are available.
In this group. Castor appears in the adult to have fusion suggested in these
bones, though not complete, and the Hystricoid Myocastor seems precisely
similar, another interesting case of parallel development in these two un-
related genera. The fibula is becoming reduced in Ctenomys. But much the
most interesting result obtained is that in the skeletons seen of the three genera
of Chinchillidae, Chinchilla, Lagidium and Lagostomus, the fibula, though still
not fully fused, is excessively reduced, slender and threadlike; the reduction
reaching an extreme degree in Lagostomus, which one might have expected to
be not the case.
It must be noted that, with regard to the formation of the forefoot, the ex-
treme reduction, almost or completely to disappearing point of the pollex, is
such a usual feature that no attention has been paid in the classification to this
character.
HYSTRICOGNATHI
Lower jaw highly specialized, either by distortion outwards of the angular
process by masseter lateralis, or by a conspicuous ridge extending along outside
of mandible below level of the toothrow for attachment of masseter medialis.
Angular portion of mandible never pulled inwards (inflected), and lower incisor
root never forming conspicuous knob beside condylar process.
1. Bathyergomorph Series
Infraorbital foramen not or scarcely transmitting muscle.
First Superfamily. BATHYERGOID.\E
Mandible with angular portion strongly distorted outwards by specialized
limb of masseter lateralis. Infraorbital foramen not or scarcely transmitting
24 OUTLINE OF CLASSIFICATION HERE ADOPTED
muscle. Zygomatic plate narrow, completely beneath the infraorbital foramen,
not broadened for attachment of masseter lateralis.
Skull much modified for fossorial life. Jugal moderately long, or in Hetero-
cephalus shortened, and Murine. Kullae without special peculiarities.
Cheekteeth i, t, or in Heliophohius, at full dentition, ;'; (in this genus
the cheekteeth are normally not all in place together). Cheekteeth rooted, but
extremely hypsodont, near simplification in pattern.
A tendency present for the upper incisors to extend behind the toothrow,
at extreme development into the pterygoids.
Fibula fully fused with the tibia below. Digits of hindfoot five. External
form much modified for underground life.
Malleus and incus fused (Tullberg). Radiale and intermedium separate
(Tullberg; fused in all other members of the Order examined by him except
Ctenodactylidae).
Family i. BATHYERGIDAE
With the characters of the Superfamily.
Group Bathyergi
Bathyergus, Heliophobius, Georychiis, Cryptomys.
Group Heterocephali
Heterocephahis.
(A key to all generic groups here recognized will be given below, when
dealing with the families.)
2. Hystricomorph Series
Infraorbital foramen much enlarged for muscle transmission.
Second Superfamily. HYSTRICOIDAE
Mandible with angular portion distorted outwards by specialized limb of
masseter lateralis. Infraorbital foramen very large, always transmitting muscle.
Zygomatic plate remaining narrow, and beneath the infraorbital foramen, not
broadened for attachment of masseter lateralis.
Skull usually broad, lacking interorbital constriction in the majority.
Paroccipital process usually well developed, prominent. Jugal normally not
approaching lachrymal ; frequently with downwardly or upwardly directed
process present. Bullae variable; in certain groups much inflated.
Cheekteeth |, the premolars not reduced in size in the majority; the
pattern flatcrowned, reduced heptamerous, laminate, or sometimes approaching
complete simplification.
Fibula not becoming fused with the tibia high on the leg, and usually, but
not alwavs, not specially reduced. Digits of hindfoot 5, 4, or 3.
Malleus and incus fused (Tullberg).
OUTLINE OF CLASSIFICATION HERE ADOPTED 25
Family 2. ECHIMYIDAE
Cheekteeth rooted or rootless; when rootless or strongly hypsodont the
pattern not a series of transverse plates. Digits of hindfoot 5, excepting
Thryonomyinae. Bullae well inflated, sometimes extremely so. Feet never
abnormally specialized for arboreal life. Spiny covering present or absent,
but when present, never to an extremely specialized degree. External form
never modified for cursorial life. Paroccipital process long, either curved
forwards under the bullae, or lengthened and standing apart from bullae.
Zygoma very generally complex, with downwardly or upwardly directed
processes present.
Subfamily FXHI.MYINAE
Cheekteeth not hypsodont, and not simplified in pattern; usually the pattern
is reduced heptamerous. Bullae not abnormally inflated, except Clyomys.
Paroccipital process curved forwards under the bullae. Externally Rat-like
or slightly modified for arboreal life, sometimes with spiny covering developed.
Ecliimys, Isothrix, Diplomys ; Proecliimys, Hoplomys, Cercomys, Euryzygma-
tomys, Clyomys, Carterodoti, Mesomys, Lotichothrix.
Subfamily CAPROMYINAE
Like the Echimyinae, but cheekteeth evergrowing, the re-entrant folds well
filled with cement, so that simplification of pattern is suggested. Upper molars
with more than one external re-entrant fold. Paroccipital process usuallv but
not always standing apart from bullae. External form arboreal or terrestrial;
fur not developing spines.
Procapromys (not seen), Capromys, Geocapromys.
Subfamily PLAGIODONTINAE
Cheekteeth evergrowing, the upper series with only one external re-entrant
fold, which is unusually deep and placed obliquely; the single inner re-entrant
fold also unusually deep, and running parallel to the outer fold. Paroccipital
process considerably lengthened. Zygoma simple.
Plagiodontia.
Subfamily DACTYLOMYINAE
Cheekteeth rooted, abnormally broadened, nearly prismatic in appearance,
and evidently with pattern not changing much during life. Paroccipital process
either standing apart from the bullae, or curved forwards under them. Habits
(said to be) arboreal; a tendency present towards elongation of the two central
digits of forefoot and hindfoot.
Thrinacodus, Dactylomys, Kannabateomys.
Subfamily MYOCASTORINAE
Cheekteeth extremely hypsodont, but not evergrowing, reduced heptamerous
in pattern, the pattern changing little during life. Bullae reminiscent of the
26 OUTLINE OF CLASSIFICATION HERE ADOPTED
type found in Castoridac, though less specialized than in that family. Skull
prominently ridged for attachment of muscles. External form considerably
specialized for aquatic life. Paroccipital process much lengthened, the lateral
process of paroccipital process enlarged.
Mvocastur.
Subfamily ABROCOMINAE
Cheekteeth evergrowing, the upper series more or less simple, the lower
series prismatic, and complex. Auditory bullae much inflated, the paroccipital
process curving forwards under them. Part of lachrymal canal open on side
of rostrum.
Al'iocoma.
Subfamily OCTODONTINAE
Cheekteeth evergrowing, both upper and lower series nearly or completely
simplified. Bullae and paroccipital process as in Abrocominae. No part of
lachrymal canal open on side of rostrum. External form generalized or modified
for underground life.
Octoinvs, Acoiiaemys, Octodoii, Octodontomys ; Spahicopus ; Ctenoiiiys.
Subfamily PETROMYINAE
Cheekteeth strongly hypsodont, nearly complete simplification in pattern,
but not evergrowing. Auditory bullae much inflated. External form without
special peculiarities.
Petromus.
Subfamily TMRYONOMYINAE
Digits of hindfoot reduced to four. Cheekteeth moderately hypsodont,
rooted, the re-entrant folds well marked and surrounded by heavy enamel.
External form heavy, terrestrial-fossorial. Auditory bullae of moderate size;
paroccipital process lengthened, standing apart from the bullae. Incisors
much thickened, the upper ones heavily three-grooved. Skull massive, ex-
cessively prominently ridged for muscle attachment. Shoulder-blade abnormal
(TuUberg).
Tlirvoiionivs.
Family 3. DINOMYIDAE
Cheekteeth evergrowing (?) or excessively hypsodont, the pattern a series of
transverse plates. External form heavy, terrestrial. Digits ot hindfoot four.
(Incisors thick, bullae moderate, paroccipital process not lengthened, no part of
lachrymal canal open on side of rostrum, and angular portion of mandible
powerfully distorted outwards, compare Chinchillidae.)
Dinoinys.
Family 4. ERETHIZONTIDAE
Externally more specialized than in the Echimyidae; feet becoming abnor-
mally modified for arboreal life, the hallux in progressive forms being replaced
with a broad movable pad (and skeleton of foot correspondingly much modified).
OUTLINE OF CLASSIFICATION HERE ADOPTED 27
Tail muscular, prehensile in progressive genera. Fur conspicuously spinous,
the spines not long, and not modified into thick circular quills. Bullae relatively
large; paroccipital process not lengthened. Zygoma usually simple. Check-
teeth typically with wide re-entrant folds (parallel — .\nomaluridae), or in C/iae-
tornys with structure much as in complex-toothed Echimyinae.
Subfamily CIIAETOMYINAE
Orbit almost surrounded by greatly thickened jugal and short postorbital
process. Cheekteeth with narrow re-entrant folds. Spiny covering weakly
developed. (Feet at highest specialization.)
Chaetomys.
Subfamily ERETHIZONTINAE
Orbit large; skull without postorbital process; jugal not specially thickened.
Cheekteeth with very wide re-entrant folds. Spiny covering at maximum
specialization. (Feet moderately to extremely specialized.)
Erethizon, Echinoprocta, Coendou.
Family 5. DASYPROCTIDAE
External form highly modified for cursorial life; digits of hindfoot reduced
to three ; clavicles suppressed. Part of lachrymal canal open on side of rostrum.
Bullae relatively large; paroccipital process not specially lengthened. Cheek-
teeth strongly hypsodont, but not evergrowing, the re-entrant folds narrow,
isolating on crown surface with wear as narrow islands.
Myoprocta, Dasyprocta.
Family 6. HYSTRICIDAE
External form heavy, terrestrial; digits of hindfoot five. Fur always con-
spicuously spinous, in progressive species attaining specialization in this respect
not known elsewhere in the Order. Tail always with modified bristles or quills
present. Spines of bodv usually partly modified into thick circular quills.
Skull in progressive species characterized by great inflation of nasal bones.
Auditory bullae relatively small. No part of lachrymal canal open on side of
rostrum. Paroccipital process not specially lengthened. Zygoma simple.
Cheekteeth moderately to extremely hypsodont, but not evergrowing, their
pattern paralleling that present in Dasyproctidae. Clavicles present but in-
complete. Lungs abnormal (Tullberg). Centrale not free (Tullberg; this
character unique in the Order so far as known except Cuniculidae).
Group Atheruri
Trichys ; Atlieruriis.
Group Hystrices
Thecurus, Ilystrix.
28 OUTLINE OF CLASSIFICATION HERE ADOPTED
Family 7. CUNICULIDAE
Skull extremelv modified bv growth of conspicuous bony cheekplate, a
structure not known elsewhere in the Order. Cheekteeth strongly hypsodont,
but not evergrowing, their pattern hke that present in Dasyproctidae and
Hystricidae, but rather more complex. External form heavy, terrestrial.
Digits of hindfoot five. Clavicles present, but incomplete. Paroccipital process
considerably lengthened. Centrale not free (Tullberg; on this character see
remarks under family Hystricidae).
Cuniciihis.^
Family 8. CHINCHILLIDAE
Cheekteeth evergrowing, the pattern a series of transverse plates. Lower jaw
with angular portion rather weaklv distorted outwards, the jaw to a certain degree
transitionar}' towards that of the Cavioidae. Digits of hindfoot four or three
(probablv three functional only in all genera). Jugal tending to approach the
lachrymal, or to come in contact with it. Some part of lachrymal canal open
on side of rostrum. Incisors relatively thin. A tendency present towards
extreme inflation of mastoids and bullae. Paroccipital process relatively short
(Chinchilla, Lagidium), or considerably lengthened (Lagosioiiiiis). Fibula
extremely slender, much reduced (skeletons of the three genera have been
examined).
Group Chinchillae
Chinchilla, Lagidiuin.
Group Lagostomi
Lagostomus.
Third Superfamily. CAVIOIDAE
Essential characters as in Hystricoidae except : angular portion of mandible
not distorted outwards by specialized limb of masseter lateralis; masseter
medialis the chief agent in modifying form of lower jaw, the outer side of which
has a long and conspicuous ridge extending below and beside toothrows for
attachment of this muscle. Cheekteeth evergrowing, relatively simple, but
with sharp folds present, the effect more or less prismatic. Malleus and incus
fused (Tullberg).
Family 9. CAVIIDAE
With the characters of the Superfamily. Paroccipital process moderately to
extremelv enlarged; bullae normally prominent. Cheekteeth strongly uni-
laterally hypsodont. Digits of hindfoot reduced to three; external form ambu-
latory or cursorial. Clavicles suppressed. Tibia and fibula as in normal
Hvstricoidae. Part of lachrymal canal open on side of rostrum, except in
Dolicliotis.
^ = "Coelogeiiys.''
OUTLINE OF CLASSIFICATION HERE ADOPTED 29
Subfamily CAVIINAE
Paroccipital process not excessively lengthened. M.3, upper series, not
enlarged. Pattern of cheekteeth relatively simpler. Palate shortened from
before backwards.
Group Caviae
Cavia, Galea, Caviella ; Kerodon.
Group Doiichotides
Dolichotis.
Subfamily HYDROCHOERINAE
Paroccipital process excessively lengthened. M.3 upper series extremely
enlarged and elongated. Cheekteeth with more complex pattern. Palate not
shortened from before backwards. (Size largest in the Order.)
Hydiochoerus.
SCIUROGNATHI
Lower jaw not highly specialized, never with the angular portion distorted
outwards, and never with long conspicuous ridge extending below level of
toothrow for attachment of masseter medialis. Angular portion of mandible
may be stronglv pulled inwards (inflected). In some genera, the root of the
lower incisor forms conspicuous knob beside the condylar process.
1. Sciuromorph Series
Infraorbital foramen not or scarcely transmitting muscle.
Fourth Superfamily. APLODONTOIDAE
Infraorbital foramen not transmitting muscle. Masseter lateralis not ex-
tending attachment on outer side (forepart) of zygomatic plate, which remains
narrow and unspecialized, completely below the infraorbital foramen. Mandible
with angular portion inflected to an abnormal degree.
Skull fossorial in aspect. Jugal lengthened. Bullae with neck directed
horizontally outwards, and region of braincase greatly widened.
Cheekteeth ;, evergrowing, near complete simplification of pattern.
Fibula not reduced, nor fused with the tibia high on the leg. Malleus and
incus free (Tullberg).
Family 10. APLODONTIIDAE
With the characters of the Superfamily.
Aplodontia.
(This family is doubtfully referred to the Sciuromorph series, and is regarded
as one of the most isolated and primitive living Rodents.)
30 OUTLINE OF CLASSIFICATION HERE ADOPTED
Fifth Superfamily. SCIUROIDAE
Infraorbital foramen not or scarcely transmitting muscle. Masseter lateralis
superficialis with anterior head distinct from zygoma, and masseter lateralis
extending its line of attachment on to zygomatic plate, which is to a greater or
lesser degree broadened and tilted upwards, the muscle typically rising upwards
on zygomatic plate to superior border of rostrum. Mandible with angular
process sometimes strongly inflected (Cynomvs); usually with a tendency for
this formation to be present.
Skull with, in progressive species, well developed postorbital processes
present (these processes always traceable). Jugal long, usually approaching
or reaching the lachrymal. Bullae without special modifications, usually
prominent.
Cheekteeth I or 4, cuspidate, very rarely approaching simplification,
in which cases (Larisciis, Rheithrosciurus) the original pattern may usually be
traced; the pattern normally a series of transverse ridges and corner cusps
(in the upper series), the lower series most often basin-shaped, with cusps at
each corner. Cheekteeth rooted, brachyodont or hypsodont.
Fibula not fused with the tibia. Digits of hindfoot five. External form
modified for arboreal or terrestrial life; a flying-membrane may be present,
attached to sides; tail always completely haired.
Malleus and incus free (Tullberg).
Family 11. SCIURIDAE
With the characters of the Superfamily.
Group Pteromyes
Belomvs, Trogoptenis, Pteromyscus, Petaiirista, Aeromys, Pteiomys, Hyhpetes,
Petinomxs, Eoglaucomys, Glaucomys, Petaurillus, lomys, Eupetaurus .
Group Sciuri
Mxosciurus, Xaniwsciunis, Saurillus ; Microsciurm, Syntheosciurus, Sciunis,
Tamiasciurus, Callosciwus, Funambuliis, Dremomys, Ratiifu, Menetes, l.arlsciis,
Glxphotcs, Rheithrosciurus, Rliinosciiirus, Hyosciurus, Heliosciurus, Paraxcrus,
Funisciurus, Protoxcrus, Myrsilus, Epixerus ; Atlantoxerus, Xerus, Spermo-
philopsis ; Sciurotamias, Tawias ; Citelliis, Marmota, Cynomys.
Sixth Superfamily. CASTOROIDAE
Zygomasseteric structure essentially as in Sciuroidae. Mandible without
special peculiarities.
Skull with no well-marked postorbital process; jugal lengthened, approach-
ing lachrymal, and extremely broadened anteriorly. Bullae with neck directed
outwards and upwards.
Cheekteeth \, extremely hypsodont, but not evergrowing, the pattern
OUTLINE OF CLASSIFICATION HERE ADOPTED 31
flatcrowned, reduced heptamerous, the enamel folds narrow, the pattern
chan^infj little during life.
I'ihula not fused with the tibia, but tending to become reduced, so that in
adult life fusion is suggested.
Digits of hindfoot five. Externally much specialized for aquatic life. Caudal
vertebrae broadened; tail naked, much broadened and flattened (unique in the
Order in structure and appearance).
Malleus and incus free (Tullberg).
Family 12. CASTORIDAE
With the characters of the Superfamily.
Castor.
Seventh Superfamily. GEOMYOIDAE
Zygomasseteric structure closely resembling that of the Sciuroidae; infra-
orbital foramen extremely reduced, and rather more modified; mandible with
angular portion somewhat reduced. Large externally-opening cheekpouches
present. Skull highly fossorial (Geomyidae), or becoming specialized by ex-
treme inflation of auditory bullae and braincase, and narrowing of rostrum
("saltatorial type") (progressive Heteromyidae). Jugal always short; sometimes
the zygomatic arch may be complete without it; in Heteromyidae, the whole
zygoma is threadlike.
Cheekteeth }, hypsodont, usually near complete simplification of pattern,
and often evergrowing.
Fibula (so far as known) fully fused with the tibia, high on the leg. Digits of
hindfoot five, or in saltatorial genera may be reduced to four.
Malleus and incus free (Tullberg).
Family 13. HETEROMYIDAE
Mastoids in progressive genera greatly inflated. Zygoma extremely narrowed.
Infraorbital canal "with orifice protected from muscle pressure bv counter-
sinking in a vacuity which extends transversely through rostrum" (Miller &
Gidley). External form Murine or modified for saltatorial life. Cheekteeth
rooted except in Dipodomys, and as a rule less simplified than in Geomyidae.
Subfamily HETEROMYIN.\E
Cheekteeth preserving pattern for a longer time; auditory bullae not specially
inflated; form IXIurine.
Heteromys, Liom\s.
Subfamily DIPODOMYINAE
Cheekteeth losing their pattern earlier; auditory bullae and mastoids con-
siderably to abnormally inflated; external form Murine or saltatorial. (In
Dipotlomys the cheekteeth are rootless and simple.)
Group Perognathi
Perognathus ; Microdipodops .
32 OUTLINE OF CLASSIFICATION HERE ADOPTED
Group Dipodomyes
Dipodomys.
(The arrangement of this family is based on the classification of Wood,
>935-)
Family 14. GEOMYIDAE
Mastoids not inflated. Zygoma robust, the jugal extremely shortened.
Infraorbital foramen with its "orifice protected from muscle pressure by
countersinking in an oblique sulcus" (Miller & Gidley). External form and
cranial characters highly modified for underground life. Cheekteeth in Hying
genera always eyergrowing and always, excepting the premolar, completely
simple.
Thoiiiomys, Geomvs, Piippogeomys, Cratogeom\s, Platygeomys, Orthogeomys,
Heterogeomys, Macrogeomvs, Zygogeomys.
2. Myomorph Series
Infraorbital foramen always clearly enlarged for muscle transmission.
Eighth Superfamily. ANOMALUROIDAE
Infraorbital foramen much enlarged for muscle transmission; masseter
lateralis not extending its line of attachment on to forepart of zygomatic plate,
which remains completely beneath the infraorbital foramen, and is narrow.
Mandible without special peculiarities.
(It may be noted that in this and the next three superfamilies, Pedetoidae,
Ctenodactvloidae, and Dipodoidae, the zygomasseteric structure as regards the
arrangement of the forepart of the skull (infraorbital foramen and zygomatic
plate) is yery similar to that of Hystricoidae and Cayioidae).
Skull not abnormal in the typical subfamily; in the Idiurinae, much modified
by abnormal enlargement of infraorbital foramen, much constricted palate,
thickened incisors, etc. Jugal long. Bullae large, but not abnormally inflated.
Cheekteeth |, rooted, not hypsodont, the pattern typically reduced
heptamerous, the re-entrant folds wide.
Fibula, so far as known, not fused with the tibia (no skeletons of Zenkerella
and Idiiirus available for examination). Digits of hindfocrt fiye. External form
suited to arboreal life; usually a flying-membrane present, attached to sides.
Underside of the tail with scaly outgrowths near the body. Malleus and incus
free (Tullberg).
Family 15. ANOMALURIDAE
With the characters of the Superfamily.
Subfamily ANOMALURINAE
Infraorbital foramen not greatly enlarged. Palate not much narrowed.
Incisors moderate; toothrows not reduced. "Anterior point of masseteric
OUTLINE OF CLASSIFICATION HERE ADOPTED 33
insertion on mandible beneath hinder part of M.i " (Miller & Gidley). (Flying-
membrane present.)
A noiiui/iinis, Anomalurops.
Subfamily lUlLKINAE
Infraorbital foramen extremely enlarged, the zygomatic plate projected
forwards to a point nearly immediately behind the incisors. Palate much
narrowed. "Anterior point of masseteric insertion on mandible in front of
P.4" (Miller & Gidley). Incisors much thickened from before backwards.
Toothrows strongly reduced. (Flying-membrane present or absent.)
Zenkerella; Idiurus.
Ninth Superfamily. PEDETOIDAE
Zygomasseteric structure not essentially different from that of the Anoma-
luroidae (infraorbital foramen extremely enlarged, the zygomatic plate projected
forwards, much as in Idiurinae).
Skull with extremely broad frontals; jugal much thickened, lengthened,
ascending to lachrymal; mastoids extremely inflated.
Cheekteeth J, evergrowing, and near complete simplification in pattern.
Fibula reduced and fully fused with tibia high on the leg. Digits of hind-
foot four. Externally much modified for bipedal saltatorial life. Metatarsal
bones normal, not tending to fuse (compare specialized Dipodidae).
Malleus and incus free (Tullberg).
Family 16. PEDETIDAE
With the characters of the Superfamilv.
Pedetes.
Tenth Superfamily. CTENODACTYLOIDAE
Zygomasseteric structure not essentially different from that of the Anoma-
luroidae in so far as it affects the general shape of the skull. Infraorbital
foramen large but not abnormally so. Mandible with the angular portion
drawn backwards to a degree, but not distorted outwards; coronoid process
absent; a weak short ridge may be developed, reminiscent of the "medialis
ridge" of Cavioidae, though much less developed than in that superfamily.
Skull considerably modified, flattened, the jugal long, the zygoma in two
portions, a horizontal and a vertical (parallel — Dipodinae). Auditory bullae
and mastoids much inflated.
Cheekteeth at full dentition i, the premolars lost in the adult, or J
(Pectinator). The teeth evergrowing, and near complete simplification of
pattern.
Fibula not fused with the tibia. Digits of hindfoot reduced to four.
E.xternal form without special peculiarities; tail fully haired; some of the
digits with stiff bristle-hairs present (parallel — Petromus, Octodontinae,
Chinchillidae).
3 — Lning Kodents — 1
34 OUTLINE OF CLASSIFICATION HERE ADOPTED
Malleus and incus fused (Tullberg).
Radiale and intermedium can be separate (fused in all other Rodents
examined by Tullberg except Bathyergidae).
Family 17. CTENODACTYLIDAE
With the characters of the Superfamily.
Pectinator, Ctenodactyhis, Massoiitiera, Felovia.
Eleventh Superfamily. DIPODOIDAE
Zygomasseteric structure not essentially different from that of Anomaluroi-
dae, the infraorbital foramen always large, sometimes extremely so, the zygomatic
plate always completely beneath it. In primitive genera the infraorbital
foramen is conspicuously wider below than above (compare the primitive
Muroidae, Graphiurus, Deomys). Mandible with angular portion weak,
sometimes perforated; occasionally strongly inflected, and usually with this
formation suggested.
Skull in progressive species characterized by much broadened frontals,
greatlv inflated mastoids and auditory bullae, and specialized zygoma. In
primitive species, the skull more or less Murine in general aspect (except the
infraorbital foramen). Jugal long, usually extending to the lachrymal.
Cheekteeth ^ or I, the extra premolar when present much reduced in
size; the teeth rooted, cuspidate, with re-entrant folds which are fairly well
open as a rule; occasionallv becoming flatcrowned, in which case the re-entrant
folds are narrow (specialized Zapodinae).
Fibula much reduced, fused high on the leg with the tibia. Digits of hind-
foot in primitive species five; or may become reduced in progressive forms to
three functional, or three only. External form much modified for bipedal
saltatorial life, excepting the genus Sicisla.
In specialized genera, the three central metatarsal bones fuse to form a
cannonbone.
Malleus and incus free (Tullberg).
Family 18. DIPODIDAE
With the characters of the Superfamily.
Subfamily SICTSTINAE
External form not modified for saltatorial life. Zygoma simple. Auditory
bullae not inflated. Metatarsal bones normal. Cheekteeth quadritubercular,
cuspidate, not tending to become flatcrowned.
Sicistci.
Subfamily ZAPODINAE
External form modified for saltatorial life (as in all remaining members of the
family). Cheekteeth semi-hypsodont, not quadritubercular, tending in pro-
gressive species to become flatcrowned, in which case the re-entrant folds are
OUTLINE OF CLASSIFICATION HERE ADOPTED 35
narrow, and usually isolate on crown surface as islands. Zygoma simple, un-
specialized; auditory bullae not inflated; metatarsal bones normal.
Eozapus, Zapus, Napaeozapus.
Subfamily CARDIOCRANIINAE
Auditory bullae and mastoids abnormally inflated. Zygoma in two portions,
a horizontal and a vertical, these portions connected by a curvature. Metatarsal
bones normal, not fused. Cheekteeth (apparently) as in Dipodinae. Digits of
hindfoot mav be reduced to three.
Cardiocranius (not seen), Salpingotus.
Subfamily EUCHOREUTINAE
The three central metatarsal bones fused to form cannonbone. Auditory
bullae much inflated. Zygoma unspecialized, simple, like that of Zapodinae.
Cheekteeth cuspidate, with unusually high cusps. Rostrum much lengthened.
Ear much enlarged. M.3 (evidently) vestigial. (Hindfoot with three functional
digits.)
Euchoreutes.
Subfamily DIPODINAE
Three central metatarsal bones fused to form a cannonbone. Auditorv
bullae and mastoids considerably to excessively inflated. Zygoma in two
portions, a horizontal and a vertical, these portions not connected by a curvature.
Cheekteeth with moderate cusps; rostrum weak; ear less enlarged; M.3 not
vestigial. (Hindfoot with three functional digits.)
Group Allactagae
Allactaga, Alactagulus, Pygeretmus.
Group Dipodes
Paradipiis ; Dipiis, Scirtnpoda, Ereniodipus (the last not seen), Jaculus.
Twelfth Superfamily. MUROIDAE
Zygomasseteric structure primitively {Graphiurus, Deomys), nearly as in
Sicistinae, except that the infraorbital foramen is less enlarged, and is not
conspicuouslv wider below than above.
In two hundred other genera examined belonging to the group, the zygomatic
plate is broadened and tilted upwards to a greater or lesser degree; masseter
lateralis extends its line of attachment on to zygomatic plate, and masseter
lateralis superficialis has its anterior head distinct, so far as known, from zvgoma
(see figures in Tullberg). Infraorbital foramen alwavs transmitting muscle,
never extremely enlarged, usually less so than in the preceding families of
Myomorph Rodents; and sometimes (Rhizomyidae) becoming much reduced.
Mandible with angular portion not distorted outwards; in Muscardinidae this
portion is relatively weak, may show signs of inflection, and may be perforated.
Skull usually with constricted frontals; auditory bullae in the majority not
36 OUTLINE OF CLASSIFICATION HERE ADOPTED
much enlarged, but may become so (Gerbillinae); or may be much reduced,
as in Phloeomys, Lophiomys, certain species of Rattiis, and others. Jugal
typically considerably shortened; but long in Muscardinidae, Tachxorvctes, etc.
Cheekteeth \ (Muscardininae, Graphiurinae), f (the greater part of the
superfamily), 'i (genus DcsmoiiiUiscus), or H (Rhvnchomys, some Hydromyine
genera); rooted except in Mxoipalax, Rhomhomys, and the majority of the
Microtinae.
Fibula so tar as known reduced, and fused high on the leg. Digits of hind-
foot fiye \yith one exception (Mahuotlnix). External form as a rule small,
generalized; sometimes highly modified for underground lite (Spala.x, to a
lesser degree Mxospcila.x, Ellohitis, Pronietlieomxs, Notiomxs, etc.); sometimes
highly specialized for aquatic life, cranially as well as externally (Ichtlixomxs
and allies, Hydromys, Crossuinys); sometimes specialized for arboreal life, with
fully opposable hallux {Chiropodomys, Chirumyscus, Hapalomys and others);
in one case, Notomys, apparently fully specialized for bipedal saltatorial life.
Spiny coyering may be moderately deyeloped (Acomxs, some species of Rattiis,
Plalacanthomys). Tail typically naked, scaly; uniformly haired in most
Muscardinidae, CraUromxs, Lophiomys, one species of .Xcotonia, most Ger-
billinae, and others.
Malleus and incus free (Tullberg).
Caecum suppressed in Muscardinidae, so far as known, except in Txph/omys;
becoming much reduced in IcJitlixoinxs (Thomas).
Family 19. MUSCARDINIDAE
Skull without special modification; jugal usually relatiyclv long; bullae large
and to a degree inflated except in Platacanthomyinae.
Cheekteeth \ or jj, primitiyelv with basin-shaped crowns and comer cusps,
becoming flatcrowned in progressiye genera, in which case they become a
series of relatiyely narrow transyerse ridges (which are always traceable, eyen
in primitiye torms), the general dental effect strongly reminiscent of that of
the Sciuridae.
Mandible with angular portion sometimes inflected, and sometimes with
perforation.
Externally slightly specialized for arboreal life; tail fully haired except in
Mxoniimiis (not seen) and Txphlo)nxs. Cardiac portion of stomach with horny
layer absent (Tullberg). (This character not known regarding Platacantho-
myinae; but present, so far as knovyn, in all other members of the superfamily.)
Caecum suppressed, so far as known, excepting in Txphlomys.
Subfamily GRAPHIURINAE
Zygomatic plate remaining completely beneath infraorbital toramen;
masseter lateralis not extending its line of attachment on forepart of zygomatic
plate. (Cheekteeth |).
Graphiiirus.
OUTLINE OF CLASSIFICATION HERE ADOPTED 37
Subfamily MUSCARDININAE
Zygomatic plate tilted strongly upwards; masseter lateralis extending line
of attachment on to forepart of zygomatic plate. Cheekteeth |. Bullae large,
normally. Cheekteeth when flat-crowned with the depressions (between the
ridges) not tending to become isolated with wear. Palate without a series of
foramina (or a single large pair) situated between the toothrows.
Myomimiis (not seen), Eliomys, Dyiomys, GUridus, Glis ; Muscardinus.
Subfamily PLATACANTHOMYINAE
Like the Muscardininae except: premolars suppressed, cheekteeth (flat-
crowned) with the depressions (between the ridges) tending to become isolated
on crown surface with wear; bullae small; palate with a large pair of foramina,
or a series of foramina, between the toothrows. (Zygomatic plate much
narrowed, parallel-Hydromyinae). (A caecum is present in Tvphlomys.)
Platacanthomys, Typhlomys.
Family 20. LOPHIOMYIDAE
Like the Muridae (below, no. 23), but more specialized; skull with temporal
fossae roofed over by bony plates rising from jugal, frontal, and parietal, a
structure not known elsewhere in the Order. Cheekteeth i; pattern as in
cuspidate Cricetinae. External form modified for arboreal life. (Hallux
partly opposable; bullae much reduced.)
Lophiomys.
Family 21. SPALACIDAE
Like the Muridae (below, no. 23), but more specialized; skull and external
form extremely modified for underground life, the eyes suppressed. Infraorbital
foramen relatively large, and zygomatic plate nearly horizontal and below it
(secondarily acquired ? masseter lateralis superficialis with anterior head dis-
tinct, as in Muridae, as figured by Tullberg). Cheekteeth with re-entrant folds
which isolate on crown surface in adult.
Spalax.
Family 22. RHIZOMYIDAE
Like the Muridae (below), but zygomasseteric structure unusual; infra-
orbital foramen extremely reduced, owing to the fact that masseter lateralis
rises to an abnormallv high degree on zygomatic plate (which is \er\ strongly
tilted upwards), extending its line of attachment beside the infraorbital foramen
on its inner side. Externally and cranially more or less modified for fossorial
life; cheekteeth with re-entrant folds isolating as islands on crown surface in
adult. (Infraorbital foramen not V-shaped below.)
Rhizomys, Cannomys.
(End of First Volume)
38 OUTLINE OF CLASSIFICATION HERE ADOPTED
Family 23. MURIDAE
Infraorbital foramen typically specialized into a wider upper portion for
muscle transmission and a narrower lower one for nerve transmission, its lower
border very generally V-shaped (not nearly straight as it is in Rhizomyidae).
Jugal normally strongly reduced (except Tachyoryctes and some genera from
Madagascar). Cheekteeth laminate, cuspidate, heptamerous or prismatic, but
never reminiscent of those of Sciuridae, i.e. never agreeing in pattern with
those of Muscardinidae. External form various, but when subfossorial, eyes
retained, and zygomatic plate not specially narrowed nor turned downwards
(compare Spalacidae). Temporal fossae never roofed in by bony plates (com-
pare Lophiomvidae). Masseter muscle, so far as known, not rising beside
infraorbital foramen on its inner side (compare Rhizomyidae).
The order in which the subfamilies are listed here is provisional.
Only valid genera which have been actually examined are included in the
present list.
Subfamily DEOMYINAE
Zygomatic plate remaining completely beneath the infraorbital foramen.
(Pattern of cheekteeth as in Dendromyinae).
Deoinxs.
Subfamily MURINAE
Zygomatic plate (as in all remaining subfamilies) broadened and tilted
upwards to a greater or lesser degree.
Cheekteeth laminate or cuspidate; when laminate, the laminae tightly pressed
together; when cuspidate, the cusps of the upper molars arranged in three
longitudinal rows.
Group Kliuri
Eliunts.
Group Anisomyes
Anisoiiixs.
Group Alures
Hapalomvs, Ponoriomvs, Leiionivs, CJiiriipadoinys. V(iii<iclc!ina, Microinys,
Apodemus, T/iaiiiiioiiivs, Graniiiionivs ; Caipomys. Batuiiiys, Pitluiheir,
Ciotennnvs, Hvoinvs, Malluiiivs, Coii/liinis, Zyzomys, Laoniys, Mcscinhriomys ;
Oeiwmvs, Dasvmvs, Mvlomvs, Arricanthis, Leinniscomys, Rliabdomys, Prlomys,
HyhowYS, Hadroiiivs, Millardia, Pyromys, Daciioniys, Eropcphis, Steiio-
ceplialcmys, Acthomys, Thallomys, Kattiis, Gyuiiiys, Lcporillus, Pseiidumys,
Apomys, Melomys, Uromys, Cuclumys, MaUuomys, Hiieromys, Zelotumys,
Chiromysciis, Leggadwa, Miis, Miiriculus, Hylenomys, Mycteroinys, Colomys,
Nesoromys, Macruromvs, Crunomys ; Notomys : Mastacomys ; Golunda ;
Acomxs, Lhanonns ; Lopluiroiiiys ; EchiatJirix ; Bmidlcota, Nesokia ; Beainys,
Saccostomus ; Cricetomys ; Pliloeomys.
OUTLINE OF CLASSIFICATION HERE ADOPTED 39
Subfamily RHYNCHOMYINAE
Like the Murinae, but incisors and cheekteeth (f ) so reduced as to appear
almost functionless.
Rhynchomys.
Subfamily HYDROMYINAE
Like the Murinae, but cheekteeth (often |) with a pattern characterized
by a series of basin-shaped lobes (evidently the outer row of cusps of Murinae
have become suppressed).
Zygomatic plate much narrowed, though strongly tilted upwards. Infra-
orbital foramen may be more enlarged than is usual. M.3 when present
vestigial.
Xeromys, Leptomys, Chrotomys, Celaenomys ; Parahydromys, Crossomys,
Hydromys.
Subfamily DENDROMYINAE
Like the Murinae, but cheekteeth with the inner row of cusps of upper
molars becoming suppressed; !\L tl vestigial.
Dendromiis, Steatotnys ; Malacothrix ; Prionomys ; Petromyscus.
Subfamily OTOMYINAE
Cheekteeth with pattern of a series of transverse plates; in the upper series
M.3 becoming the dominant tooth; cheekteeth hypsodont. Bullae may become
much inflated.
Otomys, Parotomvs.
Subfamily CRICETINAE
Cheekteeth laminate, cuspidate, prismatic or heptamerous; when cuspidate
the cusps arranged in two longitudinal rows, when laminate the laminae separ-
ated by wide folds; when prismatic, cheekteeth rooted, and skull not much
modified by ridges for muscle attachment (compare Microtinae).
Oryzomys, Neacomys, Megalomys, Nectomys, Thomasomys, Rhipidomys,
Phaenomys, Chilomys, Nyctomys, Tylomys, Ototylomys, Rhagomys, Nesomys,
Reithrodontomvs, Peromysciis, Baionivs, Calomvsciis, Onvc/ioinvs, Akodon,
Zvgodontomvs, Microxus, Leiioxus, Oxvmvcteriis, Hlarinomvs, Xulioinvs, Scap-
teromvs, Sculinomys, Cricetulus, Pliodapus, Cricetiis, Mesucricetus, Mvs/romys,
Hesperomvs, Eligmodoiilia, Graomvs, Phvllotis, Cliinchillula, Ireiiomvs, Rcitli-
rodoii, F.uneomvs, Chelemvscus, Xeotomys, Sigmomys, Sigmodori. Ilulochiltis,
Andinomys, Neulomodun, Aeotoma, Hodumys, Nelsonia ; Hypogeomys ; Rheomys,
Ichthyomys, Anotomys.
Subfamily GYMNUROMYINAE
Cheekteeth flatcrowned, laminate, the laminae excessively tightly packed
together, the pattern a series of isolated folds, these folds line-like and extremely
narrowed. M.3 slightly larger than M.2, and M.2 slightly larger than M.i.
Gymnuromys.
40 OUTLINE OF CLASSIFICATION HERE ADOPTED
Subfamily GERBILLINAE
Skull specialized hy inflation of auditory bullae and braincase, and narrowing
of rostrum (" saltatorial tvpc "). Cheekteeth tending to become a series of trans-
verse plates, these separated bv wide folds, in progressive genera; in primitive
forms, the teeth are cuspidate at birth, the cusps arranged in two longitudinal
rows, in the upper molars. I\1.3 usuallv strongly reduced. (The cheekteeth
are evergrowing in the genus Rhombomys.) External form modified tor
terrestrial plains or desert life, perhaps saltatorial in some cases; tail usually fully
haired; limbs often lengthened to a certain degree.
Microdillus, GerhiUiis, Tiiten'l/iis, Tatera; Desmodil/iis, DeswodilUscus, PacJiyu-
romxs ; AminodiUiis ; Merioncs, Biacliiones, Psanwiomvs ; Rhunibomys.
Subfamily TACHYORYCTINAE
Cheekteeth moderately or strongly hypsodont, rooted, the pattern consisting
of thick curved parallel ridges of enamel extending across crown surface.
External form generalized or fossorial.
Group Brachyuromyes
Brachxuromxs.
Group Tachvoryctae
(Infraorbital foramen V-shaped below, compare Rhizomvidae.)
Taclixorxctes.
Subfamily MYOSPALACIXAE
Cheekteeth evergrowing, the pattern prismatic. Infraorbital foramen rela-
tixely large, zygomatic plate not strongly tilted upwards. Skull and external
form modified for underground life, the lambdoid crest slanting forwards about
to level of posterior zvgomatic root (parallel-Spalacidae).
Mx(}spalax.
Subfamily MICROTINAE
Cheekteeth prismatic in pattern, frequently but not always evergrowing.
Infraorbital foramen small, narrowed; zygomatic plate strongly tilted upwards.
Skull much modified bv ridges for muscle attachment (tendency to develop
median interorbital crest, stjuamosal crests, etc.). Lambdoid crest not slanted
forwards to level of posterior zygomatic root.
Group Brachvtarsoniycs
Briuhxtiiisoinvs.
Group Lemnii
Dicrostonyx ; Svnaptomvs, Myopiis, Lemmus.
Group Microti
Clet/nionoinys, Asclnzoinvs, Eutlicnomxs, AntcUomxs, Altico/a, Uypiiacnus ;
OUTLINE OF CLASSIFICATION HERE ADOPTED 41
Dolomys ; P/teniicomvs ; Anicola, Pitvmvs, Blanfordimvs, Phaiomvs, Seodon,
Pedomys, Orthriomys, Ilerpetumys, Proedromys, Microtus, Lasiopodomys ;
Lagurus ; OndatrOy Neofiber ; Prometheomys, Ellobius.
The "Family Nesomyidae " or subfamily Nesomyinae of some authors in
which all the Rats of Madagascar are included is here regarded as not definable.
Nor can all these Rats be referred to the Cricetinae, as is often done, for it
seems clear that for the most part they are not closely related to each other.
Excellent figures of zygomasseteric structure of all the leading families of
Rodentia arc published by Tullberg, Nova Acta Reg. Soc. Sci. Upsaliensis, 3,
XVIII. 1900 (1899).
Before dealing in detail with tlie various families, it may be mentioned that
in this work, fifty-two families and subfamilies are recognized in the Order,
and that about three hundred and thirty-six genera have been examined,
included in the keys, and retained as valid.
ZYGOMASSETERIC STRUCTURE
Without entering into any detailed account of the variations of the arrange-
ment of the jaw-muscles of the Rodentia, it is necessary to note certain characters
by which these muscles modify the skull, and with which it is usually possible
at once to identify the "superfamily" position of any living Rodent.
There are three parts of the skull which become greatly affected in the
various families, namely the infraorbital foramen, the zygomatic plate, and the
formation of the mandible.
The formation of the mandible has usually been used by most authors to
divide the " Hystricomorph " series from the remainder of the others; Tullberg
made this his major division of the Order, and divided it into "Hystricognathi"
(including Bathyergidae) for forms in which the mandible has the angular
process lifted outwards by the specialized limb of masseter lateralis superficialis,
and the " Sciurognathi," in which this does not take place.
The angular portion of the mandible is as just described, to a greater or
lesser degree, but (with rare exceptions) strongly and clearly marked, in the
families Bathyergidae, Hvstricidae, Erethizontidae, Echimyidae, Dinomyidae,
Cuniculidae, Dasvproctidae, and Chinchillidae. It is at its weakest in the
C'hinchillidae, and in this family there is some approximation towards that
structure, next to be described, found in Caviidae. It is also relatively weakly
developed in more primitive Hvstricidae, as Atlicruriis; but in most of the
genera comprising the above-mentioned families (including Petiomiis which I
refer to the Echimyidae, but which has frequently been lumped with the
Ctenodactylidae), it is strongly and clearly marked, and reaches its highest
degree of strength in such forms as Tlirvoiiomvs, Mvocastor, Capromys and
certain Echimyidae, and in the Bathyergidae.
The angular portion of the mandible is not lifted outwards by the lateralis
muscle in anv other family or genus of Rodent, so far as I have seen.
In the Caviidae (Ciivici, Galea, Keiodon, Cavie/la, Dolichotis, Hydrochoents),
the mandible is not by anv means tvpicallv Hystricoid, though these Rodents
have universally been placed in the Hystricoid series; here it is according to
Miller & Gidley the medialis portion of the masseter which influences the
jaw; a very deep ridge is developed along the jaw slightly below the level of
the toothrows; though this structure is suggested in Ctenodactylidae and more
so in Chinchillidae, the Caviidae have as far as I have seen an entirely unique
formation of the lower jaw in degree of development. VVaterhouse suggested
that if the lower incisors of the Caviidae were longer and continued further
backwards the mandible would be typically Hystricoid, and included them in
his Hystricoid series. Be that as it may, unfortunately the mandible is not so,
and therefore Caviidae cannot be looked upon as typical Hystricoid Rodents
to-day, whatever they may have been derived from.
In other families of Rodentia, the mandible may have the angular portion
42
ZYGOMASSETERIC STRUCTURE 43
pulled inwards instead of lifted outwards, or may be comparatively without
peculiarity.
The pulling inwards is developed to a most abnormal degree in the Aplo-
dontiidae. It is also to be found in certain Sciuridae, certain Dipodidae; it is
evidently nearly as in A[>loduntia in the genus Cynomys (Sciuridae); and in the
genus Cardiocranius (Dipodidae; not seen).
The mandible of the Ctenodactylidae, so often placed in the Hystricoid
series, is abnormal, but not in the least like the Hystricoid type. The coronoid
is suppressed; the angular process drawn backwards to a degree; and a faint
medialis-ridge, reminiscent of that of the Caviidae may be traced.
I can call to mind no special peculiarities with regard to the mandible of
the vast number of genera and species I have examined in the Muridae. Rarely
the coronoid is suppressed. In the Aluscardinidae, and in certain Dipodidae,
the angular portion mav be perforated. In the former family sometimes traces
of the pulling inwards of the angular, so highly developed in Aplodontia, will
be seen. The mandible mav be noted as weak in the Heteromyidae.
In some genera with the "non-Hystricine" mandible, the lower incisor
extends so far backwards that it forms a conspicuous process between the
condylar and angular processes; examples are Spalax, Geomyidae, Rhizomyidae,
Sesokia, etc. This never occurs in genera with Hystricine type of mandible.
Pedetes, sometimes placed in the Hystricoid series (as by Thomas), has certainly
not a Hystricine type of mandible, with its reduced relatively small angular
process.
In mandible structure, therefore, Rodents divide very broadly speaking into
three classes :
Angular process lifted up and distorted outwards by specialized limb of masseter
lateralis superficialis;
Bathyergidae, Ilystricidae, Erethizontidae, Echimyidae, Dinomy-
idae, Cuniculidae, Dasyproctidac, Chinchillidae.
Angular process never lifted up as above described.
Lower jaw deeply modified by conspicuous ridge extending below level of
toothrows on outer side, for attachment of masseter medialis:
Caviidae.
Lower jaw without extreme modifications, except in certain cases by root of
lower incisor; or by strong inflection of angular process (e.xtreme
only in Aplodontiidae):
.\plodontiidae, Sciuridae, Geomyidae, Heteromyidae, Castoridae,
Dipodidae, Ctenodactylidae, .Anomaluridae, Pedetidae, Muscar-
dinidae, Spalacidae, Lophiomyidae, Rhizomyidae, .Muridae.
The infraorbital foramen is enlarged to transmit the masseter muscle in a
very large number of Rodents. Degree of enlargement, and shape and size of
this foramen varies exceedinglv.
Even in those forms which arc regarded here as not transmitting muscle,
in two families, Sciuridae and particularly Bathyergidae, is certain variation
44 ZYGOMASSETERIC STRUCTURE
found. In Protoxeriis and Tamias (Sciuridae), the infraorbital foramen is more
enlarged than in the other Squirrels, and probably may transmit a small strand
of the muscle. In these cases however it is not so far as I can judge anything
like so enlarged and clear as in any Rodent which is regarded here as a form
with muscle transmission of this foramen present. In the Bathyergidae, certain
species of the genus Crvptoiiivs appear to be starting to transmit muscle through
the infraorbital foramen ; it may rarelv, as in C. mcUandi, even be as much en-
larged as in the much reduced type found in the Rhizomyidae. This is evidently
a variable character, and in Cryptomys the foramen may even be more enlarged
on one side of the skull than on the other, in individual cases. In the Aplo-
dontiidae, most authors state that the canal does not transmit muscle; however
in those examined it is on the large side for this section of the Order.
The infraorbital foramen does not transmit muscle in Geomyidae, Hetero-
mvidae (excessively reduced in these two families), Castoridae, .Sciuridae with
the above noted exceptions, Aplodontiidae and Bathyergidae, with the above
noted exceptions.
In all other Rodents it is clearly enlarged to do so. There are then broadly
speaking two types of infraorbital foramen structure to be discussed, with the
one exception of the Rhizomvidae. In Hystricidae, Erethizontidae, Echimyidae,
Dinomyidae, Dasyproctidae, Chinchillidae, Caviidae, Pedetidae, Anomaluridae,
Ctenodactylidae and Dipodidae, it is round, completely above the zygomatic
plate, and normally extremely enlarged. This enlargement reaches its greatest
development probably in the Pedetidae, and in the Idiurine subfamily of
Anomaluridae; and in certain sections of the Dipodidae. In the Cuniculidae,
the infraorbital foramen is secondarily reduced by the growth of the enormous
cheekfilates. In two genera of Rodents which are here referred to the Muroid
superfamilv, Deoiins (Aluridae), and Gniphiiinis (Muscardinidae), the infra-
orbital foramen, though not abnormallv enlarged, is completelv above the
zygomatic plate.
In the remainder of the Order, the infraorbital foramen, though sometimes
varying in actual size of enlargement, is never abnormally enlarged; in vast
sections of the family Muridae, it is specialized into a wider portion above for
muscle transmission, and a narrower lower one for nerve transmission. In the
Subfamily Microtinae, it has become, correlated probably with the increase in
general strength of jaw-muscles in this group, much reduced. It is abnormally
reduced in the Rhizomyidae {K/ihoiiivs and Cannom\s)\ in this group, the
zvgomatic plate is strongly broadened and tilted upwards, and the foramen
becomes reduced to a small aperture situated at the top of this plate; the
masseter muscle rises up inside of it, a condition according to Tullberg not
known elsewhere in the Order.
It mav be noted that Winge puts forward the theorv that in all Rodentia
living, Aplodotitia excepted, the infraorbital foramen has transmitted muscle,
and has become secondarily closed in the Geomyidae, Sciuridae, Castoridae,
Bathyergidae, Heteromyidae. Without wishing to enter into a discussion on
matters such as these, it appears to me to be singularly unlikely that, having
taken such a large step forward in evolution as the enlargement of this canal
ZYGOMASSETERIC STRUCTURE 45
for muscle transmission (as it seems an unusual character among Mammalia
to say the least), these families should go even further in evolution and, so to
speak, develop covering over this canal so that it does not transmit again. There
is not a wide difference in the arrangement of the zygomatic plate between
Aplodontiu and a primitive Sciurine such as Belomys; it would seem so much
more likely that the Sciurine arrangement of jaw-muscles was developed from
a type not widely distinct from Aplodontia as regards arrangement of infraorbital
foramen and zygomatic plate ; far more likely than that the infraorbital of Belomys
is secondarily closed to muscle transmission.
In the Spalacidae (Spalax alone), the infraorbital foramen is larger than
usual for a Muroid Rodent and the zygomatic plate, though to a degree
broadened, appears to be nearly flattened to a horizontal position. This how-
ever may well have been brought about by the fossorial habits of this animal.
Summarizing: the infraorbital foramen does not, or scarcely transmits
muscle in Sciuridae, Geomyidae, lleteromyidae, Castoridae, Bathyergidae,
Aplodontiidae.
It is enlarged, and usuallv very much enlarged for muscle transmission in
Hystricidae, Erethizontidae, Echimyidae, Dinomyidae, Cuniculidae
(see note above), Dasyproctidae, Chinchillidae, Caviidae, Ctenodacty-
lidae, Anomaluridae, Pedetidae, Dipodidae.
It is enlarged, but very rarely much enlarged for muscle transmission in
Muridae, Lophiomyidae, Spalacidae, Muscardinidae, and Rhizomyidae
(see note above).
The zygomatic plate is less variable in structure, broadlv speaking, than
either the infraorbital foramen or the mandible. Among the Rodents it is found
in two conditions only. It is narrow, usually very narrow, and strictly hori-
zontal, remaining completely beneath the infraorbital foramen, in Aplo-
dontiidae, Bathyergidae, Dipodidae, Anomaluridae, Ctenodactvlidae, Pedetidae,
Hystricidae, Erethizontidae, Echimyidae, Dinomvidae, Dasyproctidae, Chin-
chillidae, Caviidae, and in the genus Graphiurus (Muscardinidae), and Deomys
(Muridae).
In the Cuniculidae it is much distorted bv the growth of the bony cheek-
plates.
In other Rodentia, to a greater or lesser degree, it is broadened and tilted
upwards. In these, according to Miller & Gidley, and supported by Tullberg's
figures, masseter lateralis superficialis is distinct from the zygoma, "not attached
to any part of zygoma except occasionally to a point at extreme base of zygo-
matic plate."
In the Sciuridae, Castoridae, Geomyidae and Heteromyidae, in which the
infraorbital foramen does not transmit muscle, the zygomatic plate is very
generally strongly broadened and tilted upwards, the only exceptions being
found among the Sciuridae; such as Tamias, and most members of the
Pteromvs group except Pteromys. In these families, the lateralis muscle
rises to the superior border of rostrum and excludes masseter medialis from so
doing. In the Muridae, so far as known, except Deomvs, the Muscardinidae,
46 ZYGOMASSETERIC STRUCTURE
except Grapliiurus, the Lophionividae, the Spalacidae, and the Rhizomyidae,
the zygomatic plate is broadened and tilted upwards to a certain degree, hut
masseter mcdialis is transmitted through the infraorbital foramen so that it is
not excluded from the superior border of the rostrum, and masseter lateralis
as a rule does not extend so high on the forepart of the skull. The zygomatic
plate in these families only approaches the Sciurine type of specialization as
regards broadening in the Rhizomyidae. The degree of broadening, narrowing,
and tilting upwards varies extremely through the Muridae, as might be expected
in such a vast group. In Hydromyinae, though tilted up strongly, it is narrow.
In such genera as Oxymycterus, and Lupjiuromvs, it is very little tilted upwards;
but only in Deomys of the vast number examined does it appear to me to be
absolutely indistinguishable from the Dipodoid type as defined by Miller &
Gidley.
Notwithstanding this, although Tullberg's figures show clearly that there is a
wide distinction between Glis and Grapliiurus in the iVIuscardinidae, and between
Deomys and Oxymycterus in the JMuridae, as regards zvgomasseteric structure
of the forepart of the skull, I am not persuaded of the desirability of transferring
Grapliiurus to a separate superfamily from Glis, as was done by Miller & Gidley,
nor Deumys to a separate superfamily from the remainder of the Muridae,
although it must be admitted that to identify the superfamily relationships of
Deomys (here considered a Muroid), from Sicista (a primitive Dipodoid), is not
possible on this character alone. It would seem however that the close resem-
blance in all other main characters between Grapliiurus and sav Eliomvs, and
between Deomys and sav Deiidromus indicate that the Murine type of zygomatic
plate and arrangement of jaw-muscles has been derived from the Dipodoid
type, or vice- versa.
The zygomatic plate therefore in the Order is narrow, and completely
beneath infraorbital foramen, showing no signs of becoming broadened
and tilted upwards, in Hystricidae, Erethizontidae, Fxhimyidae,
Dinomvidae, Dasvproctidae, Caviidae, Pedetidae, Ctenodactylidae,
Anomaluridae, Dipodidae, Aplodontiidae, Hathvergidae, Chinchillidae,
Muscardinidae, part, subfamily Graphiurinae, and Muridae, part, sub-
family Deomyinae only.
It is much modified by growth of cheekplate in Cuniculidae, but presumably
possessed the above character originally as in the rest ot the Ilvstri-
coidae.
It is broadened and tilted upwards to a greater or lesser degree in Sciuridae,
Castoridae, Geomyidae, Heteromyidae, Lophiomyidae, Rhizomyidae,
Spalacidae (see note above, p. 45), Muscardinidae. part, except
Graphiurinae, and Muridae, part, all except Deomvinae.
The presumed relationship between Deomys and the Dendromyinae, and
between Grapliiurus and the remainder of the Muscardinidae indicate that it is
not wise to base superfamilv grouping on zygomasseteric structure alone, as
was done by Miller & Gidlev.
DISTRIBUTION
The Rodentia is the only Order of non-Marsupial land mammals inhabiting
Australia. The one family of Rodents, the Muridae, must have either got there
early from South-east Asia, which is the view currently held, or evolved there,
which is the view suggested in the present paper. It is curious that if the
Muridae alone came from South-east Asia, some members at least of the
families Tupaiidae, Soricidae, Erinaceidae, Galeopteridae, Viverridae, Musteli-
dae, Tarsiidac, Cercopithecidae, Tragulidae, Cervidae, Sciuridae, Hystricidae
and Manidae, to quote only some of the families widely or at least comfortably
distributed throughout the Indo-Malayan islands to the north-west of New
Guinea, did not do so. It is remarkable to say the least if not one genus of this
vast assemblage entered the Australasian region, and yet such a large number
of Muridae did so. For in New Guinea and Australia, and immediately adjacent
islands such as Ceram, there are two hundred and forty-five named forms of
Muridae, belonging to twenty--five genera and two subfamilies. It is to my
mind as likely that a large section of the Muridae evolved in Australia and came
into Asia via .some islands as Celebes and the Philippines which may have for a
time been separated from Asia and joined Australia, but later separated from
Australia and joined Asia, than that all these Australian types came from Asia
unaccompanied bv anv other genus of non-murine mammal. (The presence
of the genus Sus in New Guinea is usually held to be due to introduction.)
'I'his view suggests that the Muridae are among the most archaic of mammals,
which appears on account of their universal distribution to be likely.
The main Australasian genera of Aluridae are Rattus (many species totallv
distinct from the "ship-rats" rattus and norvegicus, and including one group,
concolor which ranges to some of the Pacific Islands), Uromys (doubtfully
distinguishable from Rattus), and the isolated and distinct genera Zysornvs,
Mesembriomvs, Notomvs, Conilunis, Leporillus, and .Mastacomvs (Australian or
Tasmanian), and Mallotnys, Hvomys, Pogonnmys, Macruromys, Xesoromys and
Anisoinys (New Guinea or Ceram). Leggadina (Australian), appears to represent
a wild ally of the cosmopolitan genus .l/iw, which is I think not indigenous to
the area under consideration. .-Ml these belong to the subfamily Murinae; the
subfamily Hvdromvinae, which is probably derived from Murinae, and closely
allied to it, has half a dozen representatives in the area, as Hydromys (Australia
and New Guinea), and the more restricted Xeromys (Queensland), Crossomys,
Parahydromys, Leptomys, etc. (New Guinea), most of which are little known
and rare.
The Indo-Malayan region presents few families, but great reduplication
of species within the larger genera. Only Sciuridae, Hystricidae, Rhizomyidae,
Muscardinidae (Malabar and South China), and Muridae have penetrated the
area, and only Sciuridae, Hystricidae and Muridae to any great extent. Roughly
twelve hundred forms are named from the area, about half of which arc Muridae.
47
48 DISTRIBUTION
In this family, of the typical subfamily, the genus Rat/us (largest genus in named
forms in the Order), has its headquarters in the present region, with oyer three
hundred and fifty named forms ranging over the whole area, and containing
in the area about twenty specific groups, eight of which range through the
greater part of the area except Peninsular India, and in some cases Celebes, two
of \vhich are peculiar to Peninsular India, and several of which are confined
either to Celebes or the Philippines. The genera Bandicota and Cliiropodomvs,
and to a lesser degree Mus, range through most of the area except that Chiropo-
domys is not known from Peninsular India nor Celebes, and Bandicota does not
range further east than Java. Apart from these the Murine genera of the Malay
Islands are rather different from those of the mainland. In the Philippines,
highly specialized genera such as Phloeomvs, Crateromvs, Carpomvs and Cru-
tiomys occur, and are not known outside the islands; they may be allied to
certain Australasian types, as the New Guinea Mal/omvs, etc. The highly
aberrant genus Echiothrix is confined to Celebes. A tew other rather unim-
portant genera are named from Sumatra, Java, Borneo, closely allied either to
Rattus or Mus, excepting Pithecheiy (Sumatra, Malacca).
In the eastern portion of the mainland which constitutes this area (Burma,
Siam, Indo-China region), a few genera as the isolated Hapalomys, and types
such as Dacnomvs and Hadromvs are confined. Vaiide/eiina, wholly Indo-
malayan, ranges into the area from Peninsular India. In South China, the
Palaearctic genera Miiioiiivs and Apudcmtis occur. In this area, and Siam, the
genus Alus appears to end its natural Eastern Range (except perhaps for its
presence in the Philippines).
Peninsular India appears to have types rather different from those of the
eastern Indomalayan ; among these may be mentioned Goliiirda and Mil/ardia,
which range more or less through the area, and in the north occurs a species
of Acomys (African and Palaearctic chiefly), and Nesokia.
A distinctly Australasian element is seen in the Philippines in the presence
of the Hydromvinae (Chrotvmvs, Celaenomys). From the same island comes
R/ivncIioinvs, which is here regarded as type of a subfamily the Rhynchomyinae.
The Muridae of the Malay Islands, other than the Philippines, all belong to
the typical subfamily. In the northern part of the mainland area, a few INIicro-
tinae, as Eothenomvs (Southern China, Burma), Neoduii (Sikkim), and some
others have their southernmost range limit in the Old World. In Peninsular
India, the Gerbillinae are represented by Tatera which occurs throughout the
area; the subfamily is not known from the remainder of the Indomalayan.
The family Rhizomvidae {Rliisoinvs, Cainiomvs) is more or less confined to
the area, ranging out of it only in parts of Szechuan, the group extends through
South China and from Nepal south through Siam to Malacca and Sumatra.
The Muscardinidae is represented by two rare genera which form a well-
marked subfamily (Platacanthomyinae), and are confined to the Malabar coast of
India (Platacantliomxs), and to South China (7\'phluiiivs). The Hystricidae very
probably evolved in the present area since all the most primitive known types
seem to be grouped in it. Two, Trichvs and I'ht'curus, are confined to the
islands (Sumatra, Borneo, and in the case of Thectirtis, the Philippines) (Trichvs
DISTRIBUTION 49
reaches Malacca). The more widelv ranging genera Atherurus and Hvstrix
occur throughout much of the area; Hvstrix seems absent only from the Philip-
pines and Celebes; Atherurus ranges from Sumatra at least, north to Assam
and South China. The species of Hystrix in the area are with the exception
of the form found in Peninsular India, which also ranges over much of Palae-
arctic South-west Asia, of the more primitive type, at any rate as regards
development of external covering of quills and spines.
The Sciuridae present a great number of forms in the area, and a high
degree of specialization. In Peninsular India, only two genera of non-flying
squirrels occur, Funambulus (confined to the area), and Ratufa, which ranges
over the whole region east to Borneo, but evidently not much in .South China,
though known from the island of Hainan. In Nepal and Burma, many more
genera occur; Callosciurus (not very clearly distinguishable from the Holarctic
genus Sciurus), heading the list with about three hundred named forms. Mar-
mota ranges into the area from the Palaearctic, in Nepal and Yunnan. Dremomys
and Menetes may be mentioned as types typical of the eastern part of the main-
land; the former ranges to Malacca and Borneo. When Malacca is reached,
many new forms start their ranges, some of which are highly specialized.
Lariscus and Rliiuosciwus, both ranging to Borneo, are among the more im-
portant. The Pygmy Squirrels of the genus \' amiosciurus go through the whole
of the larger Malay Islands, from Sumatra to the Philippines, except Celebes,
where they are represented evidently by a species, murinus, which agrees more
in characters with the allied genus Sciurillus. Other peculiar types are Rheithro-
sciurus and Glyphutes, both of Borneo. None of the above-mentioned are known
outside the Indomalayan region, except that Callosciurus has a few forms
ranging into Palaearctic China. And it seems that the further south one goes
the more highly specialized or aberrant become the distinct genera, though the
more normal types as Callosciurus and Ratufa go through much of the area, the
former even including Celebes and the Philippines. This is one of the few
regions in the world where the named forms of Sciuridae actually exceeds the
number of named forms of Murinae, for in addition to the above-mentioned,
the area seems to be the headquarters of the Flying-squirrels; Petaurista, the
giant Flying-squirrels, and smaller forms as Hylopetes and Petinomys, range
more or less throughout the whole area except that Petaurista does not occur
east of Borneo, Hylopetes does not enter Peninsular India, and neither Hylopetes
nor Petinomvs appear to go verv far into South China. Belomys is an important
genus confined to the north-eastern part of the area (Sikkim, Tongking, For-
mosa, etc.).
To the Malay Islands, some very distinct generic types are restricted, the
most noteworthy being lomys. It will be seen that, as indicated above, only
three families of Rodents have gained a real footing in this region, which is a
very different state from that present in most of the other large areas of the
World.
Palaearctic Rodents. The Palaearctic as here understood contains all
land in the Old World lying north of the Yangtsekiang River, the 30" line of
latitude through northern India (i.e. including Kashmir), and broadly speaking
4 — Living Kodents — I
so DISTRIBUTION
the remainder of South-west Asia and the coastal regions of Africa which he
north of this hne, or just south of it (as South Persia). Arabia should probably
be regarded as forming part of the African rather than the Palaearctic region.
From this area roughly eleven hundred forms are named. The Muridae is
verv much the dominant family here in that seven hundred and fifty forms
approximately belong to it. Six other families have a wide or moderate range
in the area, and two, Ctenodactylidae (coastal regions of North Africa, Ctefio-
dactvliis, Massoutiera) and Rhizomyidae (Szechuan, Rliizoniys), just touch it.
The Microtinae is here the dominant suhfamilv of Muridae; the genus
Microtus, which occurs almost throughout the whole region except most of
North Africa, being the sole Rodent genus with more than a hundred forms
named from the area alone. Other Microtine genera with a wide range are
Clethrionomys, and Arvicola, the former like Microtus extending across into
North America. Ellobim and Prometheomys, the two subfossorial \'oles, are
restricted to the area. The Lemmings, Dicrostonyx (Arctic regions), Lemimis
and Myopiis range across the northern portion. The two former also cross into
North America. I'he most interesting of the rather numerous remaining
genera in the area are Lagiinis, Alticola, Do/oiiiys, Pitvmys and Blanfordimys.
Pitviins has a wide range in Continental Europe, but is not met with further
east of the Caucasus until it turns up again in Eastern North America, though
several forms as Neodon occurring to the East in the Palaearctic are closely allied
to it. Lagurus is also known from America. Dolomvs and Blanfordimvs are
rare and local (Montenegro region, and Afghanistan region respectively).
The Murinae are well represented, but by only a very few genera, at any rate
compared with the huge numbers of genera to be found in any of the tropical
portions of the Old World. Indeed, only five have a real range in the area.
Apodemus is probably naturally the most widely distributed, as well as one of
the most primitive members of the group, and appears to extend its range even
to Iceland. Mus and Rattiis are now cosmopolitan in the Palaearctic owing to
artificial human distribution, but both probably have a naturally wide range in
the area, especially the former. Micromys ranges intermittently from England
to Japan ; and Xesokia is common in the more southern portions of the .Asiatic
part of the area (Syria (into Egypt), across Persia and Russian Turkestan to
Kashmir and Sinkiang). Of the rest, some four genera touch the coastal part
of Africa, one of which, Acomvs, is known from Crete and Syria, and three genera
range north from India into the Kashmir region, the most important of which is
Goliinda. The Subfamily Gerbillinae has a wide range in the Palaearctic east
of western Europe; Meriones is the main genus, having more named forms from
the area than any other Palaearctic Rodent except Microtus and Apodemus;
Rlwmbomxs, the most highlv specialized member of the subfamily, is from the
Palaearctic only; but apart from these no member of the group ranges as far
north as Siberia, being mostly confined to the Syrian-Persian region (as Tatera),
or North Africa (Gerbillus. Psammomys, etc., both of which range into Syria).
The peculiar " Fat-tailed (ierbils" of the genus Pnchyuromys seem more or less
restricted to the Palaearctic portion of Africa.
The subfamily Cricetinae has a fairly wide range in the area, though only
DISTRIBUTION 5>
five genera are met with, the group being primarily American ; Calomyscus,
surely a very near relative of the American Perumyscus, is restricted to Persia,
Russian Turkestan, and Baluchistan; the more typical Hamsters, which seem
not to have very near allies in North America, have a wider range; Cricetus
occurring from Central Europe as far west as Belgium, east to Central Siberia;
Cricetiihis covering a very large part of China, as well as Greece, South Russia,
Syria, S.W. Siberia, and Kashmir; the other genera occurring in the area
being Mcsocricetiis and Phodopus. In addition to the four great subfamilies of
Muridae being well represented as indicated above, there is a very interesting
subfamily confined apparently to Palaearctic China and adjacent parts of Siberia
only, the Myospalacinae, with one genus, Myospalax. The family Spalacidae,
which is here restricted to the genus Spalax alone, is purely Palaearctic, ranging
round the eastern end of the Mediterranean Sea from Hungary and the Balkan
States to Egypt, and occurring in South Russia. The family Muscardinidae,
represented by the typical subfamilv, is more or less western in general dis-
tribution, though represented in Japan. The four better known genera, Eliomys,
Dyromvs, Glis and Muscardinus all appear to meet in Central Europe, so far as
their range is concerned. Dvi'omys goes east to Tianshan and Zungaria, but
not west of Central Europe; Glis ranges to Spain and the Atlantic, also east to
the Caucasus and Turkestan; E/iomvs does not range east of European Russia,
but occurs again in Sinai and North Africa, as well as the Iberian Peninsula;
Muscardinus is not known from Spain nor east of European Russia, but ranges
naturally in England and in Scandinavia, which none of the others reach except
by introduction. The family Dipodidae has its headquarters in the Palaearctic.
Of the more primitive groups, the Sicistinae (Sicista) has the widest range,
occurring from Scandinavia, the Balkans, and Hungary, more or less across
the area evidently, in suitable localities. The Chinese Eozapus represents the
American subfamily Zapodinae (the only subfamily occurring in that con-
tinent). The Cardiocraniinae, containing two extremely rare types, Cardio-
cranius and Salpingoius, appears to be restricted to the more inaccessible parts
of Chinese Central Asia, except that a species of Salpingotus is known from
Afghanistan. Of the more specialized groups, the Euchoreutinae {Euc/ioreutes) is
restricted to the deserts of Inner China; the Dipodinae have, however, a wide
range outside Western Europe. Alhictaga and Dipus both appear to range from
South Russia across much of the Asiatic portion of the area, east more or less
to the North Chinese Pacific coast; Jaculus ranges across North Africa from
Morocco to Egypt and east as far as Persia; and generic types worthy ot note
confined to the Palaearctic with more restricted ranges are Scirtopoda, Paradipus,
and Pygeretmus.
The Sciuridae have, as usual, a wide distribution in the area; only in contrast
to the normal element (arboreal) in tropical areas, most of the Palaearctic genera
are Ground-squirrels. Citellus and Marnwta have the widest ranges, both
occurring in Europe as well as much of Asia, and both occurring again in
North America. Tamias, principally American, ranges in North Russia,
Siberia and China. Atlantoxerus, confined to Morocco and adjacent region,
represents a somewhat different type of Ground-squirrel tound chiefly in
52 DISTRIBUTION
Africa, and evidently not represented in either America or the Indo-Malayan;
Spermuphilopsis from Russian Turkestan area is probably a distant ally. Another
type of semi-terrestrial Squirrel is the Chinese Scitirotamitis, which seems nearest
to Ttimias in relationships. Tree-squirrels are represented by Sciuriis, which
occurs throughout Europe, across Russia and Northern (wooded) Siberia, and
parts of Eastern China, as well as in the Caucasus, but is absent from North
Africa, and much of the plains regions of S.W. Siberia. The Indomalayan
Callosciurus sends a few forms north into China. Of the Flying-squirrels,
Pteromys has the widest range, from Scandinavia across U.S.S.R. to Japan and
N.E. China. Contrary to Thomas's classification of Flying-squirrels, the genus
is here held to be an isolated specialized type with no very near allies, not a
near ally either of the American Glaucomys or the Indomalayan Hv/opefes.
Petaurista, from the Indomalayan, has a wide range in China, and includes
Japan and the Kashmir region, but is not known west of Kashmir nor in any
part of Siberia. Eupetaiirus is confined to Kashmir, and Trogoptcnis to parts of
China (though this genus touches the Indomalayan in some parts of China
south of the Yangtse). Eog/iiiicoinvs from Afghanistan completes the Palaearctic
list of Sciuridae. Two other families occur in the area, the Castoridae (with
one genus. Castor), now restricted to various localities in parts of Europe, such
as Scandinavia, and some of the larger Central European rivers, parts of Russia,
and the Mongolian Altai (the genus ranging across to North America), and the
Hystricidae, yvith one genus, Hystiix, specialized species of which extend from
India through Persia and Turkestan to Syria, and again in Italy, Sicily (where
they might have been originally introduced), and North-western Africa. This
section of the genus, however, finds its widest distribution in Africa south of
the Sahara.
The Nearctic regkix (Canada and the United States) contains roughly eleven
hundred named forms of Rodents distributed among eight families. The
Muridae are here in the minority as compared with all the others, only four
hundred forms appro.ximateiy belonging to them, while about six hundred and
ninety are named for the other families. This contrasts widely with the con-
dition found in the Palaearctic. Only two subfamilies of Muridae reach America
naturally at all (apart from the House-rats, and House-mice (Murinae), Raitiis
and 71///.?, which were originally introduced accidentally by man). These two
subfamilies are the Cricetinae and Microtinae. The Microtinae contain types
mostly much like those of the Palaearctic, as for example Microtus, Clethrion-
omys (ranging over much of the area), Dicrostonyx, Lemiiiiis (northern and Arctic),
Laguriis and Pitynivs (with more restricted ranges), (ienera peculiar to the area
are Ondatra (the largest member of the subfamily), Syiiaptoinxs (a Lemming),
Phenacomvs (one of the most primitive known \'oles), and A'eofiber (confined
to Florida).
The Cricetinae of North America appear to have " come up from the south "
rather than "in from the west," in that they are apparently more nearly related
to South American types rather than to Palaearctic Cricetinae. Of the seven
genera known north of Mexico, three only reach as far north as Canada, Peromys-
cus (which appears to cover the entire continent), Ncotonui, and Onychomys.
DISTRIBUTION S3
The genera Reithrodontomys, Sigmodon, and Orysomys have a fairly wide range
in the warmer parts of the United States, and all the above without exception
continue their ranges south into Mexico, all but Onychomys into Central
America, and the three last-named range into South America. Outside the
Muridae, three families occur which also range in the Palaearctic, and four are
at the present day confined to the New World.
The Sciuridac are represented, as in the Palaearctic, mostly by Ground-
squirrels, of whicli Marmota, Cite/Ins, and Tamias (all also in the Palaearctic),
each cover a great portion or most of the area. Cynomys, a rather isolated type
of Ground-squirrel, is purely North American; Tree-squirrels are represented
by Tamimciurus (American only), and the more widely ranging genus Sciurus,
which, however, covers relatively little of the area. Flying-squirrels are
represented only by Glaiicomys, which has, however, a very wide range. The
Castoridae are represented by the sole genus. Castor, which is Holarctic in
distribution. The Dipodidae are represented only by the primitive subfamily
Zapodinae {Xapaeozapiis, Zapus), which, however, covers almost the whole
area but seems not to occur south of it. We may now turn to four solely
American families. The Aplodontiidae, with one genus, Aplodontia, represents
an archaic type of Rodent confined at present to the western side of the Rocky
Mountains, but known to have occurred formerly in East Asia. The Erethizon-
tidae, a member of the Hystricoid branch, is represented by Erethison, note-
worthy as being the only Hystricoid adapted for a life in cold climates. Other
members of the family occur in the warmer portions of the Neotropical, from
Mexico southwards. The Geomyoid branch of the Order contains two families
both confined at present to America, extending south to Panama and Ecuador
though chiefiy northern in distribution. Of these the Heteromyidae is widely
distributed in the western portions of the United States, represented by the
more primitive Perognathiis, which touches western Canada, and the highly
specialized saltatorial Dipodomys and Microdipodops. The Geomvidae are repre-
sented by Thomomys, with very many named forms from the western and
central U.S.A., and ranging into western Canada, and Geomys, from the central
and eastern United States, including Florida. So far as I have traced it this
branch of the Order is not known fossil outside America.
The NEOTROPICAL ."^REA has more named forms than any other of the great
areas, if Mexico, the West Indies, and Central America are included in it.
Notwithstanding this, although South America is currently referred to as being
the " headquarters" of the Rodents, once Panama is passed there is a surprising
general similarity of type through the various groups found in that continent, the
members being cither Cricetine Muridae, or Sciuridae, or Caviidae, or members
of the supcrfamilv llvstricoidae (all of which are more or less closely allied to
each other). (A genus of Heteromyidae occurs in the extreme north.) There
is a lack of that wide divergence of structure which makes the study of Palae-
arctic, Nearctic, or African Rodents so interesting, and recalls the state of affairs
present in the Indcmalayan. In fact, so far as the "superfamilies" recognized
in this work are reckoned, fewer of them occur in South America than in any
other of the great areas except the Indo-Malayan.
54 DISTRIBUTION
Even in the Muridae there is incessant repetition of a single (Cricetine) type,
very different from the interesting differences to be met with in the different
subfamihes that occur in all the other great areas, even including the Indo-
malayan. Of about fifteen hundred named forms, roughly eight hundred are
Muridae, roughly seven hundred belong to other families.
In Central America (with Mexico and the West Indies), already a great in-
crease in Cricetinae and a great decrease in the more northern Microtinae is met
with. In the Cricetinae north of Panama, as well as the North American
genera Neotoma, Peiomvsciis, Oiivchoiiivs, Rcithrodoiitomys, Orvsomys and
Sigmodon being well represented, about fourteen more genera start their ranges
at once. Confined to the Central America area are Nvctomvs, Nelsonia, Ototy-
lomys, Scotinomys, HoJuinvs and others; while Tvloinvs, Nectomvs, and one of
the "Fishing-rats," Rluuiiivs, range north from northern South America. The
Microtinae range south to Guatemala only; the chief genera in the area being
Microtus and Pilvmys.
The Sciuridae are very much the same, as regards genera, as in the Nearctic,
except that Marmota and probably Tamimciuriis are absent. Few range south
of Mexico; but the genus Sciurus has many named forms from this area, and
extends into South America. G/aiicomys comes into Central America; while
Microsciunis comes up into Nicaragua from South America; but apart from
Sciurus and Microsciunis, no genus which occurs north of Panama crosses south
of it. The Castoridae touch extreme North Mexico. The Heteromyidae are
widely distributed through Central America, the primitive genera Hetcromvs
and Lioiiiys occurring more or less throughout, while Dipodoinys and Perognathus
are represented in Mexico. The Geomyidae are likewise common in the area,
and one genus, Macrogeomxs, ranges as far south as Panama. This group
(Heteromyidae -t- Geomyidae) is, however, not known in South America except
for a few forms of Heteromvs from the extreme northern countries.
Four families belonging to the Hystricoid branch occur in Central America ;
of these the Erethizontidae is represented by Coendou (Mexico southwards),
while the Dasyproctidae (Dasvproctti), and the Cuniculidae (Ciiiiiculus) start
their range which is, as in the case of Coendou, from Mexico southwards over
the greater part of tropical South America. The Echimyidae are represented
by three subfamilies, two of which, Capromyinae (Geocapromys, Capromys),
and Plaeiodontiinae (Plagiodontia), appear to be confined to the West Indies
(where surprisingly few genera of Rodents occur), except that a member of the
Capromyinae has been described from Venezuela ; the other, the more generalized
Echimyinae, being represented by Proechimxs, Diplortivs, and Hoploinys, from
Nicaragua southwards. The Caviidae are represented by Hydroclioerus which
extends north to Panama.
South of Panama, vast quantities of Cricetine Muridae swarm, belonging to
a very large number of named genera, which are in many cases not or barely
distinguishable from each other. These group themselves round the following
main types: Oryzomvs, which appears to occur in all parts from Patagonia to
Colombia, and has many close allies as Nectomvs, Rliipidoinvs, Tliomasomys,
and perhaps leads to the specialized and isolated North Argentine Scapteromys;
DISTRIBUTION 55
Akodon, with a very wide range in the continent, and with several allies the best
known of which is Oxvmycterus; probably leading to the specialized subfossorial
Notiomys of Patagonia; P/ivllotis, with its allies Hesperomys, Eligmodontia,
which series may lead to such dentally highly modified types as Chinchillula and
Irenomys; and Holochilus, with its allies Neotomys and Reithrodon which seem to
correspond to the Nearctic Sigmodon type.
By far the most interesting of the Neotropical Cricetines are the Fishing-
rats, Ichthyomys and Anoiornys, which must be among the most specialized of
all iMuridae, and in cranial characters parallel to a large degree the Australasian
aquatic members of the Hydromyinae.
The Squirrels of South America are, with the exception of Sciurillus from
the Guianas, which appears to be a type which one might consider archaic and
allied to the Indomalayan Naiviosciiirm, all essentially types which agree to such
a degree with the northern Sciurus that there appears no necessity to separate
them from that genus, except for a closely allied type Microsciurus . The
family ranges south to Jujuy (North Argentine) only. The type of Squirrel
found in South America suggests that the family has "come in" recently,
comparatively speaking, from the north, and has not been isolated from more or
less Eocene times or before in the Continent when it was (as generally admitted)
an island, as I suggest most of the Cricetines and Hystricoid types have. The
Caviidae, represented by two subfamilies Hydrochoerinae (Hvdroclioeriis only;
tropical portions), and the Caviinae, containing the more specialized Dolichotis
from the southern plains, and the more primitive Kerodon (Brazil), and Cavia
and its immediate allies which between them cover the continent, are confined
to the area. They are in this work not regarded as typical Hystricoidae, but
referred to a separate superfamily on account of the formation of the lower jaw.
The Hystricoid branch of the Rodents is represented in South America as follows :
Northern tropical forest area: Family Echimyidae: two subfamilies, the
Dactylomyinae, Thrinacodus and Dactylomvs; the Echimyinae, several genera
among which the arboreal Echimys and Mesomys and the terrestrial Proechimys
have the widest ranges. From South Brazil are known two rather distinct types
referable to the latter subfamily in Clyomys and Carterodon. Family Erethizon-
tidae : two very distinct subfamilies, the Chaetomyinae (Chaetomys only,
distribution evidently local), and the Erethizontinae {Echinoprocta: Colombia;
and Coendou, distribution general). Family Dasyproctidae : Dasyprocta and
Myoprocta (distribution general). Family Cuniculidae: Cuniculus (distribution
general?). The Family Dinomyidae (Dinnmys) is confined to Peru and Ecuador
region. The Subfamily Octodontinae (Echimyidae) is represented by Octodon
and Ctenomys as far north as Peru on the western side of the continent. In
Peru also Ltigidiiini represents the Family Chinchillidae. In the plains and moun-
tains of the southern part of the continent the following Hystricoid types
occur: Family Chinchillidae {Ciiinchilla, Lagidium, Lagostomiis). Family
Echimyidae : three subfamilies, Alyocastorinae (Myocastor only). Abrocominae
(Abrocoma only), and the Octodontinae, of which Ctenonivs, Spalacopus,
Acotuiemys, Octomys and Octoduii are the main genera. Besides these types, the
genus Heteromys (Heteromyidae) occurs in Colombia, \'enezuela, and Ecuador.
56 DISTRIBUTION
Africa is the only geographical area remaining to be discussed. This
continent must surely be considered the present headquarters of the Order
so tar as variation in character goes, in that it contains more superfamilies than
any other area, four (out ot eight) of which are now confined to the continent.
Roughly eleven hundred and fifty forms are named from the area; here once
again the Muridae are very much the dominant feature in that about eight
hundred of the above forms belong to the family. The African types, both of
Squirrels and Rats and even Porcupines, have a rather difl^erent aspect from
those of the Palaearctic or Indomalayan, and appear to be rather well separated
from them in general.
The Murinae possess a very large number of genera, most of which appear
to have a very wide range on the continent, and very few of which are at present
known from any other continent. The most distinct genera are Cricetoniys
and Saccostomus. Other aberrant but more typically Murine types are Lophur-
otnys, Acomys, Uraiiomvs, Mvlomys, Thamnomxs, Beamys, Dasxmys, Arvicanthis
and its immediate allies (Aizictiiithis ranging north to Kgypt and occurring in
Arabia, as does Acomvs), Oenoiiivs, Zelotomys, Colomys, etc. Besides these
occur many indigenous wild species of A/w, and various groups of Rattiis,
some of which have received generic names which appear quite unretainable.
The Dendromyinae is a group confined to the continent, very closely allied to
the Murinae, and containing Deiidromus and Steatumvs which have a wide range,
and Piionomvs and Malacotlirix, which are more restricted, the latter being one
of the most aberrant members of the whole family. Deomys, here regarded as
type of a distinct subfamilv the Deomyinae, is confined to the Congo. The
Otomyinae, with two valid genera Otvmys and Parotomys, are an interesting
group confined to the area. The subfamily Gerbillinae is very widely distri-
buted through the continent in suitable areas, containing more generalized types
in Tatera, Gerbilliis, etc., and some more local specialized genera as Dismodillus,
Dcsinodillisciis, Annnodilhis, etc. The Cricetinae is represented by one genus
only, I\Jvstrot)iys, from the south; but the Microtinae are not known e.xcept
in the Palaearctic coastal region. Even this does not exhaust the list of sub-
families, as Tachyoryctes, type of the Tachyoryctinae, though not hitherto
currently referred to the Muridae, is here regarded as a member of the family;
this genus is known from the eastern and central portion of the continent.
The Muscardinidae are represented by the subfamily Graphiurinae, the
genus GrapliiiDus ranging over most of the continent.
Eliomxs, a Palaearctic type belonging to the typical subfamily, ranges south
to the Rio de Oro. Lophiomys, here regarded as type of a family the Lophio-
myidae, is confined to the eastern part (Abyssinia, Somaliland, Kenya, Sudan).
The Dipodidae is represented by Jaciilus in the Sahara and Somaliland. The
family does not range south of this area.
The Ctenodactylidae is another northern African group, with very much
the same range collectively as the Dipodidae; the principal genera are Pectinator,
Massoutiera and Ctenodactylus. The group is known fossil from South Europe,
and from India. The Pedetidae, with one genus, Pedetes, is confined to the
continent, ranging in the south and east. The Anomaluridae is another family
DISTRIBUTION 57
peculiar to Africa, occurring mainly in the western forests, and containing two
subfamilies, the Anomalurinae {Anomalurus, Anomalurops), and the Idiurinae
{Zenkerella and Idiurus), these groups sometimes being given family rank.
The African Sciuridae consist of a relatively small number of forms, one of
which (Xeriis) is terrestrial, and is represented as indicated already in parts of
the Palaearctic, one of which, Mvosciiirus, is a pygmy form perhaps not distantly
related to the Indomalayan Sannosciiiriis, and the remainder of which are
arboreal types ot which Protoxerus, Heliusciuriis, Funisciurus, and Paraxenis
have the widest ranges. The Ilystricidae are widely distributed through the
continent ; the genus Hystrix is the dominant form of this family and here reaches
its highest degree of specialization ; while the genus Atherurus is found in the
western and central forests, with species of a rather more advanced type than
their Indomalayan cousins. Two other Hystricoid genera occur: T/irvonomys,
which is best referred to the family Echimyidae (otherwise American) as type
of a subfamily, which ranges through most of the continent, and which is known
fossil from India; and Petromus, which seems best referred to the same family
(as type of a subfamily), and which is known only from South-west Africa.
Yet another exclusively African family, the Bathyergidae, some of the most
isolated living Rodents, range collectively through most of the area: Cnptomys
has the widest range; other more highly specialized but more restricted types
are Bathyergus and Heterocephalus; and Heliophobius which appears unique in
the whole Order in dental formula has a moderate range on the eastern side.
In Madagascar Rodents are unknown save for half a dozen peculiar Muridae.
It has been the custom of late years to refer these to a subfamily (or familv)
the Nesomyinae or to place them in the Cricetinae. I have been able to find
no characters which keep them apart as a distinct subfamily, nor do they all
appear to be Cricetinae. In the present classification I have had provisionally
to refer them to no less than five dilTerent subfamilies. The names of these
genera are Eliurus, Brachyuromys, Xesomys, Gymnuromys, Hvpogeomvs and
Brachytarsomys. Their status will be discussed in the volume set aside for
RODENTS OF THE PALAEARCTIC
(OTHER THAN MURIDAE)
Genera, Prinxip.^l Species, and Approximate Ranges
SCIURIDAE
Genus Trogopterus
xanthipes. China; Tibet to Chihli.
Genus Petaurista
alborufus. China; Tibet to Hupeh.
stilcattis. China; Chihli.
albiventer group. Kashmir; Japan, Manchuria; Szechuan.
Genus Pteromys
tolans. Scandinavia across Siberia to Japan, Kansu.
Genus Eoglaucomys
fimbriatus. .Afghanistan, Kashmir.
58 DISTRIBUTION
Genus Eupetaurus
cinereiis. Kashmir.
Genus Sciiirus
vulgaris group. All Europe; Siberia, to Manchuria, Chihli, Japan.
anomalus group. Caucasus area.
Cienus Callosciurus
ntiic/el/andi group. Tibet; Chihli.
erythraeus. Szechuan. (Indomalayan type.)
Genus Dremomvs
pernyi. China; Szechuan, Ilupeh. (Indomalayan type.)
rufigems. China; Szechuan. (Indomalayan type.)
Genus Funambuhis
palmarum group. North Punjab. (Indomalayan type.)
tjenus Atlantoxerus
getulus. Morocco,
(jenus Spermopliilopsis
hptodactxlus. Afghanistan, Turkmenia.
(jenus Sciurotamias
davidianus. China; Kansu and Szechuan to Chihli.
Genus Tamias
sibiricus. North Russia, Siberia, China north of Yangtze, to Japan.
Genus Citellus
citelhis group. South-eastern Europe, Russia, Asia Minor; Shansi,
Kansu, Mongolia, Transbaikalia (dauricus, etc.).
suslicus. East Europe, South Russia.
fulvus. East Russia, Turkestan, Persia.
pygmaeus group. South Russia, Turkestan; Mongolia {pallidi-
cauda).
eversmanni . Russian Altai to East Siberia, North Mongolia.
Genus Marmota
mar/nota. Alps and Carpathians.
bobak group. Poland, Russia, Altai, North Mongolia, Kansu,
Transbaikalia, Tibet.
caudata group. Kashmir, Afghanistan, Russian Turkestan, Chinese
Turkestan.
caligata group. North-east Siberia.
CASTORIDAE
Genus Castor
fiber. Main rivers of Central Europe; Scandinavia; parts of
European Russia; Mongolian Altai.
CTENODACTYLIDAE
Genus Ctenodactxlus
gundi. North Algeria.
Genus Massoiitiera
mzabi group. North Algeria.
DISTRIBUTION ^^
DIPODIDAE
Oenus Stcista
subtilis group. Scandinavia, Denmark, Hungary, Balkans, Russia
Siberia to Lake Baikal.
concolor group. Caucasus, Altai, Kashmir, Kansu, Manchuria
aakhahn. '
Genus Eozapiis
setchuanus. Szechuan, Kansu.
Genus Cardiocrankis
paradoxus. Nanshan, Sinkiang (China)
Genus Salpingotus
kozlmi. MongoHa; Gobi.
crassicauda group. .Mongoha; Gobi, and in Afghanistan
Genus Euclioreutes "
naso. Chinese Turkestan, Inner Mongolia
Genus Allactaga
major group. Southern Russia, Russian Turkestan
s,bmca group. Eastern Caspian region to Kansu, Mongolia,
1 ransbaikalia, Chihli. *
elater group. Caucasus to Persia, Afghanistan, Kashmir, Chinese
lurkestan; also Mesopotamia [tniphratka).
wilhamsi. Caucasus, Asia Minor.
tetradactyla. North Egjpt.
Genus Alactagulus
pumilio. Caucasus, Russian Turkestan, Inner Mongolia
Genus Pygeretmus *'
platyunis. Western Russian Turkestan.
shitkovi. Eastern Russian Turkestan
Genus Dipus
'"S'tta. Caucasus across Russian Turkeston to -Mongolia and
Chihli. "
Genus Scirtopoda
telum group. South Russia, Russian Turkestan, Mongolia
Oenus Eremodipus ^
lichtensteini. Turkmenia.
Genus Jaculus
orientals. Palaearctic North Africa.
jaculus. Across Palaearctic North Africa, Syria, Persia
Uenus Faradtpus
ctenodactylus. Turkmenia.
HYSTRICIDAE
Genus Atherurus
Genus ;/• "'""'°"''"^- China; Szechuan. (Indo-Malayan type.)
subcristatus. China; Szechuan. (Indo-Malayan type.)
60 DISTRIBUTION
[Hystr:x) kuciita. Punjab, Afghanistan, Russian Turkestan, Transcaucasia,
Asia Minor, Syria, Palestine.
crislata. Sicily, South Italy, North-western Africa.
iMUSCARDINIDAE
Genus Mvomimus
personatus. Transcaspia.
Genus Eliumys
quercimis group. Continental Europe south of Baltic; Syria,
Northern Africa westwards from Tunis. Central and southern
Russia.
Genus Dyromys
nitechda. Central Europe (Switzerland), eastwards across Russia,
to N.W. Frontier (N. India), Tianshan, Zungaria. South in
Europe to Greece. Asia Minor.
Genus Glis
glis. Continental Europe south of Baltic; Asia Minor, Caucasus,
North Persia; Southern and Central Russia; South Turkmenia.
Genus Glirulus
japonicus. Japan.
Genus Muscardimis
uvelhmarius group. Europe, except Iberian Peninsula, including
England, Sweden ; parts of Russia.
SPALACIDAE
Genus Spalax
kirgisoriim group. Kirghiz Steppes(?), Syria, Palestine, North
Egypt and Libya.
monticolii group. Hungary, Roumania, Balkans, Asia Minor,
Caucasus.
nikrophthalmus group. Southern Russia, represented in Greece,
Rumania, Poland.
giganteus. Eastern Russia.
RHIZOMYIDAE
Genus Rliizomys
vestitu$. Szechuan.
RODENTS OF THE NEARCTIC
(OTHER THAN MURIDAE)
Genera, Principal Species, and Approximate Ranges
APLODONTIIDAE
Genus Aplodontia
rufa. Western U.S.A., from California into southern British
Columbia, Pacific side Rocky Mountains.
DISTRIBUTION 6i
SCIURIDAE
Genus Glaucomys
volans. Eastern U.S.A., from New York and Minnesota south-
wards, including Florida.
sabrintis. Labrador ; across most of Canada ; Alaska ; Pacific coastal
States of U.S.A., cast to Idaho; Virginia.
Genus Sciurus
carolinensis. Eastern U.S.A. and southern East Canada, west to
Minnesota, Oklahoma, including Florida.
griseiis. California, Oregon.
aherti. Colorado, Arizona, New Mexico.
niger group. Eastern U.S.A., from Texas and South Wisconsin
eastwards, including Florida; Arizona.
Genus Tamiasciurus
hudsonicus group. Most of Canada; Alaska; Western U.S.A.,
south to California, Arizona, New Mexico; Central U.S.A.
(Minnesota, South Dakota, etc.); Eastern U.S.A., south to
North Carolina at least.
Genus Tamias
alpinus. California.
minimus group. Western U.S.A., east to Wisconsin, west to
California and Pacific States; north to Yukon, Mackenzie, and
to Lake Superior.
amoenus group. Western U.S.A., coastal states, east to Montana,
north into Alberta and British Columbia.
quadrivittatus group. Montana, Idaho ; California east to Colorado ;
Arizona, Texas, New Mexico.
towiisendi group. Coastal states of western U.S.A., east to Utah,
New Mexico.
striattis group. Eastern U.S.A., west to Oklahoma and Minnesota,
south to South Carolina, north to Canada (Ontario).
Genus Ci tell us
tozvnsendii group. Washington, Oregon, Idaho, Utah.
'aashingtoni group. Washington, Idaho.
richardsoni group. California, Oregon, Nevada; Wyoming;
Saskatchewan.
parryii group. Oregon and Idaho north to Arctic Canada (xMac-
kenzie and east to Hudson Bay), and Alaska.
Iridecemlineatiis group. Western and West Central U.S.A., from
Arizona, New Mexico and Texas north to Minnesota, the
Dakotas, and Montana.
spilosoma group. Arizona, New Mexico, Texas, north to
(.')Nebraska.
Jranklinii group. Saskatchewan south to Oklahoma and Illinois.
variegatus group. Texas andColoradowesttoCaliforniaandOregon.
62 DISTRIBUTION
(Citellus) harrisii group. Texas, Colorado, Arizona, California.
tereticaiidiis group. California, Arizona, Colorado.
lateralis group, .\rizona, Colorado, Wyoming and Montana west
to Pacific coastal states, north into Canada (Alberta).
Genus Marmota
munax group. Across Canada from Labrador to Alaska, and
Eastern U.S.A., south to North Alabama, west to Kansas and
Minnesota.
flaviventris group. Western U.S.A., from South Dakota, Colorado
and New Mexico to Pacific states. Into British Columbia.
caligaUi group. Western Canada and Alaska, south to \\'ashington
and Montana, east to Alberta.
Genus Cynomys
ludovicianus group. The Dakotas and Montana south through
West Central U.S.A. to Texas and Arizona.
gunnisoni group. Slightly to the west of the range of /iidozicianin:;
Wyoming south into Arizona and New Mexico.
CASTORIDAE
Genus Castor
canadensis. "Most of North America from .Maska and Labrador
to the Rio Grande" (Anthony).
HLTEROMYIDAE
Genus Liomvs
irruratns. Southern Texas.
Genus Perognatlius
fasciatiis group. The Dakotas, Nebraska and Texas west to
Wyoming, Colorado, Arizona.
longitnembris group. California to L'tah and Arizona.
parvus group. California and Pacific states north into British
Columbia, east to Utah and Wyoming.
formosiis group. Utah, California.
hailevi group. Arizona, California.
Iiispidiis group. Kansas, Oklahoma.
penicillatus group. Texas, Arizona, California.
intermedins group. New Mexico, Arizona, California.
californiens group. California.
spinatus group. California.
Genus Microdipodops
megacephaliis group. California, Nevada, Oregon.
Genus Dipodomys
lieermanni group. California, Oregon.
spectabilis group. Arizona, New Mexico.
phillipsii group. I'exas.
merriami group. Texas and L'tah west to California.
DISTRIBUTION g
iDipodomys) ordii group. Oklahoma, Texas, Wyoming west to Oregon and
Lalifornia. °
anilis group. California.
compactiis group. Texas.
tmcrops group. Oregon, California, Arizona
desert! group. California.
^ .,„ GEOMYIDAE
Lrenus 1 homomvs
lowmendi group. California, Nevada, South Idaho
6o«<,. group. Oregon, Nevada, California, Colorado, Arizona
Mew Mexico, I exas.
alpinus group. California.
perpalUdm group. New Mexico and Utah west to California
Julvus group. Arizona, New Mexico, Texas.
umbrinus group. Arizona.
talpoides group. Colorado, Idaho and North Dakota, west to
Washington, north to Canada (Saskatchewan)
fossor group. California and Oregon east to Colorado and
Wyoming.
duuglasii group. Washington, Oregon.
monticola group. California, Oregon
^"''' AlbeZ' ^'''""^'""' O^'^g""' ^daho, Montana, north mto
bulbivortts group. Oregon.
Genus Geomys
tuza group Eastern U.S.A. (Alabama, Georgia, Florida)
Tl Upper Mississippi ^•alIey, to Kansas, Missouri Illinois;
v\est to Nebraska and the Dakotas
breviceps group^ Central U.S.A.; Nebraska south to New Mexico
Texas, and Louisiana. '
Genus Cratogeomys
castanops. Colorado, New .Mexico, Texas.
DIPODIDAE
iicnus Zapus
hudsonius group. Evidently most of Canada and U.S A east to
Labrador and North Carolina, west to .\laska, British Columbia
Genus Napaeo.aTJ ''''"' '" "' '''"'°™"' ^°"^' ^° -^^"' ^^^■^'-•
'""""t^^'Tn S'""'*' ='"'' ^•^•^- ^^^"^ 0"'^"° ^"d Wisconsin
to iNortn Carolina.
„ ^ , ERETHIZONTIDAE
Oenus Erethizon
dorsatum group. "Most of forested North America north of 40'
64 DISTRIBUTION
{Eretluzoti) and south ill the Rockies almost to Mexican bounciary"
(Anthony). North to Labrador and Alaska.
The more 1 examine North American faunal lists the more I am convinced
that too manv specific groups arc admitted, at least in the Order Rodentia.
RODENTS OF THE INDOMALAYAN REGION
(OTHER THAN MURIDAE)
Genfra, Principal Species, and Approxim.\tf. Ranges
SCIURIDAE
Genus Bclvmys
pearsoiii. Sikkim, Assam, 'I'ongking, Formosa.
Genus Trogoptenis
xanthipes. Yunnan. (Palaearctic type.)
Genus Fetaurista
petauristu group. Fukien; Formosa; Malay Peninsula, Sumatra,
Java, Borneo.
alhonifus group. Yunnan, Formosa.
puiiitdtiis. Malacca, Borneo.
tilhkciiter group. Ceylon, Peninsular India, Nepal, Burma, Siam,
Annam, Yunnan.
Genus Pteromxscus
pulverulenttis. Malacca, Sumatra, Borneo.
Genus Aeromys
tephromelas group. Malacca, Borneo.
tliomasi. Borneo.
Genus Hvlopetes
alboiiiger group. Nepal, North Burma; Philippines (iiigripes).
sagitia group. Burma, Laos, Malay Peninsula, Sumatra, Java,
Borneo, Natunas.
Genus Petinomys
fuscocapillus group. Ceylon, South India.
/;«^f«/' group. Sumatra; Philippines (cnw/ViH).
geniharbis group. Java, Borneo, Malacca. Hainan {electilis).
jf?ox!M group. Sumatra; Tenasserim.
Genus Petaurillus
hosei group. Malacca, Borneo.
Genus lomys
horsfieldi. Malacca, Sumatra, Borneo, Java.
Genus Nannosciurus
exilis group. Sumatra, Borneo, Philippines.
whiteheadi. Borneo.
melanotis group. Sumatra, Java, Borneo.
Genus Sciiirillus
muriniis. Philippines.
DISTRIBUTION g
Genus Callosciurus
macleUandi group. Nepal, Assam, Burma, Yunnan, Fukien
Haman, i'ormosa, Cochin-China, Siam, Annam
erythraeus group. Hainan, Formosa, Yunnan, Kwantung, Assam
Hurma, Siam, Annam, south to Pahang
camceps gxonY>. Chekiang; Tongking, Tenassenm, Siam south to
Malacca.
pygerythrm group. Nepal, Bengal, Assam, Burma.
quinquestnatus group. Yunnan, Burma.
prevosti group. Malacca, Sumatra, Borneo, Celebes
notatus group. Malacca, Sumatra, Java, Borneo
hippurm group. Malacca, Sumatra, Borneo, Philippines
teiiHis group. Malacca, Sumatra, Borneo.
loKi group. Malacca, Sumatra, Borneo.
leucomiis group. Celebes.
rubriventer group. Celebes.
Genus Dremomvs
pernyi group. Burma, Yunnan to Fukien, Hainan.
owstoni. Formosa.
everetti. Borneo.
lokriah group. Nepal, Assam, Burma.
rufigenis group. Malacca, Tenasserim, Burma, Annam, Laos,
Yunnan, Haman, Hupeh.
Genus Funumbtdus
palmarum group. Peninsular India, Ceylon.
layardt. South India, Ceylon.
siibliiieatus. South India, Cevlon.
Genus Ratufa
macroura. South India, Ceylon.
indica. Peninsular India.
gigantea. Assam, Nepal, Burma, Yunnan, Hainan
btcolor and other species. Burma, Tenasserim, S,am, Annam,
Malacca, Sumatra, Java, Borneo, Bali, Natunas
Oenus menetes
herdmorei. Burma, Tenasserim, Annam, Siam
Oenus Lariscus
insignis group. Malay Penii>sula (southern), Sumatra, Java, Borneo.
hoset. Borneo.
Genus Glyphotes
simus. Borneo.
Genus Rheitlirosciurus
macrutis. Borneo.
Genus Rhinosciunis
r.n H l"tic"i'd<,tus. Malay Peninsula (southern), Sumatra, Borneo.
Oenus Hyoscturus
heinrichi. Celebes.
5 — Living Rodents — I
66 DISTRIBUTION
Genus Mannutii
hobak group (liimalayaiia); Nepal, Yunnan. (Palaearctic type.)
Genus Sciurotamias
Jorresti. Yunnan.
HYSTRICIDAE
Genus Triclivs
lipura group. Malacca, Sumatra, Borneo.
Genus Athenirus
macrouriis. Hainan, Southern China, Tongking, Assam, Malacca,
Tenasserim, Sumatra.
Genus Theciirus
piwiilus group. Philippines; Sumatra {siinuitrae).
crassispims. Borneo.
Genus Hystiix
hraclivurus group. Malacca, Sumatra, Java, Borneo, Sumbawa,
Floras.
subcn'staius group. Sikkim, Assam, Burma, Tenasserim, Yunnan,
Fukien, Anhwei, Hainan, (?)Annam.
leuciiru group. Ceylon, Peninsulijr India, Nepal.
MUSCARDINIDAE
Genus Platacanthomvs
lasiunis. South Peninsular India.
Genus Typhlomys
cinereus. I'ongking, Fukien.
RHIZOxMYIDAE
Genus Rliizomys
vestitiis group. Burma; Fukien (davidi).
sinensis group. Assam, Y'unnan, Kwantung; South Siam, Perak
[paiinosus).
siimatrensis group. Tenasserim to Sumatra.
Genus Cannomys
badiiis. Nepal, Burma, Siam.
RODENTS OF AFRICA (OTHER THAN MURIDAE)
(With Arabia, but not including Palaearctic North Coastal Area)
GENKR.A, Principal Species, and Appro.ximate Ranges
BATHYERGIDAE
Genus Batliyergiis
siiillus. South Africa; Cape Pnivuice.
janetta. South Africa; Namaqualand.
DISTRIBUTION 5_
Genus Georychus
capensis. South Africa; Cape Colony.
Genus Cryptomys
mec/iou'i group. Angola, Rhodesia.
huttetttottiis group. Cape, Rhodesia, Nyasaland, Tanganyika
/fc//« section. North Nigeria, French Shari, North Congo Kala-
hari, Rhodesia, Portuguese East Africa.
zechi. West Africa (Togoland).
ochraceocinereus. Sudan.
bocagei. Angola.
Genus Heliophobim
argenteocinerem group. Portuguese East Africa, Rhodesia, Tan-
ganyika. South Congo, Kenya.
Genus Heterocephaliis
glaber group. Somaliland, Abyssinia, North Kenya.
SCIURIDAE
Genus Alyosciurus
pumilio. West Africa; Cameroons, Gaboon.
Genus Heltosciurus
gatnbianus. Gambia east to Abyssinia, south to Angola and Por-
tuguese East Africa.
ruwenzorii. Belgian Congo. Ruwenzori.
poetuis. Fernando Po, Gold Coast, Gaboon.
lucifer. Nyasaland.
Genus Paraxerus
cepapi group. Kenya {ochraceiis) south to Transvaal, Portuguese
bast Africa, Kalahari, and Ovamboland
palliaius group. Rhodesia, Portuguese East Africa, Zululand to
Kenya, Somaliland.
flavivittis group. Portuguese East .-yrica, to Tanganyika and
Kenya.
boehmi group. Congo, Sudan, Ruwenzori.
Genus Funiscmrus
lemniscatus group. Gaboon, Cameroons, Congo.
congkus group. Angola and South Congo.
pyrrhopus group. Sierra Leone east to Congo, .-Vngola, and
Ruwenzori.
Genus Protoxerus
stangeri. Gold Coast and Nigeria east to Kenya, south through
Congo to Angola.
Genus Mynilus
aubinii group. Liberia, .\shanti.
Genus Epixerus
wilsoni. Gaboon.
ebii. Gold Coast.
6S DISTRIBUTION
Cjenus Xrnis
rutilus group. Somaliland, Eritrea, Abyssinia, Kenya.
ervthropm group. Sudan, Kenya, Uganda, Sahara (Air), F^aive
Chad, to Sierra Leone.
capeiisis group. South and South-west Africa.
ANOMALURIDAK
Genus Anoinaliinis
fmseri. Gold Coast east to Uganda (PKenya), Tanganyika, south
through Congo to Angola.
peli. Guinea Coast, Ashanti.
piisillus. Congo, Gaboon.
Genus Aiiomahiiops
beecrofti. Sierra Leone eastwards to Congo.
Genus Zenkerella
!nsl£;ms. Spanish Guinea.
Genus hiiiiiiis
zenkeri. Canieroons, Congo.
macrotis group. Canieroons, Congo.
PEDETIDAE
Genus Pedetes
cafer group. Kenya and Angola to Cape Province.
CTENODACTYLIDAE
Genus Pectinatur
spekei. Abyssinia, Somaliland, Eritrea.
Genus Ctenodactylus
guiidi group. Tripoli (northern Sahara, west to Morocco).
Genus Alassoutiera
mzabi group. Sahara, south to Asben.
Genus Felovia
vae. Senegal.
DIPODIDAE
Genus Jaculus
jaciihis. Sahara, south to Asben, Sudan, Somaliland; and Arabia.
(The other species, orientnlis, appears Palaearctic in distribution.)
ECHIMYIDAE
Genus Petiomus
typicus. South-west Africa.
Genus Thryotiomys
swhideriamts group. liahr-el-ghazal and Uganda to Nigeria,
Angola, South Africa.
gregorianus group. Congo, Kenya, Nyasaland.
DISTRIBUTION 69
HYSTRICIDAE
Genus Atherurus
africanus group. Gambia, Sierra Leone, Nigeria, Congo, to Kenya.
Genus Hystrix
leucura group. Arabia.
cristata group. Senegal, Asben, Somaliland, Kenya, Uganda,
Tanganyika.
africaeaustralis group. South Africa, South-west Africa, Portu-
guese East Africa, to Tanganyika.
MUSCARDINIDAE
Genus Graphiurus
ocularis. Cape Province.
platyops group. South-West Africa, Rhodesia.
hueti group. Senegal, Liberia, Cameroons.
crassicaudatus. Liberia, South Nigeria.
surdus. French Congo.
monardi. Angola.
u'oosnami. Kalahari.
murinus group. Asben, North Nigeria, Sudan, Somaliland, Kenya,
Gold Coast south to Cape Province.
Genus Eliomys
quercinus group. (Prom Palaearctic) south to Rio de Oro.
LOPHIOMYIDAE
Genus Lophiomys
imhausi. Somaliland, Abyssinia, Sudan (Kassala), and Kenya.
RODENTS OF THE NEOTROPICAL
(OTHER THAN MURIDAE)
(According to Flower & Lydekker, Mexico should be included in this region.)
Genera, Princip.\l Species, and Approximate Ranges
SCIURIDAE
Genus SciuHllus
pusillus. (iuianas.
Genus Syntheosciurus
brochus. Panama
Genus Microsciurus
aljari group. Nicaragua south to Peru and Upper Amazon.
Genus Sciuriu
rariegatoides group. Mexico to Panama.
deppei group. Mexico, Nicaragua.
aberti group. Northern Mexico.
70 DISTRIBUTION
(.Samus) niger group. Mexico.
hoffmimi group. Nicaragua to Venezuela, Ecuador; Peru; North
Argentine (Jujuy).
aestuaiis group. Guianas, Venezuela, Eastern Brazil to Minas
Geracs.
stramincus group. Ecuador, Peru.
piicherani group. Colombia, Peru, Bolivia.
rhoadsi. Ecuador.
jlaiiimijer . Venezuela.
langsdorjfi group. Venezuela, Colombia, Peru, Ecuador, Bolivia,
Brazil to Matte Grosso.
Genus lamias
quadrivittatus group. North Mexico. (Nearctic type.)
Genus Citellus
mexicanus. Mexico.
spilosoma. Mexico.
variegatus group. Mexico.
annidatus group. Mexico.
tereticaudiis. North Mexico.
lateralis group. North Mexico.
Genus Cynomys
mexicanus. North Mexico. (Nearctic type.)
Genus Glaucomys
volans. Through Mexico to Honduras.
CASTORIDAE
Genus Castor
canadensis. Extreme North Mexico. (Nearctic type.)
HETEROMYIDAE
Genus Heteromys
anomalus group. Venezuela, Colombia, Ecuador.
desmarestianus group. Mexico to Panama.
gaumeri. Mexico (Yucatan).
nelsoni. Mexico (Chiapas).
Genus Liomys
pictiis group. Mexico.
crispus group. Mexico to Panama.
irroratus group. Mexico.
Genus Perogtiathus
fasciatus group. North Mexico. (Nearctic type.)
longimemhiis group. North Mexico. (Nearctic type.)
baileyi group. North Mexico. (Nearctic type.)
hispidiis group. Northern Mexico.
penicillatus group. North-western Mexico. (Nearctic type.)
intermedins group. Northern Mexico.
71
DISTRIBUTION
Genus Dipodomys
spectabilis group. Northern Mexico.
phillipsii group. Mexico.
merriami group. Northern Mexico.
ordii group. Northern Mexico.
deserti group. North Mexico (Sonora). (Nearctic type.)
GEOMYIDAE
Oenus 1 homomys
hottae group. North Mexico, Sonora. (Nearctic type )
perpalhdus group. North Mexico, Sinaloa. (Nearctic tvpe )
timbrtnus group. Mexico.
Genus Geomys
r.n p ^/'■^^'■^''/'^ g'-o"P- N.E. Mexico (Tamaulipas). (Nearctic type.)
yjenus. Fappo^eomvs }^'--J
hulleri group. Mexico; Jalisco.
Genus Cratogeornvs
castanops group. Through Mexico.
Genus Platygeomys
gymnurus group. South Central Mexico.
Genus Orthogeomvs
gran'dis group. South Mexico to Honduras, Salvador
uenus Heterogeomys
hispidus group. Southern Mexico.
Genus Zygogeomys
trichopus. Mexico; Michoacan.
Genus Macrogeomys
heterodus group. Nicaragua, Costa Rica, Panama.
. CAVIIDAE
Genus Cavta
aperea group. North Argentine (Tucuman, Corrientes), north to
,-. ^ , ^eru, Colombia, Venezuela, British Guiana.
Genus Galea
./.mV^group. Southern Brazil, Bolivia, Argentine (to Upper Rio
Genus Caviella
australis. Argentine to Patagonia.
shiptoni. Argentine (Catamarca).
niata. Bolivia.
Genus Kerodon
rupestris. Eastern (?) Brazil.
Genus Dolichotis
patagona group. Argentine to Patagonia (Cordoba southwards )
saltmcola group. Argentine.
72 DISTRIBUTION
Genus Hydrochoerus
hydrochaeris group. WarmcT portions of South America (exact
range not traced), north to Panama. Known to occur in
Brazil, Paraguay, British Guiana, Venezuela.
CHINCHILLIDAE
Genus Chinchilla
latiiger. Northern Chile.
Genus Lagidium
viscaccia group. Peru, Bolivia, Argentine, Chile (south to 50° S.).
Genus Lagostomus
maximus. Argentine.
DINOMYIDAE
Genus DiiKimvs
hranickii. Peru, Colombia, Ecuador, Upper Amazonia.
ECHIMYIDAE
Genus Echimys
dasythrix group. East Brazil; Bahia to Rio Grande do Sul.
hlainzilki group. East Brazil; Bahia to Parana.
thomasi. Island oflF Bahia, East Brazil.
armatus group. Guianas, Brazil (North ?), Venezuela.
chrvsurus group. Dutch Guiana, N.E. Brazil (Para).
satiirnus. Ecuador.
grandis group. Peru, Upper Amazonia.
Genus Isothrix
pictiis group. East Brazil (Bahia).
histriatus group. Peru, Venezuela, Brazil south to Matto Grosso.
Genus Diplomys
caiiiceps group. Panama, Colombia.
Genus Pioechimvs
cayennensis group. Nicaragua southwards to Guianas, Peru,
Bolivia, Minas Geraes.
cauicollis. Colombia.
iheringi. Island off Sao Paulo, Brazil. (Sao Sebastian Island.)
setosus group. East Brazil; Bahia.
Genus Hoplomvs
gvmnuriis group. Nicaragua, Panama, Ecuador.
Genus Cercomxs
cunicularius. Paraguay, and East Brazil (Minas Geraes, Bahia,
Pernambuco).
Genus Eurvzxgomutomys
spinosus. Paraguay, South-eastern Brazil.
Genus Clvomvs
lattceps. S.E. Brazil (Santa Catharina).
DISTRIBUTION
73
Genus Carterodon
sulcidens. South Brazil; (?Lagoa Santa).
Oenus Mesomys
Genus L.„.;^Kf '""^^ ^^^°"'^= '^"""^'"^ ^^^'^ *° Ecuador. Peru.
emiliae. Central Brazil; Rio Tapajoz.
Genus Procapromys
geayi. Venezuela.
Genus Capromys
pilorides. Cuba.
melanurm. Cuba.
nana. Cuba.
Genus Geocapromvs
brounii. Jamaica.
Genus P/.^iZ' " '""''■ '"'" '^'^"' ^"'^ "°"''"^^^)' -^ Bahamas.
aediim. Dominican Republic.
Genus Thrinacodiis
albicauda group. Colombia, Venezuela
Genus Dactylomys
dactylinus. Ecuador, Bolivia, Brazil (Amazonia).
peruanus. Peru.
Genus Kannabatcumys
amblyonyx. Paraguay, S.E. Brazil (Sao Paulo)
Genus Myocastor '
corpus. Chile, Patagonia, Paraguay, Argentina.
Genus Abrocoma
bennetti. Chile.
cinerea. Northern Argentina.
Genus Octomys
mimax. North Argentine (Catamarca, San Juan)
Genus Aconaemys '
Genus Octodm'" ^'™"^' ^°"'^"" ^"''''^' Argentina (Andes).
deous group. Chile, Peru.
Genus Octodontomvs
i^liroides. Bolivia.
Genus Spalacoptts
cyamis group. Chile.
Genus Ctenomvs
magellanicus section. Paraguay, North Argentina to Patagonia
torquatus section. South Brazil, Bolivia, North and Central
-Argentme.
leiicodon. Bolivia.
opimus section. South Peru, Bolivia to Chile and Patagonia
bolivtensis section. Bolivia.
74 DISTRIBUTION
ERETHIZONTIDAE
Genus Cluietomys
subspinosus. Brazil; tropical? (exact locality not traced).
Genus Echinoprocta
rufescens. Colombia.
Genus C'oeiidoii
prehensilis group. Colombia, Brazil (Matto Grosso, PPernambuco),
Bolivia.
bicolor group. Bolivia, Peru, Ecuador, Panama (rothschildi).
mcxicaimm group. Mexico, Panama.
paranaycnsis group. Paraguay, S.E. Brazil, Eastern Brazil ?
vestitus. Colombia, Venezuela.
DASYPROCTIDAE
Genus Mvoprocta
acoucliy. Cayenne, Amazonia.
pratti. Peru, Ecuador, Colombia, east to Manaos region, Brazil.
Genus Dasvprocta
punctata. Mexico to Panama.
rariegata. Peru, Ecuador, Colombia, Bolivia, Matto Grosso.
aguti. Guianas, Brazil. Allied forms in Lesser Antilles.
CUNICULIDAE
Genus Citniculus
paca group. Mexico, Panama, Ecuador, Colombia, Brazil, Cayenne.
Probably south to Paraguav.
taczauoivskii group. Ecuador, Venezuela. ?Peru.
DISTRIBUTION OF RODENTS:
SPECIAL WORKS OF REFERENCE
Miller: Catalogue of Mammals of Western Europe, 1912.
Vinogradov: Rodents of U.S.S.R., 1933, Tab. Anal. Faune de L'URSS, Inst. Zool. Ac.
Sci. 10, p. I.
Tate: Some Muridae of the Indn-Australian Region, Bull. .-^mer. Mus. Nat. Hist.
LXXII, p. 501, IQ36.
St. Leoer: Key to Families and Genera African Rodents, 1931, P.Z.S., p. 957.
Hollister: Smiths. Inst. Bull. 99, 1919; East African Mammals in U.S. National
Museum.
Miller: List of North ."Xmencan Recent Mammals, 1923, Smiths. Inst. U.S. Nat. Mus.
Bull. 128.
Anthony: Field Book of North American Mammals, Putnam, 1928.
Flower: Mammals of Egypt, P.Z.S., p. 369. 1932.
Wrouchton: Indian Mammal Survey. Joum. Bombay N.H. .See. XXVI, No. 2, 191S
{1919), p. 352.
Aharoni ; Aluridac of Syria and Palestine, Zeitschr. filr Siiugethierk. Bd. 7, 1932.
Robinson & Kloss : Nominal List of Oriental Sciuridae, 1918, Rec. Ind. Mus. XV, IV,
p. 171.
DISTRIBUTION 7S
Robinson ; List of Sumatran Mammals, Journ. Fed. Malay States Mus., VIII, appendix,
1918.
Taylor: Mammals of Philippine Islands, -Manila, 1934.
Iredale & Troughton : Check List of Mammals recorded from Australia. Memoir VI,
'934-
Gyldenstolpe : Neotropical Cricetinae, Kungl. Svenska. Vetens. Hand. II, no. 3, 1932.
Tate; Taxonomy of Neotropical Hystricoid Rodents, 1935; Bull. Amer. Mus. Nat. Hist.,
LXVIII, p. 295.
Blanford: Fauna of British India, 188S, London.
Jones : Mammals of South Australia, Adelaide, 1923.
Shelford: a Naturalist in Borneo, igi6.
Gee: Bull. Dep. Biol. Yenchiang Univ., 1929-30, 1,2; List of Mammals occurring in
China.
Shortridce: Mammals of South-West Africa, London, Heinemann, 1934.
Allen, G.M.: Check List of African Mammals, Bull. Mus. Comp. Zool. LXXXIII, 1939.
Rummler: Die Systematik und Verbreitung der Muriden Neuguineas, Send. Mitt.
Zool. Mus. Berlin, 23, heft i, 1938.
Osgood, W. H. : Mammals of the Kelley-Roosevelts and Delacour Asiatic Expeditions,
Field. Mus. Publ. Zool. iS, 1932, pp. 193-339.
Dammermann : On the Zoogeography of Java, Treubia, Vol. XI, livr. i, 1929, Appen-
dix I, Mammals, pp. 33-39.
ScHWARZ, E. : On the Evolution and Radiation of Mammalian Faunae. .\ct. Zool.
Stockholm, 5, 1924, pp. 393^423.
Order RODENTIA
Key to Superfamilies here recognized
Lower jaw much specialized, either by distortion outwards of the angular
portion, by specialized limb of masseter lateralis superficialis, or
by a conspicuous ridge extending along outside of jaw below level
of toothrow, for attachment of masseter medialis.
Infraorbital foramen not or scarcely transmitting muscle. Fibula
reduced and fully fused with tibia. (Masseter lateralis chief
agent in modifying form of mandible.)
Superfamily Bathyergoidae
Infraorbital foramen much enlarged for muscle transmission. Fibula
rarely reduced, not fully fused with tibia.
Masseter lateralis chief agent in modifying form of mandible.
Superfamily Hystricoidae
Masseter medialis chief agent in modifying form of mandible.
Superfamily Cavioidae
Lower jaw not much specialized, never with angular portion distorted
outwards and never with deep ridge extending along outside of jaw
for attachment of masseter medialis.
Infraorbital foramen not or scarcelv transmitting muscle.
Zygomatic plate completely beneath the infraorbital foramen.
Superfamily Aplodontoidae
Zygomatic plate more specialized, broadened and tilted upwards to
a greater or lesser degree, never completely beneath infra-
orbital foramen.
Jugal bone long, usually extending to lachrymal; fibula so far as
known not fully fused with tibia. No externally-opening
cheekpouches present. Cheekteeth normally complex.
Skull without well-marked postorbital processes ; jugal much
broadened; cheekteeth extremely hypsodont, not
cuspidate in pattern; external form much modified
for aquatic life, tail broadened, flattened, naked, the
vertebrae broadened. Superfamily C.astoroidae
Skull with postorbital processes, well developed in the
majority; jugal not specially broadened, cheek-
teeth usually not hypsodont, cuspidate in pattern;
78 RODENTIA
external form never modified for aquatic life, tail
normal, always fully haired. Superfamily Sciuroidae'
Jugal bone strongly reduced, never approaching lachrymal,
the zygoma sometimes complete without it, the whole
zygoma in some cases reduced, threadlike. Fibula, so far
as known, fully fused with the tibia. Large externally-
opening cheekpouches present. Cheekteeth with ten-
dency to become simplified. Superfamily Geomyoid.'VE
Infraorbital foramen always enlarged for muscle transmission.
Zygomatic plate very generally tilted upwards and broadened to
a greater or lesser degree (two exceptions out of approxi-
mately two himdred genera). The infraorbital foramen
never much enlarged. Fibula fused with the tibia.
Superfamily Muroidae
Zygomatic plate never tilted upwards, always narrow and com-
pletely below the greatly enlarged infraorbital foramen.
Premolars becoming suppressed, either absent, vestigial, or shed
in the adult.
Fibula fused with the tibia; cheekteeth rooted, complex in
pattern; angular portion of mandible weak, not
drawn backwards. Superfamily Dipodoidae
Fibula free from tibia; cheekteeth evergrowing, simplified
in pattern; angular portion of mandible strong,
drawn backwards to a certain degree.
Superfamily Ctenodactyloidae
Premolars not suppressed, not shed in the adult, normally as
large as the molars, and not reduced.
Fibula fused with the tibia; cheekteeth evergrowing, simpli-
fied in pattern; external form saltatorial; mastoids
much inflated. Superfamily Pedetoidae
Fibula free from tibia, so far as known; cheekteeth rooted,
complex in pattern; external form arboreal; mas-
toids not much inflated. Superfamily Anomaluroidae
I would point out, before dealing with the families and genera, that sub-
genera as here retained are equivalent in rank to sub-genera as understood by
American authors; and are not groups which must at once be given full generic
rank, as has been done so often by authors other than Americans, because they
form "natural groups" or because of convenience.
Some excellent remarks on the status of genera and sub-genera are given by
Osgood in his revision of the American genus Peromysais, to which I would
refer my readers.
' For the wide differences between Castor and the Sciuridae in external characters see
Pocock, Proc. Zool. Soc. London, p. 1171, 1922.
BATHYERGIDAE 79
The present author inclines to the view that systematic classification would
be none the worse if sub-genera were abolished altogether.
Superfamily BATHYERGOIDAE
As here understood this contains one living family.
Family BATHYERGIDAE
1S96. Thomas; Myomorpha, part; Family Bathyergidae.
1899. Tullberg: Hystricognathi ; Bathyergoniorpha, Family Bathyergidae.
1918. Miller & Gidley : Superfamily Bathy'Ergoidae ; Family Bathyergidae.
1924. Winge: Family Hystricidae, part, Bathyergini.
1928. Weber: Bathyercoidea ; Family Bathyergidae.
Geographical Distributio.n. — Africa: from Sudan, Abyssinia and Somali-
land, and from Gold Coast to the Cape.
Number of Genera. — Five.
Characters. — Zygomasseteric structure unique in the order; mandible
with angular portion distorted outwards to "allow passage
of a specialized and enlarged distal anterior limb of masseter lateralis super-
ficialis" (Miller & Gidley); paralleling the Hystricoidae in this respect, but if
anything even more developed than in the most specialized of these. Infra-
orbital foramen small, not or scarcely transmitting muscle; if so, only a small
strand in certain species, the degree of enlargement of infraorbital foramen
evidently in some cases variable individually.
Skull and external form much modified for fossorial life.
Number of cheekteeth vary-ing in the different genera; cheekteeth strongly
hypsodont, but not evergrowing; normally simplified to ring-pattern in adult
(excepting the genus Georychus).
A tendency present for the upper incisors to extend into the pterygoids.
Fibula reduced, fully fused with the tibia.
Remarks. — The peculiar jaw-muscle structure combined with the varia-
bility of the number of cheekteeth, and the variability of the
infraorbital foramen serve to isolate the Bathyergidae completely among living
rodents.
Elsewhere, there is a strong uniformity in the dental formula of any one group;
in some cases, as Sciuridae (cheekteeth ; or J), Dipodidae (cheekteeth ' or
:'), there is a difference in the formula, it is true; but in almost all cases the
extra premolar retained is vestigial and going; in a vast group like the Muridae
the formula of 'i cheekteeth is very general, only a very few Australian and
Philippine genera having it reduced to H. But in this family, three completely
different dental formulas, or possibly e\en four, are to be found in five genera.
These rodents certainly cannot be lumped in "Myomorpha," as was done
by Thomas and earlier authors, on account of the comparatively trivial character
of the fusion of the tibia and fibula ; nor can they be transferred to the Hystri-
coidea, "Hystricidae," as was done by Winge, presumably on account of the
similarity of the lower jaw in the two groups, though in VVinge's Hystricidae
8o BATHYERGIDAE
the Ctenodactylidae are included, which do not possess the Hystricoid type of
mandible. Nor does the infraorbital foramen transmit muscle here, as appar-
ently Winge is of the opinion that it does (or did), except to a very small degree
occasionally, as discussed below; nor in the Mystricoidae are the tibia and fibula
fused, though this is a character which Winge has used elsewhere as a division
in other families (Anomaluridae against Dipodidae, etc., page 7).
In zygomasseteric structure the Bathyergoidae differ from the Hystricoidae
chiefly in that in the latter group the infraorbital foramen is always very much
enlarged to transmit muscle, whereas in the present group it is usually not
enlarged at all ; this fact , combined with the lack of broadening of the zygomatic
plate present, appears to be a primitive condition.
According to Tullberg's figures, the temporalis muscle in this family appears
less reduced than is usual, taking up the whole of the hinder part of superior
portion of skull, and extending forwards nearly to level of anterior zygomatic
root (Georyclius capensis).
Digits of forefoot and hindfoot five, none reduced.
According to Tullberg, the radiale and intermedium of members of this
family are separate, alone of rodents (examined by him) except Ctenodactylidae.
Malleus and incus fused according to Tullberg, as in Hystricoidae, Cteno-
dactylidae, but unlike the remainder of the order.
Thomas, Ann. Mag. Nat Hist., 8, IV, p. 111, 1909, suggested that the
cheekteeth present in the various genera are probably as follows:
Heliopliohius
b
^15:
1.2.3.
m.
1.2.3.
Bathyergus and
Georyclius
4 ^
4
" 5:
1.2.
m.
1.2.
Heterocephalus
3
3
'•■ 3':::
I.
m. —
I.
" Foiiiarina "
{=Hetervccphalus
T
p. 34-
34-
phillipsi)
Miller & Gidley, with reference to Heliophubius which exceeds their
highest formula for a rodent (i), state: "In the Genus Heliophubius, with
the greatest number of teeth, there are never more than j' functional at one
time; the apparent addition of one tooth in the upper jaw and two in the lower
jaw to the maximtun Rodent formula is probably due to a specialized condition
of the milk-dentition."
Discussion of Genera. — The genus Bathyergus appears to have evolved in a
rather different way from the remainder of the
family in that the digging is done not so much with the incisors as with the
foreclaws. This has led to great enlargement of these claws, but not to any
great lengthening of the upper incisor roots, so that the upper incisors do not
show any inclination to extend to the back ot the palate, or the pterygoids. A
EATHYERGIDAE 8i
parallel to this, between Bathyergus (a "claw-digger") and Georychus (a "tooth-
digger"), is seen in Spalax against Myospalax; the two fossorial Microtinae
Ellobius against Prometheomys; etc.
In all other Bathyergidae, the claws remain relatively small, but the upper
incisors extend over the cheekteeth to the back of the palate or at extreme
development into the pterygoids.
lleUophobius is remarkable in that, as indicated above, it is the only rodent
known with l cheekteeth, and appears to be erupting teeth more or less
through life.
Cryptomys and Georyc/ius are closely allied types, with a dental formula of
i ; Georychus, confined like Bathyergus to a small range in South Africa, is the
only member of the family without simplified cheekteeth in the adult; Cryptomys
with a large number of named forms extends over most of the Continent.
Heterocephalus, from Abyssinia and Somaliland, is a most extraordinary
animal; alone among the rodents it has become practically naked, having lost
the fur almost entirely. Various other characters such as the fact that D.3 in
the manus is noticeably longer than D.4, the more strongly shortened and
Murine jugal, and the reduction of the cheekteeth to i, or even sometimes |,
leads me to believe that it should be separated from the rest as a "generic group."
Key to the Generic Groups of Bathyergidae
Fur reduced to a few scattered hairs. D.3 in manus markedly longer than
D.4. Jugal short, supported anteriorly by the zygomatic process of the
maxillary, its general form Murine. Cheekteeth becoming reduced
numerically: ^ or - Heterocephalus Group (Heterocephali)
3 2
Fur normal. D.3 in manus never markedly longer than D.4, usually
slightly or considerably shorter. Jugal long, forming the greater part of
the zygoma. Cheekteeth not becoming reduced numerically: -, or in
6 +
one genus, at full dentition, j. Bathyergus Group (Bathyergi)
The Bathyergus Group
Fur normal; eyes and ears, as usual in the family, greatly reduced; usually
D.2 in manus longer than D.3, the digits reduced in size from D.2 to D.5
evenly; pollex not vestigial, clawed. Hindfoot with D.3 remaining main digit,
except in Heliophobius.
General cranial characters as follows: Skull with frontals moderately or
rarely strongly constricted, nasals usually narrow; posterior root of zygoma
noticeably broad, and zygomata widely spreading. .\ prominent ridge developed
in all genera extending along centre of skull from posterior part of nasals to
lambdoid crest. Occipital region usually prominent, outstanding and strongly
ridged. Jugal long. Bullae small-moderate, not abnormal. Palate normally
excessively constricted between toothrows ; extending behind level of toothrows,
in which position it is broader, excepting Heliophobius. Incisive foramina
6 — Living KoUents — I
82 BATHVERGIDAE
obsolete. Angle of mandible powerfully distorted outwards; usually not pro-
duced far backwards, except in Bathyergus. Incisors thick, pro-odont.
Key to the Genera of the Bathyergus Group
Cheekteeth i.
.4. , .
Upper incisors not extending behind toothrows, and heavily grooved.
Foreclaws much enlarged. Angular portion of mandible pro-
duced considerably backwards. Bathyergus
Upper incisors extending behind the toothrows, in extreme develop-
ment into pterygoids, not grooved. Foreclaws not specially
enlarged. Angular portion of mandible not produced far back-
wards.
Cheekteeth simplified to ring-pattern in adult. Posterior tooth cut
early in life. Cryptomys
Cheekteeth retaining one inner, one outer fold to old age; posterior
tooth cut late in life. Georychus
Cheekteeth at full dentition -.
0
(Upper incisor extends mto pterj'goids; cheekteeth ring-shaped; fore-
claws not enlarged; angular portion of mandible not produced
far backwards.) Heliophobius
1^-- te.
Fig. I. B.^THYERGUS suiLLL's suiLLUS, Schreber.
B.M. No. 5.8.10.10. o ; -■ I.
BATHYERGIDAE: BATHYERGUS
83
Fig. 2. Bathyergus suillus suillus, Schreber.
B.M. No. 5. 8. 10.10, (J; X I.
Fig. 3. Bathyergu-s suillus, Schreber.
Mandible from below ; X i.
Genus i. BATHYERGUS, Illiger
1811. Bathyergus, Illiger, Prodr. Syst. Mamm., p. 86.
Type Species. — Mus maritimus, Gmelin.
Range. — South Africa: Cape Province, and Namaqualand; coastlands.
84 BATHYERGUS— HELIOPHOBIUS
Number of Forms. — Three.
Ch.\racters. — Skull essentially as described above: frontals in the rs'pe
species much constricted; occipital region extremely ridged
and powerful in old age. .\ngular portion of mandible produced considerably
backwards, the mandible being perhaps proportionately larger in relation to the
upper part of the skull than in any other member of the order.
Upper incisors one-grooved, lowers plain; roots of upper incisors not
extending to nor approaching posterior part of toothrow.
Cheekteeth hypsodont, wider than long, when cut, with an inner and outer
fold, which quickly wear down so that the tooth is ring-shaped in adult. Infra-
orbital foramen normal (small).
Size largest tor the family in the type species; ear conch absent: tail as long
as hindtoot, thick and flat, with long hairs growing outwards each side giving
a feather-like effect. Claws immensely developed in forefoot, particularly of
digits 2. I and 4; D.2 longer than D.3. Hindfoot with the centre digit longest,
D.2 shghtlv longer than D.4, hallux slightly longer than D.5. Claws of hindfoot
medium. Pollex with short claw, not vestigial.
Two well-marked species are known, the "giant" suillus, and the moderate-
sized ^(jHf/^rt, which appears to have a less heavily ridged skull.
Forms examined: suillus. janetta.
List of X.\.med Forms
I. B.\THYERGUS SUILLUS SUILLUS, Schreher
1782. Saugt. IV, p. 715. pi. 204B.
South Africa: Cape.
S>"non\-m: maritimus. Gmelin, 17SS, Linn. Syst. Nat. i, p. 140.
africatm. Lamarck. Voyages de Thunberg au Japon. etc.. 4,
34S, 179b.
;. B.-\THYERGUS SUILLUS INTERMEDIUS, Roberts
1926. .Ann. Transvaal Mus. XL p. 261.
Klaver, Cape Province.
5. B.A.THYERGUS J.A.NETT.\, Thomas S; Schwann
1904. .-^bstr. Proc. Zool. Soc. London, no. 2, p. 6.
Port Xolloth, Little Xamaqualand.
Genus 2. HELIOPHOBIUS, Peters
1846. Heliophobius, Peters, Monats. Her. Akad. Berlin, p. 259.
1890. Myosc.\lops. Thomas, Proc. Zool. Soc. London, p. 448. Xew name to replace
Heliophobius on the assumption that it was preoccupied by Heliophobus, Boisduval.
Type Species. — Heliophobius argenteocinereus, Peters.
Range. — Eastern and Central Tropical Africa: Kenya, Tanganyika, South
Congo, Xorth Rhodesia, Xyasaland.
Number of Forms. — Eight or nine are recognized.
Char.\cters. — Cheekteeth at full dentition ". The teeth are very infre-
quently all in place together; the anterior premolars being
shed before the posterior molars are cut. In fifty skulls available for examina-
tion only one No. 18. 6. 15. 6 has all six teeth in place together (one side of the jaw
HELIOPHOBIUS 85
only). 1 have not seen one with the six lower teeth in place together. The
normal number in place at once appears to be either i or ', but sometimes
there may be ', etc. and frequently there will be 5 teeth on one side of the jaw
and 4 on the other. The last tootli appears to be cut late in life. The teeth
when cut are with one external and one internal fold, but soon simplify to a
ring-pattern.
Upper incisor roots extending into pterygoids. Palate excessively narrow,
differing from that of Bathyergus, Cryptomys and Georychiis in that it does
not e.xtend behind the toothrows. Infraorbital foramen very small. Other
essential characters of skull as already described.
Tail and ears obsolete. Claws not excessively lengthened. Hindfoot differ-
ing from that of liatliycraus in that D.2 is the main digit rather than D.3, as in
the forefoot, though the hallux remains slightly longer than D.5.
Forms seen : albifrons, angoniciis, argenteocinereus, emini, kapiti, marungensis,
robustus, spalax.
List of Named Forms
Mr. R. W. Hayman has been kind enough to look through this genus for
me and reports that all the named forms "cannot in my view be more than
races of argenteocinereus, except spalax, which has the narrow posterior nares
reaching the level of last molars, and is distinguishable on this from all the
others." I fully agree with this conclusion.
1. HELIOPHOBIUS SPALAX, Thomas
1910. Ann. Mag. Xat. Hist. 8, VI, p. 315.
Taveta, near Kilimanjaro.
2. HELIOPHOBIUS .\RGENTEOCINEREUS .ARGENTEOCINEREUS, Peters
1852. Reise nach Mozambique, ZooL Saug. p. 140.
Tette, Lower Zambesi.
3. HELIOPHOBIUS ARGENTEOCINEREUS ANGONICUS, Thomas
1917. Ann. Mag. Nat. Hist. 8, XX, p. 314.
Bua River, Angoniland, East Rhodesia.
4. HELIOPHOBIUS ARGENTEOCINEREUS ROBUSTUS, Thomas
1906. Ann. Mag. Nat. Hist. 7, XVII, p. 179.
Mpika, N.E. Rhodesia.
5. HELIOPHOBIUS ARGENTEOCINEREUS MARUNGENSIS, Noack
1887. Zool. Jahrb. Syst. II, p. 223, pi. ix, fig. 25.
Marungu, South-east Congo.
6. HELIOPHOBIUS ARGENTEOCINEREUS EMINI, Noack
1894. Zool. Jahrb. Syst. VII, p. 559.
Simba Mucnna, near Mpwapwa, Tanganyika.
7. HELIOPHOBIUS .\RGENTEOCINEREUS K.\PITI, Hillcr
1909. Smiths. Misc. Coll. LII, part 4, p. 469.
Kapiti Plains, Kenya.
8. HELIOPHOBIUS ARGENTEOCINEREUS ALBIFRONS, Gray
1864. Proc. Zool. Soc. London, p. 123.
"East .Africa."
Synonym : ? pallidus. Gray, 1864, P.Z.S. London, 9. 124. "East Africa."
86 GEORYCHUS— CRYPTOMYS
9. HELIOPHOBIUS MOTTOULEI, Schouteden. (Not seen)
1913. Rev. Zool. Afr. 2, p. 203.
Kilong\ve, near Lake Kisale, Belgian Congo.
Genus 3. GEORYCHUS, Illiger
181 1. Georvchus, Illiger. Prodr. Syst. Mamni. p. 87.
Type Species. — Miis capensis, Pallas.
Range. — South Africa: Natal and Namaqualand to the Cape.
Number of Forms. — Three.
Characters. — Like Cryptomys, next to be described, but upper cheek-
teeth with one narrow inner and outer fold each, the folds
persisting; lower cheekteeth with one outer fold persistent and one inner fold
which tends to become weak or obsolete. Posterior cheekteeth cut late in life.
Upper incisor roots extending into pterygoids. Infraorbital foramen normal
(small). Externally with no special peculiarities; claws not enlarged; the digits
arranged about as in Bathyergus.
Forms seen: caperisis, canescens.
List of Named Forms
1. GEORYCHUS CAPENSIS CAPENSIS, Pallas
1779. Glires, pp. 76. 172, pi. VII.
Cape Colony.
Synonyms: buffoni. Cuvier. Ann. Sci. Nat. 1834, i, p. iq6.
hucops, Lichtenstein, Forsters Desc. Arnrn. Iter. ad. Maris
Aust. Teras Suscepto, p. 364, 1844.
2. GEORYCHCS CAPENSIS CANESCENS, Thomas & Schwann
iqo6. Proc. Zool. See. London, p. 165.
Knysna, South Cape Colony.
3. GEORYCHUS CAPENSIS Y.\TESI, Roberts
IQ13. Ann. Transvaal Mus. IV, p. 92.
Trans\aal.
Genus 4. CRYPTOMYS, Gray
1864. Cryptoiuys, Gray, Proc. Zool. Soc. London, p. 124.
1S64. CoETOMYS, Gray. Proc. Zool. Soc. Loiidon, p. 125- (Based on coeciitwns and
da?nayetisis).
Type Species. — Georvchus holosericeus, Wagner.
Range. — Africa, widely distributed: Togoland, Nigeria and Bahr-el-Ghazal
to the Cape; evidently not occurring in Kenya, nor Abyssinia,
nor vSomaliland.
Number of Forms. — Approximately forty-nine have been named.
Characters. — Skull, excepting in some species the infraorbital foramen,
without special peculiarity; about as usual in the family;
frontals not much constricted; mandible with angular portion not much pro-
duced backwards. Upper incisors plain, their roots extending behind the
CRYPTOMYS 87
toothrow. Cheekteeth }, a simple ring in adult, one inner, one outer fold
when unworn.
Claws normal; length of digits as Bathyergus, or with considerable tendency
towards D.2 and D.3 in the manus being subequal, or even in some seen D.3
is very slightly the longer. D.2 and D.3 in the hindfoot also often subequal.
Mammae usually 2 — i = 6 (Thomas). Tail shorter than hindfoot.
Infraorbital foramen variable, sometimes even individually. In some cases,
as in the giant species, tuellandi, it becomes as large relatively as in some Muroid
rodents, as Rhizomys, and surely must transmit muscle. Skull no. 20.1 1.3.227
at the British Museum shows a specimen in this state. In C. coecutiens, as
figured by Tullberg, a small strand of muscle passes through the foramen.
The infraorbital foramen is normal (small), or but very slightly enlarged in
damaremis, lugardi, beirae, zeclii (type not seen), molyneaiixi, micklemi,foxi, lechei,
kummi, and tchytei (slightly enlarged); in the type of nimrodi, the infraorbital
foramen of one side of the skull is small, on the other side slightly enlarged (which
proves that no specific groups may be based on this character) ; it is very little
enlarged in ansorgei (one of the giant mechoui group) ; moderate in blainei (same
group); relatively large in bocagei, coecutiens (type not seen), darlingi, jorisseni;
largest in amatus, hottentotus (type not seen), mellandi and mechowi (type not
seen).
The above notes, except when stated to the contrary, are based on type
skulls; there may be some individual variation perhaps within some of the species.
Four forms, mechowi, mellandi, ansorgei and blainei, separate rather sharply
as a group from the others on account of their relatively very large size.
Forms seen: amatus, ansorgei, beirae, blainei, bocagei, coecutiens, damarensis,
darlingi, foxi, holosericeus, hottentotus, jorisseni, kummi, lechei, lugardi, mechowi,
mellandi, micklemi, molyneauxi, nimrodi, talpoides, whytei, zechi.
List of Named Forms
Mr. R. W. Hayman has kindly looked through the large collection of the
genus Cryptomys at the British Museum, with a view to getting it into some
semblance of order, and reports as follows :
"The British Museum material of this genus seems to me to be divisible
into five groups, based primarily on presence or absence of white head-spot (this
is more reliable than was expected), secondarily on colour and size. Cranial
characters seem to be unreliable and cannot be correlated with the groupings
given here. It is obvious that manv of the so-called species listed here will
eventually be relegated to sub-specific rank.
I. Without head-spot.
(a) mechmci group. Large to very large, head and body 200 up to 260. \'ery
pale brown in all forms.
1. mechou'i, Peters. North Angola.
2. mellandi, Thomas. North Rhodesia and Angola.
3. ansorgei, Thomas & Wroughton. Central Angola.
4. blainei, Hinton. Central .\ngola.
88 CRYPTOMYS
(b) hotteritoius group. Small to medium-sized, head and body 100-150.
Drab or lawn, exceptionally blackish (talpoides).
5. hottentotus, Lesson. Cape Colony and Natal.
6. /(. talpoides, Thomas & Schwann. Cape Colony.
7. h. occlusus, Allen & Loveridge. S.W. Tanganyika Terr.
8. Ii. zobytei, Thomas. N.W. Nyasa, N.E. Rhodesia. (Treated as a
race of hottentotus by Allen & Loveridge, Bull. Mus. Comp. Zool.
Harv., LXXV, No. 2, p. 125.)
Fig. 4. Cryptomys d.\m.4Rensis, Ogilby.
B.M. No. 25.12.4.190, $; y 2.
9. eoecutiens, Lichtenstein. Cape Colony.
10. holosericeus, Wagner. Cape Colony, Orange Free State, Transvaal.
11. jorisseni, Jameson. Transvaal.
12. nimrodi, de Winton. S. Rhodesia.
13. amatus, Wroughton. N. Rhodesia and Katanga.
With head-spot.
(a) lechei group. Size medium to large, head and body about 125 to 200.
Colour ranging from blackish through seal-brown to slate and
silvery-fawn.
14. lechei, Thomas. N.E. Congo, N. Uganda. (Only 3 Uganda skins
seen, all lacked spot.)
15. kummi, Thomas. French Shari.
16. foxi, Thomas. North Nigeria.
CRYPTOMYS 89
17. lugardi, dc Winton. S.W. Africa and Kalahari.
18. micklemi, Chubb. N.W. Rhodesia. (St. Leger, Proc. Zool. Soc.
London, 1932, p. 964, considers micklemi lugardi.)
19. molyneauxi, Chubb. N.W. Rhodesia.
20. darlingi, Thomas. S. Rhodesia.
21. beirae, Thomas. Portuguese East Africa.
Fig. 5. CRYPTONrys DA^L\RENSIS, Ogilby.
B.M. No. 25.12.4.190, 9; X 2.
(A) diimarensis group. Size medium; head and body about 150. Pale sandy
brown.
22. damarensis, Ogilby. S.W. .\frica.
23. ochraccocinereus, Heuglin. Bahr-el-Ghazal, Sudan.
24. zechi, iVIatschie. Togoland, West Africa.
Authentic examples of 23 {ochraceocinereus) not seen. This is
periiaps a more artificial group than the preceding one, but Nos.
22, 23 and 24 do not seem to fit elsewhere.
3. With or without head-spot.
bocagei group. Colour cinnamon to drab; size small to medium, up to 150.
Head spot very variable.
25. bocagei, de Winton. Angola.
26. kubtingensis, IVIonard. Angola."
On account of the variability of the head-spot I think it will be desirable to
treat the hottentotus, lechi, damarensis and bocagei groups as sections of one
specific group, particularly bearing in mind the amount of variability met with
in the genus lleliupliobius in this character. I must add that it was I who origin-
ally suggested to Mr. Hayman that the presence or absence of this spot might
be used to divide Cryptomys into groups.
go CRYPTOMYS
The mechowi group is unquestionably very distinct from the remainder.
mechoioi Group
1. CRYPTOMYS MECHOWI, Piters
1881. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 133.
Malanje, North Angola.
2. CRYPTOMYS MliLLANDI, Thomas
igo6. Ann. Mag. Nat. Hist. 7, XVII, p. 178.
Mpika, N.-E. Rliodesia.
3. CRYPTOMYS .YNSORGEl, Thomas & Wroughtoii
1905. Ann. Mag. Nat. Hist. 7, XVI, p. 175.
Bihe, Central .\ngola.
4. CRYPTOMYS BLAINEI, Hiiit.)n
1921. Ann. Mag. Nat. Hist, g, VII, p. 372.
Loando River, Central .\ngola.
hottentotus Group
(typical section)
5. CRYPTOMYS HOTTENTOTUS HOTTENTOTCS, Lesson
1826. Voy. Coq. Zool. i, p. 166, pi. ii, fig. 2.
Paarl, Cape.
6. CRYPTOMYS HOTTENTOTUS T.-\LPOIDES, Thomas & Schwann
igo6. Proc. Zool. Soc. London, p. 166.
Knysna, Cape Colony.
7. CRYPTOMYS HOTTENTOTUS OCCLUSUS, Allen & Loveridge
1933. Bull. Mus. Comp. Zool. LXXV, no. 2, p. 125.
Uzung%ve Mountains, S.-W. Tanganyika.
8. CRYPTOMYS HOTTENTOTUS WHYTEI, Thomas
1897. Proc. Zool. Soc. London, p. 432.
Karonga, Lake Nyasa.
g. CRYPTOMYS COECUTIENS, Lichtenstein
1827. Brants. Muiz. p. 37.
Natal.
Synonym: hiilnigi, Smitli, 1829, Zool. Journ., p. 439. A synonym of
C. hottentotus, fide G. M. Allen.
10. CRYPTOMYS HOLOSERICEUS, Wagner
1842. Schreb. Saugt. Suppl. Ill, p. 373.
Graaf Reinet, Cape Colony.
11. CRYPTOMYS NIMRODI, de Winton
i8g6. Proc. Zool. Soc. London, p. 808.
Bulawayo, Rhodesia.
12. CRYPTOMYS .\MATUS, Wroughton
1907. Manchester Mem. 51, no. 5, p. 28.
.Alala Plateau, North Rhodesia.
13. CRYPTOMYS JORISSENI, Jameson
igog. Ann. Mag. Nat. Hist. 8, IV, p. 466.
Waynek, Waterhurg District, Transvaal.
CRYPTOMYS 9"
(lechei section)
14. CRYPTOMYS I.ECHEI, Thomas
1895. Ann. Mag. Nat. Hist. 6, XVI, p. 241.
Bellima, Monbuttu, N.E. Congo.
15. CRYPTOMYS KUMMI, Thomas
igii. Ann. Mag. Nat. Hist. 8, VH, p. 592.
French Shari Protectorate, Ironside Plateau, about 8° N. 22' E.
16. CRYPTOMYS FOXI, Thomas
1911. Ann. Mag. Nat. Hist. 8, VII, p. 462.
Panyani, North Nigeria.
17. CRYPTOMYS LUG.U^DI, de Winton
1898. Ann. Mag. Nat. Hist. 7, I, p. 253.
Kalahari, between Palapye and Ngami.
Synonym: micklemi, Chubb, 1909, Ann. Mag. Nat. Hist. 8, III, p. 35.
Upper Zambesi.
i8. CRYPTOMY'S MOLYNEAUXI, Chutb
1908. Ann. Mag. Nat. Hist. 8, II, p. 451.
Loano Valley, N.-\V. FLhodesia.
IQ. CRYPTOMYS DARLINGI, Thomas
1895. Ann. Mag. Nat. Hist. 6, XVI, p. 239.
Salisbury, Rhodesia.
20. CRYPTOMYS BEIR.\E, Thomas & Wroughton
1907. Proc. Zool. Soc. London, p. 780.
Beira, Portuguese East Africa.
{damarensis section)
21. CRYPTOMYS DAMARENSIS, Ogilby
1838. Proc. Zool. Soc. London, p. 5.
Damaraland.
22. CRYPTOMYS ZECm, Matschic
1900. Sitz. Ber. Ges. Nat. Fr. Berlin, no. 4, p. 146.
Middle Volta, Togohind.
23. CRYPTOMYS OCHI^ACECJCINEREUS, Heuglin
1864. Nov. Act. Ak. Caes. Leop. Dresden, XXXI, p. 3.
Bahr-el-Ghazal, Sudan.
(bocagei section)
24. CRYPTOMYS BOCAGEI, de Winton
1897. Ann. Mag. Nat. Hist. 6, XX, p. 323.
Hanha, Angola.
25. CRY'PTOMYS KLB.VNGENSIS, Monard
1933. Bull. Soc. Neuchatel. Sci. Nat. 57, p. 58.
Cubangu River, Mossamedes, -Angola.
There then remain to be discussed twenty-three "species" (?) of Roberts.
Some comments on some of these have already been made by Oldfield Thomas,
Ann. Mag. Nat. Hist. 8, XX, p. 444, 1917.
92 CRVPTOMYS
It is useless attempting any remarks on these, as all are unrepresented; they
are therefore listed alphabetically.
2b. CRYPTOMYS ABERUANS. Roberts
1913. Ann. Transv. Mus. IV, p. 98.
Port St. Johns, Cape Province.
-7. CRYPTOMYS ALDUS, Roberts
1913. .Ann. Transv. Mus. IV, p. 100.
Wynberg, Cape Colony.
28. CRYPTOMYS ANOMALUS, Roberts
1913. Ann. Transv. Mus. IV, p. 96.
Transvaal, Pretoria.
2v. CRYPTOMYS AREN.ARIUS, Roberts
1913. .Ann. Transv. Mus. IV, p. 96.
Transvaal, Pretoria.
30. CRYPTOMYS BIG.\LKEI, Roberts
1924. Ann. Transv. Mus. X, p. 73.
Glen, Orange Free State.
31. CRYPTO\n'S CRADOCKENSIS, Roberts
1924. Ann. Transv. Mus. X, p. 73.
Cradock, Cape Province.
32. CRYPTOMYS JAMESONI, Roberts
1913. .Ann. Transv. Mus. IV, p. 95.
Trans\'aal, Johannesburg.
33. CRYPTOMYS JUNODI, Roberts
1926. .Ann. Transv. Mus. XI, p. 260.
Masiene, Portuguese East Africa.
34. CRYPTOMYS KOMATIENSIS, Roberts
191 7. .Ann. Transv. Mus. V, p. 272.
Amhemburg, Transvaal.
35. CRYPTOMYS LANGI, Roberts
1929. .Ann. Transv. Mus. XIII, p. 119.
Keerkloof, Natal.
36. CRYPTOMYS MAHALI, Roberts
1913. .Ann. Transv. Mus. IV, p. loS.
Transvaal.
37. CRYPTOMYS MELANOTICUS. Roberts
1926. .Ann. Transv. Mus. XI, p. 260.
Makoetsi River, N.E. Transvaal.
38. CRYPTOMYS MONT.ANUS, Roberts
1926. Ann. Transv. Mus. XI, p. 260.
Klapperklop, Pretoria, Transvaal.
39. CRYPTOMYS NATALENSIS, Roberts
1913. Ann. Transv. Mus. IV, p. 94.
Natal, Wakkcrstroom, Transvaal.
CRYPTOMYS 93
40. CRYPTOMYS ORANGIAE, Roberts
1926. Ann. Transv. Mus. XI, p. 259.
(Jlcn, Orange Free State.
41. CRYPTOMYS PAI.KI, Roberts
1917. Ann. Transv. Mus. VI, p. $■
Vaal River, Transvaal.
42. CRYPTOMYS PRKTORIAE, Rolierts
1913. .Ann. Transv. Mus. IV, p. 99.
Transvaal, Pretoria.
43. CRYPTOMYS RUFULUS, Roberts
1917. Ann. Transv. Mus. V, p. 272.
Tzaneen, Transvaal.
44. CRYPTOMYS STELLATUS, Roberts
1917. Ann. Transv. Mus. V, p. 272.
Komatipoort, Transvaal.
45. CRYPTOMY'S TR.ANSVAALENS1S, Roberts
1924. .Ann. Transv. Mus. X, p. 73.
Pretoria district.
46. CRYPTOMYS VANDAiMI, Roberts
1917. Ann. Transv. Mus. V, p. 273.
Leydsdorp, Transvaal.
47. CRYPTOMYS VETENSIS, Roberts
1926. Ann. Transv. Mus. XI, p. 259.
Vet River, Orange Free State.
48. CRYPTOMYS VRYBURGENSIS, Roberts
191 7. Ann. Transv. Mus. V, p. 274.
Vryburg, British Bechuanaland.
Addendum ;
CRYPTOMYS NATALENSIS NEMO, G. .M. Allen.
1939. Bull. Mus. Comp. Zool. LXXXIII, p. 429.
Manetsi River, near Malala, Zoutspansberg district, Transvaal.
Synon>Tn: pallidus, Roberts. 1917, Ann. Trans. Mus. V, p. 278. Not
of Gray.
The Heterocephaliis Group
Cheekteeth ij or |, simplified in adult. Size smaller than in other
members of the family. Fur practically absent, the hairs occurring singly,
scattered, throughout the body, most developed on the feet. Tail longer than
hindfoot. Eyes and ears extremely small, no ear conch. Forefoot with five
digits, the centre of which is the longest. D.5 and especially the pollex shorter
than D.4 and D.2, which are subequal. Hindfoot like forefoot, but hallux
about as long as D.5.
Essential cranial characters as in Bathyergus group, but jugal reduced; more
Murine in appearance; palate not continuing behind molars, and in appearance
rather less constricted normally than in other genera; upper incisors extending
behind the toothrows.
94
HETEROCEPHALUS
Genus 5. HETEROCEPHALUS, Ruppell
1S42. Heterocephalis, Ruppell, Mus. Scnckenberg. Abh. 3,
1903. FoRNARiNA, Thomas, Proc. Zool. Soc. London, p.
phillipsi, Thomas.)
Type Species. — Heterocephalus glaber, Ruppell.
R.^NGE. — Known from Abyssinia, Somaliland, Kenya.
Number of Forms. — Four are here listed.
Heft 2, p. 99.
336. {Heterocephalus
Fig. 6. Heterocephalus glaber glaber, Ruppell.
B.M. No. 32.2.19.9, ?; > 2h-
CH.'UiACTERS. — As indicated above. Frontals little constricted; nasals
appear rather broader than in other genera; palate shorter
than in other genera except Ihliophobiiis. Cheekteeth simple in adult, the usual
folds found elsewhere in the family present when unworn ; normally ij; in
H. phillipsi, so far as known, reduced to f , evidently at a certain age or stage
of wear; this species is represented in the British Museum only by three skulls;
two of these have two upper teeth on each side, the third has three upper cheek-
teeth on one side, the posterior one minute, two on the other side, the posterior
one apparently having been shed. This species was made the type of a genus
"Foriiaiina" by Thomas, but much more evidence on the condition of this
form is required before any generic separation can be done; I should be
HETEROCEPHALUS 9S
quite content to assume that if enough specimens could be brought to hand
phillipsi would turn out to be no more than a race of H. glaber^
In the few available for examination, including skulls which have been made
types of two or three "species," there is much variation in the size of the cheek-
teeth, M.3 being in some only slightly, and some very considerably smaller than
M.2, and of the incisors, which reach their maximum size in the type of dunni.
There is also variation in the form of the coronoid.
Hollister, 1919, East African Mammals in the U.S. National Museum,
synonymizcs several forms with the typical race; this classification is here
followed. Provisionally I list all named forms as either synonyms or races of
the type.
Forms seen : ansorgei, glaber, dunni, phillipsi.
Fig. 7. Heteroceph.\ll"s glaber glaber, Ruppell.
B.M. No. 32.2.19.9, 9; X 3 J.
List of Named Forms
I. HETEROCEPHALUS GLABER GL.A.BER, RUppell
1842. Mus. Senckenberg, .Abb. 3, Heft 2, p. 99.
Shoa, .Abyssinia.
Synonj-m: glaber progrediens, LSnnbcrg, 191 1, Kungl. Sv. Vet. Akad.
Handl.Bd.48,no.5,p. 102. North of GuasoNyiro, Kenya.
ansorgei, Thomas, 1903, Proc. Zool. Soc. London, p. 336.
Makindu district, Kenya.
stygius, .Allien, 1912, Bull. Mus. Comp. Zool. LIV, p. 444.
Neumann's Boma, Nth. Guaso Nyiro, Kenya.
* Since the above was written, the Check List of .African Mammals of G. ^L Allen has been
published ; in this H. phillipsi is considered a synonym of H. glaber.
96 HYSTRICOIDAE
2. HETEROCKPHALLIS GI.ABICR SCORTICCCII, de Heaux
1934. Atti. Soc. Ital, Sci. Nat. LXXIII, p. 2S3.
Gardo, Italian Somaliland.
(A synonym of o'. glaber, according to G. M. Allen)
3. HE'l'EROCEPHALUS GLABER DUNNI, Thomas
ujoq. Ann. Mag. Nat. Hist. S, IV, p. log.
Wardairi, Central Somaliland.
(A synonym of ^. glaber, according to G. M. Allen)
4. HETEROCEPHALUS GLABER PHILLIPS!, Thomas
iSSj. Proc. Zool. Soc. London, p. 612.
Somaliland.
{A synonym of ,i^. glaher, according to G. M. Allen)
The references and type localities for all members of the family Bathyer-
gidae are the work of Mr. R. W. I layman.
The family have been described fossil from the Oligocene of Mongolia. This
indicates a former wide distribution for the group, and contrasts with some of
the other African families as Anomaluridae and Pedetidae which do not seem to
be known outside the Continent.
BATHYERGIDAE:
SPECIAL WORKS OF REFERENCE
TuLLBERG, Nova .Acta Reg. Soc. Sci. Upsaliensis. XVIII, ser. 3, no. i, 1899.
HoLLiSTER, .Smiths. Inst. Bull. 99, p. 159, 1919; East African Mammals in the U.S.
Nat. Mus. Note on status of some forms of Heterocephalus.
St. Leger, Key to Families and Genera of African Rodentia, Proc. Zool. Soc. London,
1 93 1, p. 976.
Thomas, Ann. Mag. Nat. Hist. ser. 8, vol. IV, p. 1 10, IQ09. Note on dental formula in
the family.
Superfamily HYSTRICOIDAE
This group is equal in rank to the "lateralis-scries" of the superfamily
Hystricoidae of Miller & Gidley; or to the "Hystricomorpha" of authors not
including Caviidae (Cavia, Galea, Caviella, Kerodoii, Dolichatis, Ilydrochomis)
nor Ctenodactylidae; it is here divided into seven families.
1896. Thomas: Hystricomorpha, part, included Pedetidae, Caviidae, Ctenodactylidae.
1899. TuUberg: HvsTRicoGNATHi : Hystricomorpha; part, included Caviidae.
1918. Miller & Gidley: Superfamily Hystricoid.'\e, part, lateralis series. (Included,
as medialis series, the Caviidae.)
1924. Winge : Family Hystricidae, part, included IJathyergidae, Ctenodactylidae and
Caviidae of this work.
1928. Weber: HYSTRICOIDAE, part; included Caviidae, Ctenodactylidae.
Geographical Distribution. — The greater part of the American Con-
tinent from Canada to Patagonia (evidently
absent only from certain areas of southern U.S.A.); the greater part of the
African Continent; the Indo-Malayan region, from the Himalayas to Ceylon
and from Southern China to Borneo and the Philippine Islands; represented
in the Palaearctic in Italy, coastal regions North-west Africa, and in South-
western Asia (north into southern Siberia).
HYSTRICOIDAE 97
Characters. — Zygomasseteric structure differing from that of all members
of the order, except Bathyergidae, in that the lower jaw,
paralleling the Bathyergidae, has the angular portion of the mandible distorted
outwards, to a greater or lesser degree, "to allow passage of a specialized and
enlarged distal anterior limb of masseter lateralis superficialis, its general
direction parallel with zygoma" (Miller & Gidlev); combined with the fact
that the infraorbital foramen is very much enlarged to allow passage of masseter
medialis; the zygomatic plate is narrow, and remaining completely below it,
the general arrangement of the forepart of the skull as regards muscle insertion
(infraorbital foramen, zygomatic plate), essentially as in Anomaluridae, Cteno-
dactylidae, Pedetidae and Dipodidae.
Skull normally specialized, with broad frontals, which rarely show much signs
of interorbital constriction, a tendency present towards complexity of zygoma
(Echimyidae), and lengthening and specialization of paroccipital processes.
Dental formula: i. — c. -> p. -> m. - ^ 20.
I o "^ I 3
Cheekteeth usually flatcrowned, usually hypsodont, often evergrowing,
not cuspidate in adult.
A tendency present towards reduction of the digits of the hindfoot (in some
forms, Dasyproctidae, Lagostomits, hindfoot with three digits only).
The malleus and incus are fused according to Tullberg, though in some
cases apparently not completely so.
The tibia and fibula remain distinct, or are not fully fused.
Remarks. — This group has been recognized as one of the major groups of
the order by all authorities. But many forms which appear to
me not to belong are currently included in it. The Hystricomorpha or Hystri-
coid series are always described as with the angular process distorted outwards,
as indicated above; if this is a sufficiently important character on which to base
superfamily grouping, and it apparently is so (Tullberg divided the whole order
into two great groups, Hvstricognathi and Sciurognathi, based on its presence
or absence), it seems clear that forms which do not agree in mandible structure
with typical Hystricoidae must be excluded from that superfamily, no matter
what their ancestors may have been. This takes the Caviidae into another
branch of the order, as thev cannot by the longest stretch of imagination be
regarded as with typically Hystricoid mandible formation. The close associa-
tion ot Caviidae with such forms as Dasyprocta and Cunicuhis by manv authors,
Tullberg among them, has long struck me as extremely unnatural; Cuniculus is
of course one of the most isolated and aberrantly specialized living rodents, and
has not even the feet structure and external specialization of Caviidae and
Dasyprocta; but the last two named, both clearly parallels in evolution, both
highly modified for cursorial life, with digits of hindfoot reduced to three, are
yet so clearly totallv different when lower jaw structure and dental structure are
looked into. Such an association appears as unnatural to me as dumping Castor
and Myocastur into a family together because both swim !
Of the forty-three genera belonging to the group, thirty-six are confined to
the neotropical region. One is peculiar to North America ; of the remainder, two
7 — Living Rodents — I
98 HYSTRICOIDAE
are African, two (Hystricidae) the Malay Islands, the remaining two (Hystricidae)
cover a wide area in the southern palaearctic, the African and the Indo-malayan
regions.
This group, together with the Cavioidae, contains all the giants of the order
(except Castor), and exhibits some extreme tvpes of external specialization.
Taking all their characters into consideration, it appears to me that this group is
without doubt, broadly speaking, the most highly specialized and progressive
branch of the order, particularly such forms as Ilvstrix cristata, though in all
cases such species grade down quicklv to relatively low primitive allies.
The division of the group into families is not easy. As many as thirteen
have been recognized by various authors. It appears to me to be both unneces-
sary and inconvenient to divide them into as many as this; particularly taking
into account that all seem closely allied to each other, and that elsewhere vast
groups like the Muridae (with Cricetinae, Microtinae, Gerbillinae, Myospa-
lacinae, etc., etc.), are usually retained as one family. Flower & Lydekker,
1891, Mammals Living and Extinct, recognized (of Hystricoidae as here under-
stood) only five families, the Octodontidae (=Echimyidae of this paper),
Hystricidae, Chinchillidae, Dasyproctidae and Dinomyidae. Thomas in 1896
very properly separated the American "Porcupines" from the Hystricidae as a
distinct family; and Miller & Gidley, Pocock and others have recognized
Cuniculiis as type of a distinct family. With these two modifications I retain
the classification of Flower & Lydekker.
The Echimyidae as here understood contain a large quantity of neotropical
rodents and two African genera ( Thrvoiioinys, Petroinm) ; broadly speaking this
group contains forms which have not become much modified externally (excep-
tions: Mvocastor, Tlirvoiioinvs, Dactv/oiiivinae, part), but which seem to have
their specialization in the skull characters (lengthened or specialized paroccipital
process, some tendency to enlargement of bullae, very general tendency for
complex zygoma, etc.); and in dental characters (such as the rootless simple
teeth in Octodontinae). Normally the size is relatively small, though Mvocastor
and Tlii'vonomxs provide exceptions to this. The relationships of the various
groups will be discussed below.
Apart from the evergrowing plain laminate cheekteeth, there is little to dis-
tinguish the Dinomyidae from them; and certain species of Echimyidae mav
attain laminate cheekteeth, though this not combined with extreme hypsodonty.
But Dinomys is evidently rather an isolated specialized type, and is best retained
as a family. The Chinchillidae are dentally like Dinomys, but cranially with
their poorly-ridged mandible, the tendency either to lengthening of paroccipital
process or extreme inflation of mastoids and bullae, they stand rather apart not
onlv from Dinomvs but as a group from all other Hystricoids apparently; the
functional digits of the hindfoot are reduced to three. The lirethizontidae are
more primitive than the Echimyidae cranially and dentally (as a rule), but very
much more highly specialized externally, the feet attaining arboreal specializa-
tion not seen elsewhere in the order, and the spiny covering of the body being
in a very different class from that of the few spiny members of the Echimyidae,
in all but the very lowest. The Hystricidae are held not to be closely related
HYSTRICOIDAE 99
to the Erethizontidae, but rather to Dasyproctidae ; once again their lower
members are less specialized than in Echimyidae, as regards cranial char-
acters, their higher ones very much more so, with tendency to extreme inflation
of nasals without parallel in living rodents. The external covering presents
extreme specialization in development of spiny covering, hut goes through an
interesting series of grades of development, so that the lowest is much less
specialized in spiny covering than the higher members. The Dasyproctidae
are very similar to the Ilystricidae in cranial and dental characters; externally
they are very different, and not less specialized in their way, being modified for
cursorial life, with three digits only to the hindfoot, and in external form calling
to mind a type that primitive ancestral ungulates must have at one time passed
through in their evolutionary history. Finally the Cuniculidae in the develop-
ment of their vast bony cheek-plate present cranial characters very widely different
from any other rodent.
Key to thk Families of Hystricoidae
Entire zygomatic region abnormally modified by growth of bony cheek-
plates. (Cheekteeth strongly hypsodont, the folds of the teeth
isolating as islands on crown surface; form heavy; digits not
numerically reduced on hindfoot, the claws thick, more or less
hoof-like.) Family Cuniculidae
Zygomatic region not abnormal, always without bony cheek-plates.
Cheekteeth evergrowing, or extremely hypsodont, the pattern one of a
series of transverse plates.
Mandible with angular process strongly distorted outwards ; incisors
powerful; form heavy, limbs not lengthened; hindfoot with
four well-developed digits; no tendency present for exces-
sive enlargement of bullae and mastoids nor for lengthening
of paroccipital process. Family Dino.myidae
Mandible with angular process rather weakly distorted outwards;
incisors relatively thin or medium; form more slender,
limbs to a degree lengthened; hindfoot with three well-
developed digits; a marked tendency present either to
excessive enlargement of bullae and mastoids, or to
lengthening of paroccipital process. Family Chinchillidae
Cheekteeth when evergrowing or extremely hypsodont never with
pattern of a series of transverse plates.
Hindfeet excessivelv specialized for arboreal life, or becoming so;
the hallux being replaced by a broad movable pad. (Fur
conspicuously spinous, the spines short; bullae rather large,
prominent; cheekteeth either with very wide re-entrant
folds, or nearly laminate in structure, rooted; no lengthening
of paroccipital process; zygoma simple.)
Familv ERETHIZONTIDAE
loo HYSTRICOIDAE
llindteet never excessively modified for arboreal lite, hallux never
replaced by broad movable pad, and never suppressed in
arboreal genera.
External form modified for cursorial life, the digits of the hind-
foot reduced to three, the claws thick, more or less hoof-
like. (Cheekteeth semi-rooted, the re-entrant folds isola-
ting as narrow islands on crown surface in adult; fur not
developing spines.) Family D.\syproctid.\e
External form never modified for cursorial life; digits of hind-
foot never reduced to three; claws not hoof-like in struc-
ture.
Externally showing a progressive series of modification of fur
into spiny covering, at extreme development highly
specialized (the spines long); bullae relatively small;
paroccipital process not specially lengthened nor
modified; zygoma simple. (Cheekteeth semi-rooted
or rooted, the re-entrant folds isolating as narrow
islands on crown surface in adult; a tendency present
towards extreme inflation of nasals in progressive
species; form heavy, terrestrial.) Family Hystricidae
Externally without extreme modifications (in one case to a
degree specialized for aquatic life); spiny covering
of body when present relatively weak (as compared
w ith Hystricidae) ; bullae relatively large ; paroccipital
process enlarged, either curving forward under the
bullae or lengthened, tending to stand apart from
them; zygoma very generally with upwardly directed
process on posterior border, or downwanilv directed
process on posterior border, or both. (Cheekteeth
various; sometimes evergrowing, when evergrowing
most often approaching or reaching complete simpli-
fication of pattern ; w^hen with a pattern of islands
isolating on crown surface in adult, usually brachv-
odont.) Family I'Iciiimyidae
These families it will be noticed are based chiefly on the external characters.
So many fossils are known belonging to this group that care must be taken if
defining the families on cranial and dental characters, as it may be that in many
cases fossil forms will prove intermediate between certain groups, or break
these characters down. Also the cranial characters of these Flystricoids are,
generally speaking, so similar that if not known living, all except Ciiiiiculus could
readily be referred to one family. The extreme external specializations reached
by some members of this group are in my opinion just as important as any
cranial or dental character.
ECHIMYIDAE loi
Family ECHIMYIDAE
i8q6. Thomas: Hystricomorfha, part, Family Octodontidae, part (included Cteno-
dactylidae) ; subfamilies Echimyinae (included Dactylomyinae), Capromyinae
{Capromys, Plagiodontia, Myocaslor, Thryonomys), Octodontinae (included
Abrocoma), Ctenodactylinae, part {Petromus).
1899. Tullberg: Hystricomorfha, part, Family Echinomyidae ; subfamily "Myopo-
tamini" (= Myocastorinae) ; subfamily Echinomyini (groups, Echinomyes,
Echimyinae and Dactylomyinae as here understood) ; and Octodontes (Octo-
dontinae and Abrocominae as here understood). Family Aulacodidae ( = Thryo-
nomyidae). Family Pctromyidae.
igi8. Miller & Gidley. Superfamily Hystricoidae, part; Family Echimyidae, part
(included Chaetomys); Subfamily Echimyinae ("Spiny-Rats, Hutias," etc.);
subfamily Octodontinae. Family Petromyidae. Family Myocastoridae. Family
Thryonomyidae. Family Abrocomidae.
1924. Winge: Family Hystricidae, part; Capromyini (Capromys, Plagiodontia,
Myocastor, Thryonomys); Octodontini, groups Octodontes (Octodontinae and
Abrocominae as here understood), Echinomyes (Dactylomyinae and Echimyinae as
here understood); Ctenodactylini, part (Petromus).
1928. Weber: Hystricoidea, part. Family Capromyidae (Capromys, Plagiodontia,
Myocastor); Family Octodontidae (Ecliimys, Octodon, Ctenomys); Family Cteno-
dactylidae, part (Petromus) ; Family Thr\onomyidae.
Geographic.\l Distribution. — Neotropical region, from Nicaragua south-
wards to Patagonia; Cuba and the West
Indies; Africa widel)- distributed south of the Sahara.
NiMBER OF Genera. — As here understood the family contains twenty-eight
genera, one of which, Procapromys, has not been
examined and is not represented in London.
Char.\cters. — Zygomasseteric structure typically Hystricoid in formation.
Cheekteeth when evergrowing never a series of transverse
plates (compare Chinchillidae, Dinomyidae); feet never abnormally modified
for arboreal life (compare Erethizontidae); zygomatic region without bony
cheek-plate (compare Cuniculidae); external form never modified for cursorial
life, digits of hindfoot more than three (always five except four in Thr\onom\s)
(compare Dasyproctidae); bullae prominent, and paroccipital process length-
ened, and zvgoma usually more angular, also tail never with specialized quills
or bristles in spiny genera, and spinv covering when present usually not highly
developed (compare Hystricidae).
As thus defined the group includes the great central mass of genera of Hystri-
coid rodents which have not become abnormally specialized in any external
particulars. The cheekteeth may be evergrowing (.\brocominae, Octodontinae,
Plagiodontinae, Capromvinae), extremely hypsodont (Myocastorinae, Petro-
myinae), or moderately so but rooted (the remainder). In the Octodontinae, the
structure approaches complete simplification of pattern; in the Abrocominae
the upper cheekteeth are simplified, but the lower series remains complex;
certain simplification has taken place in Plagiodontinae, which appears unique
as regards dental characters, and in Petromyinae. These subfamilies have
one external fold only in the upper cheekteeth; all other subfamilies have
102 ECHIMYIDAE
more than one, typically three. The skull is normally remarkable for the par-
occipital processes, which may be extremely lengthened', as in Myocastorinae,
moderately so and standing apart from the bullae (Caprominyae, Plagiodontinae,
Thryonomvinae), or curved forwards to a greater or lesser degree under the
bullae (Echimvinae, Petromvinae, Abrocominae, Octodontinae). This fact has
led some authors to form two families, Capromyidae and Octodontidae, but in
the Dactylomyinae, as proved by British Museum material, either condition may
exist; Kannabateomxs and Tlirinacudus, and some specimens of Dactxlomxs agree
with the Echimyinae, but some large skulls of Dactxlomxs are quite indistinguish-
able in paroccipital structure from Capromyinae, in which group there is also
some variation apparently.
The zygoma is usually rather broad, and frequently of a complex type, with
a downwardly directed process on posterior lower border, and sometimes an
upwardly directed one present above also.
The bullae may be much inflated, as in Abrocominae, Octodontinae,
Petromvinae; they are relatively large as a general rule.
The external form varies; in Myocastorinae, which contain relatively very
large forms, it is modified for aquatic life, with enlarged hindfeet, most of the
digits of which are webbed; this group has also bullae, which recall the type
found in Castoridae, though less specialized than in that family. Elsewhere
the genera are not aquatic. Some genera of Octodontinae are remarkable as
being the only Hystricoids which have taken to a subfossorial life {Ctenomys,
Spalacopiis, Acouaemxs); Spalacopus must be about the smallest living Hystri-
coid genus. A tendency to develop spiny covering, most pronounced in
Mesomxs, Lonc/iothrix and Hoplomxs, is present in some of the Echimvinae; the
spiny covering is, broadly speaking, very rudimentary compared with Hystri-
cidae ( Tn'clixs perhaps excepted), and even Erethizontidae.
The Dactylomyinae present a curious feature in that except in Tluiniicodiis the
third and fourth digits of fore- and hindfeet are much elongated; these animals
are said to be arboreal in habit. In all the genera of the family four functional
digits in manus and five functional digits in pes are present excepting Thryono-
myinae, in which the manus has three main digits only, D.5 being excessively
shortened, though bearing a thick claw, and the hallux is entirely suppressed;
this group possesses an abnormally massive and heavily ridged skull, perhaps
more so than in any other living rodent, and extremely thickened three-grooved
upper incisors. The form is heavy, and of terrestrial or slightly fossorial type.
I provisionally divide the group into nme subfamilies, most of which have
at some time or other been regarded as distinct families. But if a vast group like
the Muridae are kept together as one family, and the Hystricoid branch is
divided up into about seventeen families, the classification of the Order does not
appear very consistent. The simple-toothed Octodontinae are connected with
the main branch by such forms as Abrocoma and Plai;iodonlia\ the African
genera appear to possess no essential characters which will keep them apart as
families distinct. Petromus has of late been associated with the Ctenodactylidae
by some authors, but the typical Hystricoid mandible of Petromus differs very
widely from that of Ctenodactylidae, which are fully discussed elsewhere and
ECHIMYIDAE 103
which are here considered not related to the Hystricoid group. Thryonomys is
undoubtedly a very distinct type, but the only character which seems valid to
keep it apart as a family is the formation of the shoulder-blade, which, accord-
ing to Tullberg, differs considerably from that of other Echimyidae; but un-
fortunately this character cannot be examined throughout the genera at the
British Museum, Petromus, for one (one of the most important genera), being
not represented so far as this character is concerned; so that until the skeleton
of all the genera here included can be examined it seems more reasonable not
to separate any group on this structure alone. The digits of Thryonomys are
reduced, but this is an acquired character.
Myocastor, which is sometimes made the type of a family, is undoubtedly
a highly specialized form, but aquatic specialization alone is not sufficient to
base family groups on unless accompanied by some definite cranial or dental
characters (compare, for instance, other families, many of which have aquatic
offshoots beside normal generalized types, as Hydromyinae, Cricetinae, etc.);
and the structure of the paroccipital process, though highly specialized in
Myocastor, is too variable elsewhere in the group.
Miller & Gidley divided the Hystricoid lateralis series (^Hystricoidae as
here understood) into two groups based on the lachrymal canal, which is stated
to be closed in one branch, but open in the other branch on the side of the ros-
trum. Abrocoma as thus defined comes in a different group from the remainder
of those here. But this character seems not too constant elsewhere, for instance
both conditions are to be found in the Caviidae of Miller & Gidley, and I do
not attach too great importance to this character. The family as here under-
stood in fact is the family Octodontidae of Flower & Lydekker, and earlier
authors, except that of course the Ctenodactylidae are removed.
The subfamilies must be regarded as provisional; though easily recognizable
in living genera, it may be that among the large number of neotropical Hvstri-
coid fossil rodents some forms would be found which are intermediate between
some of the groups in the characters here noted.
Key to the Subf.^milies of Echimyidae
Cheekteeth becoming strongly simplified, the outer side of the upper
series with not more than one re-entrant fold.
The lower cheekteeth prismatic and angular in appearance; the upper
series eight-shaped. Part of lachrymal canal open on side of
rostrum. (Cheekteeth evergrowing; bullae much inflated;
zygoma more or less simple; form terrestrial; digits of hindfoot
five; skull not heavily ridged for muscle attachment.)
Subfamily Abrocomin.^e
[Abrocoma)
The lower cheekteeth not essentially different in pattern from the
upper scries; no part ot lachrymal canal open on side of rostrum.
Cheekteeth rooted, the pattern ultimately wearing out; inner side
of upper and outer side of lower teeth elevated. (Bullae
I04 ECHIMYIDAE
much inflated, the paroccipital process joining them; digits
of hindtoot five; zygoma more or less simple; skull not
heavily ridged for muscle attachment; form terrestrial, tail
fully haired.) Subfamily Petromyinae
(Petronnis)
Cheekteeth evergrowing, the pattern not or scarcely changing
during lite. Inner side of upper and outer side of lower
teeth not elevated.
Folds of cheekteeth very deep and long, set obliquely; each
upper tooth with a well-marked outwardly pointing
projection on external side. (Paroccipital process length-
ened, standing apart from bullae, which arc not extremely
inflated; zygoma simple; form generalized, tail naked;
skull not heavily ridged for muscle attachment.)
Subfamily Plagiodontinae
{Plagiodontia)
Folds of cheekteeth not specially deep and long, set less
obliquely, or not so. Upper cheekteeth either eight-
shaped, or "kidney-shaped." (Bullae normally much
inflated, the paroccipital process curved forwards under
them to a greater or lesser degree, and joining them;
zygoma complex, always with an upwardly pointing
process on posterior border; skull normally not heavily
ridged for muscle attachment (sagittal crest if present
weak); external form terrestrial, or modified for sub-
fossorial life; digits of hindfoot five.)
Subfamily Octodontinae
(Oitoinvs, Octoilontoiiivs, Ociodon, Acoiiaemvs ;
SpdlcHopiis ; Ctenoinvs)
Cheekteeth less simplified, the outer side of the upper molars with
at least two, typically three, re-entrant folds.
Externally considerably modified for acjuatic life (bodily size
largest of family); paroccipital process much elongated;
cheekteeth extremely hypsodont, but not evergrowing, with
well-marked re-entrant folds which are long retained, the
teeth decreasing in size markedly from M.3 forwards; skull
considerably ridged for muscle attachment (palate con-
stricted anteriorly; digits of hindtoot five; zygoma complex).
Subfamily Myocastorinae
(Mvocdstor)
External form never modified for aquatic life; paroccipital process
less or not elongated; cheekteeth less or not decreasing in
size from M.3 forwards.
ECHIMVIUAE 105
Skull massive, abnormally heavily ridged for muscle attach-
ment; incisors thick, the upper ones heavily three-
grooved; forefoot with three functional digits (D.5
clawed but extremely shortened); hindfoot with four
digits only; cheekteeth rooted; external form heavy,
terrestrial-tossorial (bullae not much inflated, paroccipital
process lengthened to a degree, and standing apart
from bullae; zygoma much broadened, jugal nearly
reaching lachrymal). (Shoulder-blade differing from
American genera (Tullberg).) [Subfamily Thryonomyinae
( Tliryonomys)
Skull much less heavily ridged for muscle attachment ; incisors
not three-grooved (plain except in Carterodon); forefoot
ahvays with four and hindfoot always with five functional
digits; jugal usually complex, with upwardly or down-
wardly projecting process present on posterior border,
and normally not approaching lachrymal.
Cheekteeth extremelv broadened, the re-entrant folds deep
and persistent, the teeth rooted; palate with a ten-
dency towards anterior constriction; paroccipital
processes tending to stand apart from bullae in larger
species, or curved forwards under them in smaller
forms; tendency present towards considerable elong-
ation of middle digits of forefoot and hindfoot; (fur
not spiny). Subfamily Dactylomyinae
{Thrinacodus, Dactylomys, Kannabateomys)
Cheekteeth not speciallv broadened; no tendency present
towards any lengthening of the digits.
Cheekteeth evergrowing, the folds filled up with
cement, the teeth llatcrowned; fur not developing
spines; paroccipital processes usually standing
apart from the bullae (not always); palate
slightly constricted anteriorly; form usually not
rat-like (modified for arboreal or terrestrial
life). Subfamily Capromyin'ae
(Procapromys (not seen), Capromys, Geocapromys)
Cheekteeth rooted, the folds normally isolating as
narrow islands in adult, or rarely {Echimys and
allies) more persistent, the dental pattern more
complex, or in extreme development becoming
a series of transverse plates. External form
usually rat-like; modified for arboreal, terres-
trial, or slightly for fossorial life; a tendency
present for the fur to be spiny; paroccipital
io6 ECHIMYIDAE: ECHIMYINAE
processes curved forwards under the bullae, which
are large but not abnormally so, excepting
CIvoinys; palate not constricted anteriorly.
Subfamily Echimyinae
(Efhtinys, ho/hrix, Diploniys ; Proechimvs,
Hdplomxs, Eiirvivgoiiiat<»iiYS, (Ivomys, Cartcr-
odon, Cercomvs, Mesomvs, Lonchothrix)
Subfamily ECHIMYINAE
Geographical Distribution. — Neotropical region from Nicaragua south
to Paraguay and South Brazil.
Number of Genera. — Eleven.
Char.\cters. — Cheekteeth rooted, not specially broadened, flatcrowned
with inner and outer re-entrant folds which become isolated
on crown surface with wear; or with a heavier dentition, more complex, and
tending to become a series of transverse plates (Echiinys, Isotlirix, Diplomys).
Bullae prominent ; paroccipital processes curved forward under the bullae. Skull
with broad frontals, little or not constricted. Externally more or less rat-like;
a tendency present for development of spiny covering, which is in rare cases
strongly developed.
Compared with the Capromvinae, the cheekteeth are brachyodont and rooted,
and of a less simple general appearance, the pattern changing during the animal's
life; compared with the Dactylomyinae, there is no broadening of the cheek-
teeth, which never show traces of the almost prismatic pattern peculiar to
Dactylomyinae; and no digit elongation takes place; compared with the Myo-
castorinae, cheekteeth more brachyodont, and pattern not long maintained;
also externally not modified for aquatic life; skull much less heavily ridged, size
smaller, form much less heavy, and paroccipital processes not lengthened; com-
pared with Thryonomyinae, skull much less heavily ridged; forefoot with four
and hindfoot with five clearly developed digits.
In the remainder of the subfamilies the cheekteeth are more simplified.
There seems in this group a tendency for the tail to be lost during life,
paralleled by the Old World porcupine Trichys; in the case of Trichys it is sug-
gested that the males may bite oft the tail of the female when courting; but
whether this might apply to this group is not known.
The subfamily contains two sections, in one of which the teeth are much
heavier, more complex, the folds usually not isolating as islands; in extreme
development the teeth become a series of transverse plates. The genera
contained in this section have the feet modified tor arboreal life.
EcHlMYS is the main genus, with five or more rather well-defined specific
groups, and a wide range in South America ; a species, E. arimitiis, is said to occur
in the Lesser Antilles (Martinique). The fur may be strongly spinous, or weakly
so; IsoTHRix appears to be indistinguishable cranially or dentally from Echiniys,
but has soft fur, and a bushy tail; Diplomys is a (chiefly) Central American
ECHIMYINAE 107
genus in which hoth lower and upper molars are a series of transverse plates,
though in a section (jf Kchini\s the upper molars arc already so, so that Diplomys
is not widely separated from Echimvs.
The simpler-toothed branch of the subfamily, in which the dentition is
lighter, and the folds isolate as islands in the adult, contains eight genera;
I'ROEClllMYS is much the most widely distributed, having a range coincident with
with that of the subfamily, and very many named forms; this genus has spiny
fur in adult, though not highly developed, and is terrestrial; Hoplomvs stands
near Proechimys, but the spiny covering is very much more developed, and the
cheekteeth are more complex; the genus ranges from Nicaragua to Ecuador.
EuRVZYGOMATOMYS is like Proechimys, but is more modified for fossorial life
(though not highly so); the tail is strongly reduced; the cheekteeth are more
simplified than in normal Proechimys, and the skull is less ridged posteriorly,
but the zygoma is greatly broadened. Clyomys is near Euryzygomatomys,
but alone in the group has abnormally inflated bullae. Carterodon is not
unlike the last two named in cranial characters, but has deeply grooved incisors
(unique in the subfamily), and softer fur. These three genera are rare and have
a restricted range in Brazil, the last two being very little known. Cercomy'S
agrees with the last named in essential cranial and dental characters, but is not
in any way modified for fossorial life; the tail is long and fully haired, the fur
soft. Mesomys is an arboreal type ranging across the tropical portions of north-
ern South America; the fur is densely spiny, the teeth are of the Proechimys
type. Finally Lonchothrix, rare and little known, is much like Mesomys
externally except for the heavily tufted tail, but dentally differs from all in the
considerable width of the re-entrant folds of the molars.
Key to the Genera of Echimyinae
Cheekteeth lighter, with narrow folds, which typically become isolated
as islands in adult; the teeth never tending to become a series of
transverse plates.
The hindfcet broadened, noticeably modified for arboreal life. (Fur
densely spiny; D.5 hindfoot relatively long; tail long, usually
longer than head and body.)
Cheekteeth with strong wide folds; tail conspicuously tufted
terminally. LoNCHOTHRIX
Cheekteeth with weaker narrower folds; tail not or scarcely tufted
terminally. Mesomy'S
The hindfeet narrow, long, terrestrial in type. (D.5 hindfoot relatively
short; tail in general shorter than head and body.)
Jugal thickened anteriorlv; zygomatic region noticeably broadened,
Tail strongly reduced.
Upper incisors grooved ; fur soft ; foreclaws less enlarged. Carterodon
Upper incisors plain; fur bristly; foreclaws to a greater or lesser
degree enlarged.
io8 ECHIMYINAE: ECHIMYS
Bullae abnormally inflated. Clyomys
Bullae moderate. Euryzygomatomys
Jugal not or rarely thickened anteriorly, zygomatic region narrower.
Tail not strongly reduced. (Upper incisors plain; bullae
never abnormally inflated.)
Fur soft; tail well haired; palatal foramina broadened; folds of
cheekteeth tend less to isolate. Cercomys
Fur spinv; tail poorly haired; palatal foramina not noticeably
broadened; folds of cheekteeth tend to isolate to a greater
degree.
Modification of hair into spines at highest development;
cheekteeth more complex, outer side of upper cheek-
teeth with clear traces of four folds. Hoplomys
Modification of hair into spines much less developed ; cheek-
teeth relatively simpler, outer side of upper cheek-
teeth with clear trace of three folds or less. Proixhimys
Cheekteeth heavier, with more persistent folds, the general effect comple.x;
the folds isolating late, or not isolating; or cheekteeth tending to
become a series of transverse plates. (Feet adapted for arboreal life.)
The lower molars a series of transverse plates, as well as the upper
series. Diplomys
The lower molars not becoming specialized into a series of transverse
plates; (the upper cheekteeth may or may not be so).
Fur bristly or spiny; tail not bushy. EcHIMYS
Fur soft; tail bushy, Isothrix
(k-nus I. I'X'HIMYS, Cuvier
1S09. EcHiMVS, Cuvier, Bull. Soc. Philom. XXIV. p. 3134.
181 1. LoNCHERES, Illigcr, Prodr. Syst. Mamm. et Avium, p. 90. (Myoxiis clirysurns,
Zimmermanii).
1837. Nelomys, Cuvier, Ann. Sci. Nat. Paris, 2, VIII, p. 370. (Nelomys blaiiivitlei, Cuvier).
1841. Phvllomvs, Lund, ,Afh. K. Danske Vid. Selsk, 4, VIII, p. 243. (FhyUoiiiys
brazilicnsis, Waterhouse.)
1840. EcHINOMYS, Wagner, .Abb. Bayer. Akad. 3, p. 203. (Emendation of Eciiimys.)
Type Species. — Myoxus chrysurus, Zimmermann.
Range. — Neotropical; Colombia, Ecuador, Peru, Southern Brazil, Eastern
Brazil, Amazon region, the Guianas, Venezuela. E. armatus is
recorded from Martinique, Lesser Antilles.
Number of F'orms. — Twenty-two are named.
Characters. — Skull with broad frontals, and as a rule well marked supra-
orbital ridges; in larger species the parietals are well ridged.
Infraorbital foramen with no canal for transmission of nerve. Bullae prominent,
the paroccipitals curved forward under them. Palate varying in width in
ECHIMYS 109
different species, but tends to be narrow. Palatal foramina normally short.
Zygoma relatively broad, the jugal usually with process both above and below
on posterior border. Mandible well twisted and ridged.
Cheekteeth complex; apparently usually there are two outer and two inner
folds in the upper series, and there is a strong tendency for these teeth to be
divided into a series of transverse plates. They are completely so in some
Southern Brazilian types as mediits, thumasi, lamarum, dasythrix, etc., for which
Thomas revived the generic name Nelomys. Tate synonymizes this with
Echimys; some of the northern species come so near this formation that I do
not think the name can be used. The lower cheekteeth are characterized by one
outer and two inner folds, except the premolar, a complex tooth with three or
sometimes four inner folds; the folds of the lower teeth are usually rather well
open.
Externally, the feet are broad and modified for arboreal life, with D.5
relatively long, and claws prominent; the fur is always spiny, though the spines
vary in their development in different groups, being in some very strong, in
others relatively weak. The length of the tail is little shorter than head and
body to longer than this measurement; it may be well haired or nearly naked.
Not very much material of this interesting genus is available for examina-
tion, but the forms seen seem to divide sharply and easily into five main groups.
These groups should be regarded, however, as provisional, as it may be with
more material that some of the characters would not be constant.
The blaiinillei group as here understood is equivalent to part of the genus
Nelomys of Thomas; containing species in which the upper molars are appar-
ently completely specialized into transverse plates; the tail is longer than the
head and body, and is more or less completely haired; blainvillei has the spines
strongly developed in the one skin seen; medius has the spines weakly developed;
thomasi is like medius but much larger (hindfoot 45 as against about 38 or less
in allies). E. braziliensis is listed as "hairy-tailed" by Tate, and is provisionally
included here, as it was based on the genus " PhvUomys" which is considered
by Thomas synonymous with Xelomys.
The dasythrix group contains the rest of the genus Nelomys of Thomas; it
is closely allied to the last, and from the same area (Southern Brazil), but the
tail appears to be not longer than the head and body, and is naked. The spines
are strong. It may be that blainvillei might be considered an intermediate form
between these two groups.
The armatus groups contains forms in which the upper teeth are normally
not separated into transverse plates; the tail is naked, or in punctatus rather
intermediate between the "hairy-tailed" and "naked-tailed" types. The
spines are strong, well developed. E. carrikeri, not seen, appears to stand near
punctatus from the description ; longirostris is stated to be near armatus, as is
obscura, according to Tate. I can see no specific difference between armatus
and occasius, and treat the latter as a subspecies. £. y?aM(yi« is described as an
insular form of punctatus.
The clirysurus group contains two striking forms; the tail is longer than
the head and body, coloured white from about half its length or more to the
ECHIMYS
end; a white headspot present; spiny covering strong to extreme (at maximum
for the genus). The tail is well haired ; the dentition agrees with arniatus.
Fig. 8. Echimys armatus armatvs, GeofFroy.
B.M. No. 3.4.5.40, ?; ,■ li.
Fig. 9. Echimys armatus armatus, GeofFroy.
B.M. No. 3.4.5.40, 9; > I J.
E. satiirmis, a giant species (hindfoot 51), was thought by Thomas to be near
the above group; it is evidently known by one skin only. I am inclined for now
to refer it to xht grandis group (below), on account of the much less development
ECHIMYS 1 1 i
of the spiny covering, which is relatively weak. There is no white headspot.
The tail is only white at the extreme tip. With a larger series it may be that
saturnus could be shown to belong to the chrysurus group, but with the limited
material at present at hand there is no doubt that it is very different from that
group. (The tail is long and fully haired.)
The grundis group contains western forms with tail sub-equal to or shorter
than head and body, not white terminally, and well haired. Spiny covering
weak or weak-medium. E. grandis is a very large form (hindfoot 53); dentition
usually as in armatus group.
\^^^
a b
Fig. 10. EcHiMVs armatus armatus, Geoffroy.
Cheekteeth: a, upper; b, lower. B.M. No. 3.4.5.40, ?; X 7.
The spines may vary through the animal's life, and be an age character
throughout this family, within closely related species; but I do not think there
is any question of such a difference between groups listed here as "strong-
spined" or " weak-spined." In adult grandis, for instance, the spines are very
weak; in adult chrysurus abnormally strong.
Other species not represented at the British Museum I am unable to allocate
to groups.
Forms seen; armatus, blaimillei, braziliensis, chrysurus, dasythrix, "gutanae,"
grandis, lamarum, medius, occasius,paleacea,punctatus, rhipidurus, saturnus, thomasi.
List of N'.\.med Forms
(The references and type localities for all members of the family Echimyidae
have been collected for me bv Mr. G. W. C. Holt.)
,12 ECHIMYS
Not seen uiitl not allocated to t^roups
1. I-;CH1MVS SEMIVILLOSUS, Gtoffroy
1838. Revue Zool. i, p. lOi.
New Grenada, Colonibia.
2. ECHIMYS MACRL'RA, Wagner
1842. Archiv. fur. Naturg. i, p. 360.
Borba, Rio Madeira, Brazil.
3. ECHIMYS UNICOLOR, Wanner
1842. Archiv. filr. Naturg. i, p. 361.
Brazil.
4. ECHIMYS NIGRISPINA, Wagner
1842. Archiv. fiir. Naturg. i, p. 361.
Ypanema, Sao Paulo, Brazil.
dasythrix Group
5. ECHIMYS DASYTHRIX. Ilensel
1872. Abh. Akad. Berlin, p. 49.
Rio Grande do Sul, Brazil.
6. ECHIMYS LAMARUM, Thomas
1(316. Ann. Mag. Nat. Hi.st. 8, XVIII, p. 297.
Lamarao, Bahia, Brazil.
blainvillei Group
7. ECHIMYS BLAINVILLEI, Cuvrer
1837. Ann. Sci. Nat. Paris, 2, VIII, p. 371.
Small Island near Bahia, Brazil.
S. ECHIMYS MEDIUS, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. 239.
Roca Nova, Parana, Brazil.
9. ECHIMYS THOMASI. Ihirint:
1S97. Revista Paulista, ii, p. 171.
Island of Sao Sebastiao, near Bahia, Brazil.
10. IXHIMYS BRAZILIENSIS. Waterhnuse
1848. Nat. Hist. Mamni. ii, p. 330.
Lagoa Santa, Mmas Geraes, Brazil.
armatus Group
11. IX'HIMYS ARMATUS ARMATCS, Geoffroy
1838. Revue Zooliigique, i, p. loi.
Cavenne, I'rench Guiana.
Synonym: guiamie, Thomas, 1888, Ann. Mag. Nat. Hist. 6, II, p. 326.
British Guiana.
castaneiis, Allen & Chapman, 1893, Bull. Amer. Mus. N. H.
V, p. 222. Princetown, Trinidad.
12. ECHIMYS ARMATL'S OCCASIUS, Thomas
1921. Ann. Mag. Nat. Hist. 9, VII, p. 450.
Gualea, Mt. Pichincha. Ecuador. (This locality is queried by Tate, I935-)
ECHIMYS— ISOTHRIX 113
.3. IXHIMYS LONGIROSTRIS, Anthony
1921. Amer. Mus. Nov. no. 19, p. 5.
Kartabo, Uritish Guiana.
14. IXHIMY.S ()BSCUR.'\, Wanner
1840. Abh. Akad. Wiss. Munch, iii, p. 196.
Brazil.
15. KCHIMYS PUNCTATUS, Thomas
1899. Ann. Mag. Nat. Hist. 7, III, p. 153.
Caicara, Rio Orinoco, Venezuela.
16. ECHIMYS KL.WIDUS, Hollister
1914. Proc. Biol. Soc. Washington, XXVII, p. 143.
El Valle, Margarita Island, Venezuela.
17. ECHIMYS C.-XHRIKERI, Allen
191 1. Bull. Amer, Mus, Nat. Hist. XXX, p. 251.
San Esteban, near Venezuela.
chrysurus Group
18. ECHIMYS CHRYSURUS, Zimmermann
1780. Geogr. Gesch. ii, p. 352.
Surinam (Dutch Guiana).
Synonym: cristatus, Desmarest, 1817, Nouv. Diet. d'Hist. Nat.
2nd ed., X, p. 55.
19. ECHIMYS PALEACEA, Illiger
181 1. Prodr, Syst, Mamm. et Avium, nom. nud. 1820, Lichtenstein, Abh. Ak. Wis.
Berlin, p. 187.
Province of Para, Brazil.
grandis Group
{satiirnus section)
20. ECHIMYS SATURNUS, Thomas
1928. Ann. Mag. Nat. Hist. 10, II, p. 409.
Ecuador, Rio Napo, Prov. del Oriente.
(typical section)
21. ECHIMYS GRANDIS, Wagner
1845. Archiv. fiir Naturg. i, p. 146.
Managueri, Upper Amazon, Brazil.
22. ECHIMYS RHIPIUURUS, Thomas
1928. Ann. Mag. Nat. Hist. 10, II, p. 291.
Pebas, Rio Maranon, Peru.
Genus 2. ISOTHRIX, Wagner
1845. IsoTHRlx, Wagner, .^rchiv. fiir. Naturg. i, p. 145.
1852. Lasiiromys, Deville, Rev. Mag. Zool. 2, IV, p. 353. {Lasiuromys villosus,
Deville.)
Type Species. — hothrix histriatus, Wagner.
Range. — Venezuela, Brazil and Peru. (South evidently to Matto Grosso.)
Number of Forms. — Eight are named.
8 — LivinR Rodents — I
114 ISOTHRIX
Characters. — Like luhiinvs cranially and dentally; (parietals ridged;
upper cheekteeth nut tending to become separated into
transverse plates).
Fur soft, showing no tendency to develop bristles or spines. Tail long,
bushy, almost Sciurine. Feet of arboreal type.
Remarks. — In a group of this description where considerable specialization
is sometimes present towards modification of fur into spines,
I think the difl'erence in the coat between Echimys and hot/nix is sufficient for
their generic separation.
Forms seen : mol/iae, iiegreiisis, oriiioci, pagurus, pictiis, rillosiis.
Two groups may be recognized among the material examined, pictiis, with
its highly specialized black (or dark brown) and white cok)ur pattern, and the
rest which are much more sober in coloration.
Mr. Tate stated that pictiis is an Echimys; but it definitely belongs here
according to our specimens; some months ago when he was in London we
looked at the species together, and he was in agreement with me on this point.
List of Named Forms
pictiis Group
1. ISOTHRIX PICTUS, Pictet
1 841. Notice An. Nouv. Mus. Geneve, p. zg.
Bahia, Brazil.
bistriatiis Group
2. ISOTHRIX BISTRI.'^TUS BISTRl.^TUS, Wagner
1845. Arch. Naturg. i, p. 146.
Rio Guapore, Brazil.
3. ISOTHRIX BISTRIATUS ORINOCI, Thomas
1899. Ann. Mag. Nat. Hist. 7, IV, p. 382.
Maipures, Upper Orinoco, Venezuela.
4. ISOTHRIX BISTRIATUS NEGRENSIS, Thomas
1920. Ann. Mag. Nat. Hist. 9, VI, p. 277.
Acajutuba, Lower Rio Negro, Brazil.
5. ISOTHRIX PACHYURA, Wagner
1845. Archiv. fiir Naturg. i, p. 146.
Cuyaba, Matto Grosso, Brazil.
Synonym: crassicaudiis, Wagner, Abli. .\kad. Miinch., p. 291, 1847.
Brazil.
6. ISOTHRIX PAGURLIS, Wagner
1S45. Archiv. fiir Naturg, i, p. 146.
Borba, Rio Madeira, Brazil.
7. ISOTHRIX VILLOSUS VILLOSUS, Deville
1852. Rev. Mag. Zool. 2, IV, p. 560.
Mission de Sarayacu, Rio Urubamba, Peru.
8. ISOTHRIX VILLOSUS M( )LLIAE, Thomas
1924. Ann. Mag. Nat. Hist. 9, XIII, p. 534.
Tushemo, Masisea, Rio Ucayali, Peru.
Tiie "hirsiitiis" of Burmeister which has been confused with this genus is
a Siginodon (Cricetinae).
DIPLOMYS— PROECHIMYS iiS
Genus 3. DIPLOMYS, Thomas
1916. DiPLOMYS, Thomas, Ann. Mag. Nat. Hist. 8, XVIII, p. 240.
Type Species. — Loncheres caniceps, Giinther.
Range. — Panama and Colombia.
Number of Forms. — Four.
Characters. — In general Hke Echimys, but the lower cheekteeth in a series
of transverse plates as well as the upper teeth. There are
four laminae in each of the upper teeth; in the lower series there are four in
P.4, three in the molars.
Externally, the fur is harsh but not spiny; tail moderately haired, but with
the scales visible ; feet of arboreal type.
Remarks. — The genus is not well represented in London; I assume the
dental characters to be constant.
Forms seen : caniceps, labilis.
List of Named Forms
1. DIPLOIVrV'S CANICEPS, Gunther
1876. Proc. Zool. Soc. London, p. 745.
Medellin, Colombia.
2. DIPLOMYS LABILIS, Bangs
1901. Amer. Naturalist, XXXV, p. 638.
San Miguel Island, Panama.
3. DIPLO.MYS D.\RLINGI, Goldman
1912. Smiths. Misc. Coll. LX, no. 2, p. 12.
Marraganti, Rio Tuyra, East Panama.
4. DIPLOMYS RUFODORSALIS, .'\llen
1899. Bull. Amer. Mus. Nat. Hist. XII, p. 197.
Onaca, Santa Marta district, Colombia.
Genus 4. PROECHIMYS, Allen
1899. PROECHIMYS, Allen, Bull. Amer. Mus. Nat. Hist., XII, p. 257.
1921. Trinomys, Thomas, .•\nn. Mag. Nat. Hist. 9, VIII, p. 140. Proechimys albi-
spiirns, GeofFroy. Valid as a subgenus.
Type Species. — Echimys trinitatis, Allen & Chapman.
Range. — From Nicaragua, Costa Rica and Panama to Colombia, Ecuador,
Peru, Bolivia, Southern Brazil, East Brazil, the Amazon region,
the Guianas, Venezuela, Trinidad.
Number of Forms. — Appro.ximately fifty are named.
Characters. — Rostrum relatively narrow and pointed; incisors typically
opisthodont; supraorbital ridges present and usually well
developed; parietals usually ridged. Canal for transmission of nerve in infra-
orbital foramen present, weak or absent, never strongly developed. Palatal
foramina well open, but not excessive; palate relatively broader as a rule than
in Echimys; toothrow rather short, far forward in skull, the pterygoid fossae
long. Jugal normally thin, but ridged posteriorly and tending to have a weak
I'ROECHIMYS
process on posterior lower border; thickened only, so far as seen, in iheringi
(considerably, but not as extremely as in Euiyzygomatomys), to a degree in the
subgenus Trinomys, and to a degree in dimidiatus. Bullae largish; paroccipital
Fig. II. Proechimys cayennensis, Desmarest.
B.M. No. 3.4.5-44. 9; X I J.
Fig. 12. Proechimys cayennensis, Desmarest.
B.M. No. 3-4.5-44. S; ' i|-
PROECHIMYS 117
processes curved forwards under them, as is usual in the subfamily. Mandible
strongly ridged, the angular process clearly lifted outwards, its lower border
broad; coronoid process low; angular portion slightly drawn backwards at
lower border.
Upper cheekteeth normally with three outer and one inner folds each,' these
soon becoming isolated as islands. A few species, which will be discussed below,
vary slightly in pattern. Lower cheekteeth normally reverse the pattern of the
Fig. 13. Proechimys cayennensis, Desmarest.
Cheekteeth: a, upper; b, lower. B.M. No. 3.4.5.44, $; x 8.
upper series. P. 4 sometimes with extra island anteriorly. Spines always present
in adult, but not highly developed, at any rate as compared with such genera as
Mesomxs and Hoplomys. Tail rather shorter than head and body, naked or
moderately haired. Hindfeet long and narrow, the outer digits shorter than the
central three, hallux shorter than D.5; forefoot not abnormal, pollex rudi-
mentar\\
Tri.nomys was erected as a subgenus for the species albispimis and setosus,
on the ground mainly that the folds of the cheekteeth are in these species
reduced from three (external) to two. (One skull seen in which the teeth are
cutting has a normal P. 4, however). The cheekteeth may varv slightly elsewhere
in the genus; P. lacillator has it appears M.2 and M.3 as Trinomrs; the type of
P. dimidiatus appears to be going the same way, though rather worn. One skull
seen of P. canicollis seems also to be transitionary towards the Trinomxs type.
It should also be noted that P. iheringi seems not quite normal dentally; the
teeth in this case appearing a little more complex than usual.
ii8 PROECHIMYS
Other characters of Trinomxs are cranial, "shorter muzzle, less developed
supraorbital ridges, and orthodont or slightly pro-odont incisors." Also the
spiny covering seems to be much more developed than in the other species.
Forms seen : albispiiuis, ho/iiiainis, brevicauda, calidior, cavciincnsis, canicollis,
centralis, cherriei, chiriquinus, chrysaeoliis, coloinbianiis, decumaniis, diinidiatus,
goeldii, gorgomie, guiarae, gidaris, hendeei, hilda, iheringi, longicaiidatus, mincae,
oris, pac/iitii, paruimensis, rattiinis, roberti, rosa, securiis, semispinostis, setostis,
sertotntis, sitnonsi, trinitatis, iirichi, vacillator, warreni.
Of the fifty named forms of this genus, twelve have not been seen.
Most of the remainder are standing at the present day as "distinct species."
A very large number, however, seem to conform to one essential type, so that
neither in external nor cranial characters, so far as I can see, are they more
than racially distinct.
P. caxeiiiieiisis, Desmarest, 1817, is the oldest name for this type of Spiny-
Rat. It is true that Tomes has pointed out characters by which his semispinosm
may be distinguished from cayeiieimsis; but these seem to me to be relatively
unimportant subspecific characters. All forms occurring north of Panama are
at present regarded as races of semispinosus.
The type and a large or moderate series of skins have been examined in
clirysaeolus, deciiiininiis, rosa, warreni, cherriei, gularis, brevicaiida, simonsi,
pdchita, hilda, bulivianus, securiis, oris and roberti, and a moderate or large series
of skins of mincae, guiarae, trinitatis, uriclii, gorgonae, longicaiidatus, and goeldii.
Not one of the above names seem to me to be retainable as full species.
There are slight colour variations present ; there are quite noticeable size
variations (but the smallest connected with the largest by intermediate forms);
there are variations in the length of the tail, though in animals of this kind,
which frequently lose the tail during life, this seems to be a character to which
too much attention should not be paid.
I propose provisionally to treat all the above-mentioned forms, and semi-
spinosus and its races, as subspecies of cayenuensis. Should this prove in any
case incorrect, it may be noted that numbers 14 to 20 are regarded now as semi-
spinosus and its races, and numbers 21 to 40 as "distinct species."
P. vacillator is kept separate on account of the dental characters noticed
above, though it must be admitted that on external characters alone it would
certainly be regarded as a race of cavennensis. P. canicollis differs in colour from
the above; hendeei and rattiinis are darker than usual, and with rather weak
spines; Thomas regarded these as forming a section of the genus; they are
accordingly kept apart as species. P. iheringi is kept apart as a group on account
of the characters of the zygoma. P. dimidiatus, of which one skull only has been
seen, might belong to that group, or to the typical one; until more specimens
come to hand the question must remain open. P. albispiniis differs from the
only other species referred to the subgenus Trinomys in colour, so far as seen.
The forms here referred as races to cavennensis are mostly supposed by
Thomas to be "species" on trifling skull characters, such as the absence of the
parietal ridges (age character .'), the length of palatal foramina, " narrow muzzle,"
etc.
PROECHIMYS 119
List or Named Forms
Subgenus Proecliimys, Allen
Not seen, and not allocated to group.
1. I'ROKCHIMYS OCHRACEUS, Osgood
191 2. Field Mus. Pub. Zool. Ser. X, p. 56.
El Panorama, Rio Aurare, Zulia, Venezuela.
2. PROECHIMYS MACROURUS, Jentink
1879. Notes Leyden Mus. i, note 23, p. 97.
Surinam.
3. PROECHIMYS O'CONNELLI, Allen
1913. Bull. Amer. Mus. Nat. Hist. XXXII, p. 479.
Villavicencio, Colombia.
4. PROECHIMYS POLIOPUS, Osgood
1914. Field Mus. Nat. Hist. Zool. ser. X, p. 141.
San Juan de Colon, Tachira, Venezuela.
5. PROECHIMYS STEEREI, Goldman
191 1. Proc. Biol. Soc. Washington, XXIV, p. 238.
Hyutanaham, Rio Purus, Brazil.
6. PROECHIMYS KERMITI, Allen
1915. Bull. Amer. Mus. Nat. Hist. XXXIV, p. 629.
Lower Solimoes, Brazil.
7. PROECHIMYS BOIMENSIS, Allen
1916. Bull. Amer. Mus. Nat. Hist. XXXV, p. 523.
Boim, Rio Tapajoz, Brazil.
8. PROECHIMYS ELEGANS, Lund
1841. Afh. K. Danske Vid. Selsk, 4, VIII, p. 245.
Lagoa Santa, Minas Geraes, Brazil.
9. PROECHIMYS LEUCOMYSTAX, Ribeiro
1914. Comm. Linhas. Telegr. Annex. 5, p. 43.
Utiarity, Rio Papagaio, Matto Grosso, Brazil.
10. PROECHIMYS MYOSUROS, Lichtenstein
1820. Abh. Akad. Wiss. Berlin (1818-1819), p. 192.
Bahia, Brazil.
11. PROECHIMYS LEPTOSOMA, Brants
1827. Muizen, p. 150.
Bahia and Sao Paulo, Brazil.
Synonym: cinnamomeus, Lichtenstein, 1830, Darstellung, pi. 36, fig. 2.
12. PROECHIMYS FULIGINOSUS, Wagner
1842. Schreber Saug. Suppl. Ill, p. 343.
Brazil.
From the descriptions, ochraceus (no. 1), o'connelli (no. 3), steerei (no. 5),
kermiti (no. 6), appear near caycnncnsis or perhaps races ; poliopus (no. 4) is
probably distinct, hoimcnsis (no. 7), is clearly distinct from others, and macrourus
(no. 2), is described as a form with an unusually long tail (head and body 221,
tail 320).
I20 PROECHIMYS
ca\ennensis Group
13. PRtJECHlMYS CAYENNENSIS CAYIiNNENSIS, Dcsmarcsl
1817. Nouv. Diet. d'Hist. Nat. 2d Ed. X, p. 59.
Guiana.
14. PROECHIMYS CAYENNENSIS SEMISPINOSUS, Tomes
i860. Proc. Zool. Soc. London, p. 265.
Gualaquiza, Eastern Ecuador.
15. PROECHIMYS C.JiYENNENSIS BURRUS, Bangs
1901. Amer. Naturalist, XXXV, p. 640.
San Migue! Island, Panama,
lb. PROECHIMYS CAYENNENSIS CENTRALIS, Thomas
1896. Ann. Mag. Nat. Hist. 6, XVIII, p. 312.
San Emilio, Lake Nicaragua, Nicaragua.
17. PROECHIMYS C.\YENNI:NSIS PANAMENSIS, Thomas
1900. .\nn. Mag. Nat. Hist. 7, V, p. 220.
City of Panama, Panama.
Synonym: centralis chiriquimis, Thomas, 1900, Ann. Mag. Nat. Hist. 7,
V, p. 220. Bogava, Chiriqui, Panama.
18. PROECHIMYS CAYENNENSIS RUBELLUS, HolHster
1914. Proc. Biol. Soc. Washington, XXVII, p. 57.
Angostura Valley, Costa Rica,
ig. PROECHIMYS CAYENNENSIS COLOMBIANUS, Thomas
1914. .Ann. Mag. Nat. Hist, 8, XIV, p. 60.
Condoto, Choco, Western Colombia.
20. PROECHIMYS CAYENNENSIS CALIDIOR. Thomas
1911. Ann. Mag. Nat. Hist. 8, VIII, p. 254.
San Javier, Lower Rio Cachavi, Ecuador.
21. PROECHIMYS CAYENNENSIS CHRYSAEOLUS, Thomas
1898. Ann. Mag. Nat. Hist. 7, I. P- 244-
Muzo, north of Bogota, Colombia.
22. PROECHIMYS CAYENNENSIS MINCAE. Allen
1899. Bull. Amer. Mus. Nat. Hist. XII, p. 198.
Minca, Santa Marta district, Colombia.
23. PROECHIMYS CAYENNENSIS GORGONAE, Bangs
1905. Bull. Mus. Comp. Zool. Harvard, 46, p. 89.
Gorgona Island, Colombia.
24. PROECHIMYS CAYENNENSIS DECUM.ANUS, Thomas
1899. Ann. Mag. Nat. Hist. 7, IV, p. 282.
Chongon, Prov. Guayas, Ecuador.
25. PROECHIMYS CAYENNENSIS ROSA, Thomas
1900. .Ann. Mag. Nat. Hist. 7, V, p. 219.
Santa Rosa, South-west Ecuador.
26. PROECHIMYS CAYENNENSIS GULARIS, Thomas
191 1. Ann. Mag. Nat. Hist. 8, VIII, p. 253.
Canelos, Rio Bobonaza, Ecuador.
27. PROECHIMYS CAYENNENSIS BREVICAUDA, Gunther
1876. Proc. Zool. Soc. London, p. 748.
Chamicuros, Huallaga River. Peru.
PROECHIMYS
28. PROECHIMYS CAYENNENSIS SIMONSI, Thomas
iQoo. Ann. Map. Nat. Hist. 7, VI, p. 300.
Perene River, Prov. Junin, Peru.
29. PROECHIMYS CAYENNKN.SIS P.-\CHITA, Thomas
1923. Ann. Mag. Nat. Hist. 9, XH, p. 694.
iPucrto Leguia, Rio Pachita, Peru.
30. PROECHIMYS CAYENNENSIS HILDA, Thomas
1924. Ann. Mag. Nat. Hist. 9, XIII, p. 534.
San Lorenzo, Rio Maranon, Peru.
31. PROECHIMYS CAYENNENSIS BOIJVl.-\NUS, Thomas
1901. Ann. Mag. Nat. Hist. 7, VIII, p. 537.
Mapiri, Upper Rio Beni, Bolivia.
32. PROECHIMYS CAYENNENSIS SECURUS, Thomas
1902. Ann. Mag. Nat. Hist. 7, IX, p. 140.
Charuplaya, Secure River, Bolivia.
33. PROECHIMYS CAYENNENSIS WARRENI, Thomas
1905. Ann. Mag. Nat. Hist. 7, XVI, p. 312.
Comaccka, Demerara River, British Guiana.
34. PROECHIMYS CAYEiNNENSlS GLAIRAE, Thomas
igoi. Proc. Biol, Soc. Washington, XIV, p. 27.
La Guaira, Venezuela.
35. PROECHIMYS CAYENNENSIS URICHI, Allen
1899. Bull. Amer. Mus. Nat. Hist. XII, p. 199.
Quebrada Seca, Prov. Sucre, Venezuela.
36. PROECHIMYS CAYENNENSIS CHERRIEI, Thomas
1899. Ann. Mag. Nat. Hist. 7, IV, p. 381.
Munduapo, Upper River Orinoco, Venezuela.
37. PROECHIMYS CAYENNENSIS TRINITATIS, Allen & Chapman
1893. Bull, Amer. Mus. Nat. Hist. V, p. 223.
Princestown, Trinidad.
38. PROECHIMYS CAYENNENSIS GOELDII, Thomas
1905. Ann. Mag. Nat. Hist. 7, XV, p. 587.
Santarem, Rio Tapajoz, Brazil.
39. PROECHIMYS CAYENNENSIS ORIS, Thomas
1904. Ann. Mag. Nat. Hist. 7, XIV. p. 195.
Igarape-Assu, near Para, Brazil.
40. PROECHIMYS CAYENNENSIS ROBERTI, Thomas
igoi. Ann. Mag. Nat. Hist. 7, VIII, p. 531.
Rio Jordao, Araguar\' district, Minas Geraes, Brazil.
41. PROECHIMYS CAYENNENSIS LONGICAUDATUS, Rengger
1830. Naturg. Saug. Paraguay, p. 236.
North of Paraguay, Matto Grosso, Brazil.
42. PROECHIMYS VACILL.ATOR, Thomas
1903. Ann. Mag. Nat. Hist. 7, XI, p. 490.
Kanuku Mountains, British Guiana.
43. PROECHIMYS HENDEEI, Thomas
1926. Ann. Mag. Nat. Hist, 9, XVIII, p. 162.
Puco Tambo, 50 miles cast of Chachapoyas, Peru.
122 PROECHIMYS— HOPLOMYS
44. PROICCHIMYS RATTINUS, Thomas
1926. Ann. Mag. Nat. Hist, q, XVIII, p. 164.
Tushcmo, Masisea, Rio Ucayali, Peru.
45. PROECHIMYS CANICOLLIS, Allen
iSgg. Bull. Amer. Mus. Nat. Hist. XII, p. 200.
Ronda, Santa Marta district, Colombia.
46. I'ROECHIMY.S DIMIDIATUS, Gunther
1S76. Proc. Zool. Soc. London, p. 747.
South Brazil.
iheringi Group
47. PROECHIMYS IHERINGI, Thomas
191 1. Ann. Mag. Nat. Hist. 8, VIII, p. 252.
Island of Sao Sebastiao, Sao Paulo, Brazil.
Subgenus 'J'rinomvs, Thomas
48. PROECHIMYS ALBISPINUS ALBISPINUS, Geoffrey
1838. Ann. Sci. Nat. X, p. 125.
Ilha de Deos, near Bahia, Brazil.
49. PROECHIMYS ALBISPINUS SERTONIUS, Thomas
1921. Ann. Mag. Nat. Hist. 9, VIII, p. 142.
Lamarao, Bahia, Brazil.
50. PROECHIMYS SETOSUS, Geoffrey
1817. Desmarest, Nouv. Diet. d'Hist. Nat. X, p. 59.
Brazil.
The anoinalm of Kuhl, 1820, Beitr. Zool., p. 17, is, according to Tate, based
on Heteromys anomahis, Thompson.
Genus 5. HOPLOMYS, Allen
1908. HoPLOMYS, Allen, Bull. Amer. Mus. Nat. Hist. XXIV, p. 649.
Type Species. — Hoplomys tniei, Allen.
Range. — Known from Nicaragua, Panama and Ecuador.
Number of Forms. — Three.
Characters. — Skull much like Proechimvs; bullae appear a little smaller.
Cheekteeth with four (at least) outer folds in the upper
series, longer than Pioec/iimvs, placed more obliquely, but isolating in the same
way, the folds stretching further across the tooth, and sometimes tending to
divide up more when isolated. Lower molars reversing the pattern of the upper
series. Frontals and parietals strongly ridged.
Feet as Proechimvs; spines much more developed, at maximum for the
family, more or less concealing the fur, coarse and strong. Tail shorter than
head and body, scaly, naked. Size about largest of subfamily (triiei: head and
body 380 mm.).
Forms seen: gvmnurus.
HOPLOMYS— CERCOMYS 123
According to Goldman (Smiths. Misc. Coll., LXIX, no. 5, p. 124, 1920)
all forms may be regarded as races of the oldest name, gynmurus.
List of N.ivmed Forms
1. HOPLOMYS GVMNURUS GYMNURUS, Thomas
1897. Ann. Man. Nat. Hist. 6, XX, p. 550.
Cachavi, North Ecuador.
2. HOPLOMYS GYMNURUS GOETHALSI, Goldman
191 2. Smiths. Misc. Coll. LVI, no. 36, p. 10.
Rio Indio, near Gatun, Canal Zone, Panama.
3. HOPLOMYS GYMNURUS TRUEI, Allen
1908. Bull. Amer. Mus. Nat. Hist. XXIV, p. 650.
Lavala, Matagalpa, Nicaragua.
Genus 6. CERCOMYS, Cuvier
1829. Cercomys, Cuvier, Hist. Nat. Mammalia, iii, pi. 60.
1881. Thrichomys, Trouessart, Cat. Mamm. Bull. Soc. litudes Sci. Angers, p. 179.
(Nelomys antricola, Lund).
Type Species. — Cercomys cunicularius, Cuvier.
Range. — East Brazil (Pernambuco) southwards (?); known also from Bahia,
Lagoa Santa and Paraguay.
Number of Forms. — Four.
Characters. — Skull broad, rather less heavily ridged than Proechimys and
Hoplomys, braincase appearing rather broader, and parietals
not or scarcely ridged. Jugal not thickened anteriorly, zygoma narrow, and
usually with weak process on lower posterior border. Infraorbital foramen with
separate canal for nerve transmission. Bullae large. Upper cheekteeth with two
outer folds and one inner one, the folds usually clear and straight, not tending
to isolate so completely as in Hoplomys and Proeclmnys. Lower cheekteeth
reversing the pattern of the upper series; P. 4 often with a small extra inner
fold.
Palatal foramina usually abnormally broadened.
E.xternally with soft fur, showing no signs of developing bristles; tail
slightly shorter than head and body, thickly haired. Feet essentially as
Proechimys.
Remarks. — This genus has in the past been compared with Dactylomys and
Myocasior; but from the dental characters and the characters of
the feet I am convinced that it has nothing to do with these genera, but seems
to bear very nearly the same relationship to Proechimys that Isothrix does to
Echimys. namely a hairy-tailed soft-furred representative. A paper has been
published (Boker, 1929, Vcrh. Anat. Ges. Jena, XXXVIII, p. 19) on the bipedal
leaping adaptations of a captivity specimen of this genus.
Forms seen : hiurentius, fostcri, apereoides.
124 CERCOMYS— EURYZYGOMATOMYS
I am convinced that all the three forms seen are not more than racially
distinct from each other, though the first two were described as species.
Thomas states (Proc. Biol. Soc. Washington, 1912, XX\', p. 115) that
apcreuides is synonymous with the earlier described ctiiiicularius.
According to Thomas there are four mammae {Iciurcntius).
List of N.-\med Forms
1. CERCOMYS CUNICULARIUS CUNICULARIUS, Cuvier
1829. Hist. Nat. Mamm. ill, fig. 276.
"Capitanerie des Mines," Brazil.
Synonym; afiercoides, Lund, 1841, Afh. K. Danskc Vidensk Selsk 4,
VIII, p. 98. Lagoa Santa, Minas Geraes, Brazil.
aniricola, Lund, 1841, Afh. K. Danske Vidensk Selsk 4,
VIII, p. 242. Brazil.
2. CERCOMYS CUNICULARIUS LAURENTIUS, Thomas
1904. Ann. Mag. Nat. Hist. 7, XII, p. 254.
Sao Louren90, near Pernambuco, Brazil.
3. CERCOMYS CUNICULARIUS FOSTER!, Thomas
1903. Ann. Mag. Nat. Hist. 7, XI, p. 227.
Sapucay, Paraguay.
4. CERCOMYS INERMIS, Pictct. (Not seen)
1841. Notice An. Nouv. Mus. Gene%'e, ii, p. 33.
Bahia, Brazil.
Genus 7. EURYZYGOMATOMYS, Goeldi
igoi. EuRYZYGOMATo.MYs, CJneldi, Bol. Mus. Paraense, III, p. 179.
Type Species. — Echirnvs spinosus, Rengger. (See Tate, 1935, Taxonomy of
Neotropical Hvstricoid Rodents).
Range. — "Probably throughout the pampas country of Paraguay,
northern Corrientes, Parana, Santa Catharina and Rio Grande
do Sol" (Tate).
Number of Forms. — Three are named.
Ch.xracters. — Skull broad, with poorly marked supraorbital ridges, rela-
tively broad rostrum, prominent bullae (these not excessively
inflated). Palate narrow and rather short; palatal foramina short and broad.
Infraorbital foramen with a separate canal for nerve transmission. Jugal long,
greatly thickened anteriorly, but with posterior projecting process not well
marked; the zygoma more robust than in the genera dealt with above. Mandible
heavily ridged and twisted; coronoid higher than in Proechhnys. Llpper cheek-
teeth with two outer, one inner folds, becoming isolated as islands with wear;
general effect nearer Cercomvs than Proecliinivs. Lower teeth with this pattern
reversed.
Fur bristly, spines about as well developed as in the less spiny members of
Proechimxs, or perhaps less so. Feet narrow, essentially of Proechimys type;
EURYZYGOMATOMYS— CLYOMYS 125
claws of forefoot slightly elongated. Tail strongly reduced, not much longer
than hindfoot, well haired.
I'orins seen : s{>inosiis, cutelliis.
1 do nut think that the above two forms are more than racially distinct from
each other.
List ov N.\med Forms
1. I;LRYZYGOM.'>iTOMYS SPINOSUS SPINOSUS, Desmarest
1817. Nouv. Diet. D'Hist. Nat. 2d Ed. X, p. 57.
Atira, 8 leagues east of Asuncion, Paraguay.
Synonym: brachyurus, Wagner, 1843, Schreber Saug. Suppl. iii, p. 346.
Brazil.
rufa, Lichtenstein, 1818, Abh. Akad. Berlin, p. iy2. Brazil.
2. KURYZYGOMATOMYS SPINOSUS CATELLUS, Thomas
1916. Ann. Mag. Nat. Hist. 8, XVIII, p. 301.
Joinville, Santa Catharina, Brazil.
3. KURYZYGOMATOMYS GUIARA, Brandt (Not seen)
1S35. Mem. Acad. St. Petersb. 6, III, p. 432.
Ypaneme, Sao Paulo, Brazil.
The status of this form appears doubtful (Tate, Bull. Amer. Mus. Nat.
Hist. LXVIII.p. 405).
Genus 8. CLYOMYS, Thomas
1916. Clyomys, Thomas, Ann. Mag. Nat. Hist. 8, XVIII, p. 300.
Type Species. — Echimys laticeps, Thomas.
Range. — Described from Joinville, Santa Catharina, Brazil.
Number of Forms. — One.
Characters. — Essentiallv like Euryzvgomatomys in cranial characters except
that the bullae are abnormally inflated, very much more so
than in other members of this subfamily, a great part visible external to the
paroccipitals when viewed from behind. Dental characters of the one skull seen
too worn for notes.
Externally like Euryzygomatomys, but foreclaws noticeably more developed.
Forms seen : laticeps.
List of N.\med Forms
I. CLYOMYS LATICEPS, Lund, 1S41, nom. nud., Thomas
1909. Thomas, Ann. Mag. Nat. Hist. 8, IV, p. 240.
Joinville, Santa Catharina, Brazil.
(1841, Lund, nom. nud.. Afh. K. Danske Vid. Selsk. 4, VIII, p. 99)
Genus 9. CARTERODON, Waterhouse
1848. Carterouon, Waterhouse, Nat. Hist. Mammalia, ii, p. 351.
Type Species. — Echimys sulcidcns, Lund.
126 CARTERODON— MESOMYS
Range. — Brazil (? Lagoa Santa).
Number of Forms. — One.
Ch.\racters. — Jugal thickened anteriorly, as in Eiiryzvgomatomys and
C/voinys; supraorbital ridges developed, and slight inter-
orbital constriction present in the one skull examined. Nasals broad. Bullae
prominent, hut not extreme. No canal tor nerve transmission in the infraorbital
foramen. Mandible heavily twisted. Zygoma with moderate process on
posterior lower border. Upper incisors one-grooved. l"hc outer side of these
teeth yellow, the inner side white, as remarked bv Waterhouse. Lower incisors
plain. Upper cheekteeth with two outer, one inner folds, the enamel sur-
rounding them thick; lower teeth reversing the pattern.
Size relatively small; fur soft, at any rate compared with most of the genera
of this group; tail shortened, well haired, evidently not so reduced as in Eury-
zygomatomys; feet narrow and long, as in Proechimys; claws moderate.
Forms seen : sukidens.
List of N.^med Forms
I. CARTERODON SULCIDENS, Lund
1841. Mh. K. Danske Vid. Selsk, 4, VIII, p. 99.
(Originally described fossil from Lagoa .Santa, Brazil.)
This genus was originally described from fossil remains, but subsequently
found living.
Genus ID. MESOMYS, Wagner
1845. Mesomys, Wagner, .-^rch. fur Naturg. i, p. 145.
Type Species. — Mesomys ecaiidatus, Wagner.
R.\NGE. — Amazonia; "From the Tocantins River to eastern Peru and
Ecuador" (Tate).
Number of Forms. — Approximately seven.
Ch,\racters. — This genus was described by Thomas as having the skull,
ears and feet of Echiwvs, but the teeth of Proechimys. Skull
with short and narrow rostrum, and well marked supraorbital ridges. Frontals
tending to be very broad, parietals not or scarcely ridged. No canal for trans-
mission of nerve in infraorbital foramen. Bullae relatively large; jugal not
specially broadened, with weak process on posterior border both below and
sometimes above. Palatal foramina short; toothrow far forward in skull.
L'pper cheekteeth of Proechimys tvpe, with narrow folds, usually traces of
four external in the upper series, the lower teeth reversing the pattern.
.Size small, usually or always under 200 mm. head and body, fur heavily
spiny, comparable to that of llophjmys; hindfeet very broad, of arboreal type,
D.5 long; claws prominent. Tail usually slightly longer than head and body,
scaly, poorly haired except terminally.
Forms seen : ferrugiiieus, liispidus. leiiiceps, spicatus, stimiihix.
The few species admitted are all very closely allied to each other.
MESOMYS— LONCHOTHRIX 127
List of Named Forms
1. MESOMYS HISPIDUS, Desmarest
1817. Nouv. Diet. D'Hist. Nat. 2d. Ed. X, p. 58.
"South America."
Synon>Tn: ecaudatiis, Wagner, 1845, Arch. fiir. Naturg. i, p. 145.
Borba, Rio Madeira, Brazil. (For status see Thomas,
Ann. Mag. Nat. Hist. 8, XVI 11, p. 298, 1916.)
2. MESOMYS FERRUGINEUS FERRCGINEUS, Giinther
1876. Proc. Zool. Soc. London, p. 750.
Chamicuros, Rio Huallaga, Peru.
3. MESOMYS FERRUGINEUS SPICATUS, Thomas
1924. Ann. Mag. Nat. Hist. 9, XIII, p. 535.
Tushemo, near Masisea, Rio Ucayali, Peru.
4. MESOMYS LENICEPS, Thomas
1926. Ann. Mag. Nat. Hist. 9, XVIII, p. 348.
Yambrasbamba, Amazonas, Peru.
5. MESOMYS STIMULAX, Thomas
191 1. Ann. Mag. Nat. Hist. 8, VII, p. 607.
Cameta, Lower Tocantins, Brazil.
6. MESOMYS DIDELPHOIDES, Desmarest. (Not seen)
1817. Nouv. Diet. d'Hist, Nat. 2nd Ed. X, p. 58.
Probably from Brazil.
7. MESOMYS OBSCURUS, Wagner. (Not seen)
1840. Abh. Akad. Wiss. Munch, iii, p. 196.
Brazil.
Genus 11. LONCHOTHRIX, Thomas
1920. LoNCHOTimix, Thomas, Ann. Mag. Nat. Hist. 9, VI, p. 113.
Type Species. — Lonchothrix emiliae, Thomas.
Range. — Described from Villa Braga, Rio Tapajoz, Brazil.
Number of Forms. — One.
Characters. — Skull closely similar to that of Mesomys. Cheekteeth com-
pared by Thomas to those of a small Erethizon ; upper molars
with three outer, one inner folds, these noticeably wide and deep; lower cheek-
teeth evidently with only two inner folds, one outer in all molars; P. 4 with
traces of four folds (three main, one vestigial); the folds wide in the lower
series.
Externally the form is striking owing to the heavily tufted tail, which is
considerably longer than the head and body, but hairy at the end only, the
upper part scaly and covered with short spines. Fore and hindfeet broad, of
arboreal type, as in Mesomys. Spines of body veni' highly developed com-
paratively, even the belly being semi-spinous.
Little appears to be known of this genus; the teeth do not appear very
128 CAPROMYINAE: CAPROMYS
typical of this section; I include it here on account of the resemblance to
Mesomys in cranial and external characters.
Forms seen : emiliae.
List of Named Forms
1. LONCHOTHRIX EMILIAE, Thomas
1920. Ann. Mag. Nat. Hist, 9, VI, p. 114.
Villa liraga, Rio Tapajoz. Brazil.
Subfamily CAPROMYINAE
Geogr.aphic.^i. Distribution. — Cuba, Jamaica, Bahama Islands, Swan
Island (Gulf of Honduras), and one
form named from Venezuela.
Number of Genera. — As here understood the group contains three genera,
of which one is not represented in the British Museum.
Characters. — Not essentially different from the Echimyinae, but cheek-
teeth evergrowing, characterized by two outer, one inner
folds in the upper series, the folds in adult completely filled with cement, the
teeth flatcrowned and changing little or not at all during the animal's life.
Paroccipital processes usually, not always, tending to stand apart from the
bullae. Form usually robust, not Rat-like; tail haired, long and prehensile or
strongly reduced; habits terrestrial or arboreal.
Key to the Genera of Capromyinae, not including the
genus Pruccipromvs (not seen)
Tail considerably longer than hindfoot; claws more prominent; (habits
arboreal). (Tail prehensile, constant ?). Capro.my'S
Tail scarcely longer than hindfoot; claws less prominent; (habits terres-
trial). Geqcafromys
Genus I. CAPROMYS, Desmarest
1822. Capbomys, Desmarest, Bull. Soc. Philom. Paris, p. 185.
Type Species.— Cupromvs foiiniieri, Desmarest=Isudoii pilorides, Say.
Range. — Cuba, including the Isle of Pines.
Number of Forms. — Six.
Ch.\racters. — Skull long and rather flat, a postorbital-like ridge can be
present; parietals may be well ridged; jugal with well marked
and strong backwardly directed process. Bullae prominent. Paroccipital pro-
cesses usually slightly lengthened, and standing apart; in the one skull seen of
C. nana (adult female), the paroccipital processes join the bullae, about as in
Echimyinae.
CAPROMYS 129
Infraorbital foramen with no canal for nerve transmission. Palate slightly
constricted anteriorly, but less so than in Myocastor or Dactylomys; palatal
foramina medium. Mandible with angular process drawn backwards, and
stronglv lifted outwards; condyle high; coronoid process low. Incisors narrow.
Upper cheekteeth as already described; the lower series reverse the pattern
of the upper series, the premolar has also a vestigial e.xtra inner fold.
Externally rather large as a rule; fur harsh; feet broad, of arboreal type, or
more or less; claws prominent, D.5 relatively long, hallux medium. A tendency
in the few skins examined for D.4 to be a little longer than D.3. Forefoot with
four digits well developed, pollex small. Tail long, haired, said to be prehensile
in at least one species, and may be so throughout the genus.
The species of Capromys were revised bv Chapman, 1901, Bull. Amer. Mus.
N'at. Hist., vol. XI\', p. 313.
Forms seen : prehensilis, pilorides, melamirm, nana.
C. Jiana, of which one skull alone is available, seems considerably smaller
than the remainder; it was originally described as fossil, but subsequently found
living. C. melanurus, of which one specimen only has been seen, has a much
more heavily haired tail than the remainder. C. prehensilis and C. pilorides are
distinguishable from each other on length of tail, and the latter is stated to have
a much heavier skull. The genus is not very well represented in London.
List of Named Forms
(The references and type localities of all Capromyinae are the work of
Mr. G. W. C. Holt.)
1. C.-\PROMY.S PILORIDKS PILORIDES, Say
1822. Joum. Acad. Philadelphia, ii, p. 333.
Cuba.
Synonym; Joitrnieri, Desmarest, 1822, Mem. Soc. Hist. Nat. i, p. 43.
Cuba.
? qiiemi, Fischer, Add. ad Synops. Mamm. 1830, p. 389.
2. CAPRO.MYS PILORIDKS RKLICTLS, .•\llen
191 1. Bull. Mus. Comp. Zool. Harvard Univ. 54, p. 207.
Casas Mountains, Nueva Gerona, Isle of Pines, Cuba.
3. CWPRO.MYS PREHENSILIS PREHENSILIS, Poeppig
1824. Joum. Acad. Philadelphia, 4, p. 11.
Wooded parts of Southern Cuba.
Synonym: poeyi, Guerin, 1834, Mag. Zool. IV, PI. XV, 5 pp., and
poeppingi, Lesson, 1842, Nouv. Tabl. Regn. .-Xnim. p. 124.
4. CAPROMYS PREHENSILIS GUNDL.\CHI, Chapman
igoi. Bull. .Amer. Mus. Nat. Hist. XIV, p. 317.
Nueva Gerona, Isle of Pines.
5. CAPROMYS MEI.ANL RLS, Peters
1864. Mon. Ber. .■ikad. Wiss. Berlin, p. 384.
Manzanillo, Cuba.
Synonym: pallidus. Peters, 1864, Mon. Ber. .Akad. Wiss. Berlin, p. 384.
Cuba.
9 — Living Rodents — I
p-IG. 14. GeOCAPROMYS BROWNII, Fischcr.
B.M. No. 1334 C.; I.
Fig. 15. GEOCiPROMYS brownii, Fischer.
B.M. No. 1334 t.; / I.
CAPROMYS— GEOCAPROMYS 131
6. CAPROMYS NANA, G. M. Allen
1917. Proc. New England Zool. Club, VI, p. 54.
Sierra de Hato Xucvo, Province of Matanzas, Cuba.
The " Capromys" elegans of Cabrera, 1901, is a member of the Murine
genus Phloeomys.
„ b
Fig. 16. Geocapromys brownii, Fischer.
Cheekteeth: B.M. No. 1334C.; X7.
Genus 2. GEOCAPROMYS, Chapman
1901. Geocapromys, Chapman, Bull. Amer. Mus. Nat. Hist. XIV, p. 313.
Type Species. — Capromys brozcni, Fischer.
Range. — Jamaica, Swan Island, and the Bahamas.
Number of Forms. — Three.
Ch.'VR.'^cters. — Like Capromys, but tail strongly shortened, scarcely longer
than hindfoot; feet with less prominent claws, and pollex
more reduced (the feet, however, do not seem very different in formation from
those of Capromys); habits terrestrial. "Dentition and cranium as Capromys,
but ascending portion of maxillary arch of zygoma wider, superior margin of
squamosal narrower, and without process, and occipital region lower" (Chap-
man). 1 am inclined to doubt the constancy of these cranial characters between
the two groups, particularly when C. nana, not known in Chapman's dav, is
compared. The skull may have a sagittal ridge in adult; it may be that this is
132 GEOCAPROMYS— PROCAPROMYS
present also in Caproinvs, but not in our small series; the paroccipitals in Geo-
capromys appear relatively short. The dentition is as in Capromvs; in the
cutting teeth of a newly born animal, the folds are quite open and well marked,
hut even as early in life as that the dentition is relatively simple. In M.i lower,
the vestigial inner extra front fold of Capromxs is more clearly marked.
The group was proposed as a subgenus, but has since been given generic
rank; the differences in habit and tail characters between the two groups seem,
I think, to warrant their separation. The broader ascending portion of the
maxillary seems very well marked in all our series of Gcocaproiiivs with one
exception, which might be wrongly identified; other than this it seems a clear
distinction between Geocaproinxs and Caproiins.
The species were revised by Chapman, 1901, Bull. Amer. AIus. Nat. Hist.,
vol. XI\', pp. 313-323. hrowni seems rather larger, and with a rather heavier skull
than thoracatus and ingrahami, which appear very doubtfully distinct from each
other. Certain cranial characters are said to distinguish hrozcnii from thoracatus,
and the form of the ear.
Forms seen : hrou-nii, thoracatus, ingraliaiiii.
List of Named Forms
1. GEOC-^PROMYS BROWXII, Fischer
1829. Syn. Mamm. Addenda, page 3S9 ( -= p. 589).
Jamaica.
Synonym: hrachyurus, Plill, 1S51, in Gosse, Nat. Sojourn in Jamaica,
p. 471. Jamaica.
2. GEOCAPROMYS THORACYTU.S, True
1SS8. Proc. U.S. Nat. Mus. XI, p. 469.
Little Swan Island, Gulf of Honduras.
3. GEOCAPROMYS INGRAHAMI, Allen
1S91. Bull. lAmer. Mus. Nat. Hist. Ill, p. 329.
Plana Keys, between Acklin Island and Mariguana, Bahama Islands.
Genus 3. PROCAPROMYS, Chapman
lyoi. Proc.\pro.mvs. Chapman, Bull. .Amer. Mus. Nat. Hist. XIV, p. 322.
Type Species. — Capromxs geaxi, Pousargues.
Range. — Described from \'enezuela, central coastal region.
Nl-.mber oe F0RM.S. — One.
Rem.\rks. — Not represented in the British Museum; differing m dental
details from Capromxs and Geocapromxs.
" Size smaller than the smallest known species of Capromxs, tail half as long
as head and body — enamel outline in the first three upper molars continuous,
with two external and one internal folds; the fourth, last molar with three dis-
tinct and disconnected transverse enamel ellipses, the posterior one about half
the size of either of the anterior two; enamel outline in the four lower molars
continuous, the first molar with three internal and one external folds, the first
PLAGIODONTINAE: PLAGIODONTIA 133
and second interior folds being more extended than in the corresponding tooth
of Capramys; the remainini; three lower molars each with two internal and one
external folds, the enamel enclosed space on the posterior margin of the last
molar being scarcely wider than the enamel itself" (Chapman).
Chapman suggests that this represents the ancestral mainland type from
which Capromys and Geocapromys descended.
List of Named Forms
I. PROCAPROMYS GEAYI, Pousargues
1899. Bull. Mus. Paris, p. 150.
Mountainous coastal region on the slopes of the range which separates
the town of Caracas from the port of La Guaira, Venezuela.
Subfamily PLAGIODONTINAE
Geographical Distribution. — Dominican Republic.
Number of Genera. — One.
Characters. — Not unlike the Capromyinae, but cheekteeth differing
markedly from any Hystricoid Rodent examined; the upper
molars with only one fold each side, these folds yery long and deep, penetrating
far into tooth, running parallel to each other and set obliquely; folds well filled
with cement, as in Capromyinae; each upper tooth with (in the one examined)
a strong outwardly-pointing external projection on the outer side, adjacent to
the external border of the outer fold. Cheekteeth eyergrowing. Lower cheek-
teeth with two long, deep inner and one shallow outer folds. Paroccipital pro-
cesses much lengthened. Jugal simpler than in Capromyinae, without processes
on upper or lower border. Tail naked.
Rem.\rks. — The status of this genus must remain provisional ; it does not
appear to agree with Capromyinae sufficiently to be included in
the same subfamily, in dental characters; but only one skull is available for
examination.
Genus I. PLAGIODONTL-\, Cuvier
1836. Plagiodontia, Cuvier, Ann. Sci. Nat. Paris, 2, VI, p. 347.
Type Species. — Plagiodontia aedium, Cu\ier.
Range. — As in the subfamily.
Number of Forms. — Two.
Characters. — Miller compared his P. hylueum with Geocapromys broumii;
the most noteworthy differences quoted were (in Plagio-
dontia) the less breadth between the lachrymals, the more anterior positions
of the swellings caused by the frontal sinuses, the zygoma much more slender,
the upper part not bearing an orbital process, the jugal slender, without posterior
concavity and posteroinferior process, the excessively long paroccipital processes,
the smaller incisive foramina, the greater width of the mandibular masseteric
134 PLAGIODONTIA— DACTYLOMYINAE
ridge. Some interorbital constriction is apparent. There is no canal in the
infraorbital foramen for nerve transmission. The skull appears depressed, or
slanting downwards, posteriorly; the palate is slightly constricted anteriorly.
The cheekteeth are as described above.
The feet are heavy, the tail naked, of moderate length, the ears small.
Claws well developed; D.5 hindfoot relatively long.
Miller suggested that the animal is more nearly related to Ade/pliuinva, a
Patagonian fossil, than to living Hutias, with which it is currently associated.
Forms seen: liyhuuin.
List of Named Forms
1. PLAGIODONTIA AKDIUM, Cuvicr
1S36. Ann. Sci. Nat. Paris. 2, VI, p. 347.
Dominican Republic.
2. PLAGIODONTIA HYLAKL'iM. Miller
1927. Proc. U.S. Nat. Mus. LXXVII, no. 16, p. 4.
Guarabo, lo miles east of Jovero, Samana Province. Dominican
Republic.
The type species does not appear to be known at present as a living animal.
Originally described in 1836, little more was heard of the genus until Miller
described livlcieum ninety years later. In 1916 Miller described some bones
taken in the Dominican Republic; the impression at that time was that the
animal was extinct.
Subfamily DACTYLOMYINAE
Gf.ographical Distribution. — South America: \'enezuela, Colombia,
Amazonia, Ecuador, Peru, Bolivia, Para-
guay, S.FL Brazil, etc.
Number of Genera. — Three.
Characters. — Cheekteeth brachyodont, excessively broad and heavy, the
pattern essentially consisting of a deep re-entrant fold in the
middle of each upper molar more or less completely dividing each tooth into
two lobes, each of which is subdivided by a broad external fold. The pattern
varies slightly within the genera, but the general somewhat prismatic effect is
unmistakable. There is a strong tendency towards anterior constriction ot the
palate, as in M\ociistur\ the paroccipital processes are usually as in Ec/iimys,
i.e. curved forward under the bullae, but in some specimens of Dactylomys
dactylinus, they stand apart from the bullae and cannot be distinguished from
those of Geocapromys, which makes the former separation of this section of
Rodents into forms with large paroccipitals (family "Capromyidae") and forms
with paroccipitals curved under the bullae (family Echimyidae, hitherto
including Dactylomys) unretainable. Skull number 22.5.4.4. in the British
Museum appears just as Geocaproinvs, in paroccipital structure. The palatal
foramina are small or nearly obsolete; the palate is very narrow and extends to
a level with hinder part ot .M.3 or slightly behind it.
DACTYLOMYINAE: THRINACODUS 135
The fur is soft, not developing spines. A feature of the group is the extreme
elongation of certain digits in the manus and pes of all except Thrinacodus, a
character very unusual or unique within the Order. I am told that these are
climbing animals, and that they grasp the branches between their third and
fourth digits. The tail is usually much longer than the head and body, and
may be heavily haired, or naked and reptilian in appearance.
If the families Capromyidae, Myocastoridae and Thryonomyidae are to be
retained as distinct from the Echimyidae, I suggest the present group should
also form a special family, Dactylomyidae. For the purposes of the present
work, however, all these groups are kept within one family, as already noted.
Key to the Gener.a of D.actylomyinae
Digits three and four of both fore and hindfeet not specially elongated,
and not broadened. THRiN.'iCODUS
Digits three and four of both fore and hindfeet much elongated, and
considerably broadened.
Palate much constricted anteriorlv; main lobes of upper cheekteeth
not united by enamel bridges. Dactylomys
Palate scarcely constricted anteriorly; main lobes of upper cheekteeth
united by narrow enamel bridges. Kannabateomys
Genus i. THRINACODUS, Gunther
1879. Thrinacodus, Gunther, Proc. Zool. Soc. London, p. 144.
Type Species. — Thrinacodus albicauda, Gunther.
R.\N"GE. — Colombia and Venezuela.
Number of Forms. — Three.
Characters. — Essential cranial and dental characters as Dactylomys, ne.xt
to be described. Palate as Dactylomys. Digit elongation at
minimum for the subfamily; the foreclaws sharper than in related genera;
polle.x as usual in the group scarcely traceable. Digits narrow; D.3 and D.4
longer than the outer digits in forefoot; D.2 longer than D.5. Hindfoot with
digits not abnormal, essentially like those of forefoot except that a short hallu.x
is present. Fur thick and soft; tail longer than head and body, moderately to
poorlv haired.
(The paroccipital processes agree with those of the Echimyinae.)
Forms seen : eda.x, albicauda.
The three forms are at present regarded as species ; I am not very- convinced
as to their distinctness from each other, and they should perhaps be regarded
as races.
List of Named Forms
I. THRIN.'VCODLS .\LBlCAUD.-\, Gunther
1879. Proc. Zool. Soc. London, p. 144.
Near Medellin, Colombia.
136 THRINACODUS— DACTYLOMYS
2. THRINACODUS APOLINARI, Allen
1914. Bull. Amer. Mus. Nat. Hist., XXXIII, p. 387.
Tomeque, Bogota district, Colombia.
3. THRINACODUS ED.A.X, Thom.ns
1916. Ann. Mag. Nat. Hist. 8, XVIII, p. 299.
Sierra de Merida, Venezuela.
Genus 2. DACTYLOMYS, Geoffroy
1S38. D.\CTYLOMYS, CJeotfroy, .^nn. Sci. Nat. Paris, 2, X, p. 126.
1916. L.\CHNOMYS, Thomas, Ann. Mag. Nat. Hist. 8, XVIII, p. 298. (Dnclylomys
peruanus, Allen). Valid as a subgenus.
Typi; Sphcies. — Dactvlomvs tvpiis, Geotfroy — iic/z/wivv dtictvliinis, I)es-
marest.
R.'iNGE. — Known from Ecuador, Peru, .Amazonia, and Bolivia.
Number of Forms. — Five.
Characters. — Skull typically with weak postorhital-like ridges, and a
sagittal crest developed in adult; paroccipital processes, as
noted above, in the type species tending to stand apart from the bullae (age
character?), or in smaller forms about as in Ecliimys. Jugal not thickened
anteriorly, but relatively broad, with small process on upper and lower border
posteriorly. Bullae moderately large. Infraorbital foramen with no canal for
nerve transmission. Palate constricted anteriorly so that the premolars almost
touch each other. Upper cheekteeth as described above, the inner side of the
lobes narrow and contracted ; lower molars with two inner folds, the hinder one
completely divitling the tooth; lower premolar with a small extra lobe anteriorly,
behind which it is not so completely divided into lobes as are the molars.
Size rather large; fur soft; tail longer than head and body, tvpically almost
completely naked e.xcept the portion joining the bodv, which is well haired.
Forefoot with the two central digits greatly lengthened, broadened to a degree;
D.2 also considerablv lengthened; D.5 short; poUex untraceable normally.
Claws weak, nail-like. Ilindfoot not very different from forefoot except that
the hallux is moderately developed, and the claws are more prominent.
Lachnomys, proposed as a subgenus bv Thomas for D. peiiiaiiiis, is given
generic rank by Tate, though it scarcely seems even a valid subgenus. The fur
is much thicker, and the tail is fully haired throughout. The dental details
given by Thomas to divide the two subgenera are not clear to me in our series.
Forms seen : dactvliiius, caiiesceiii, peiiianus.
The species bolivicnsis appears from description to be very closely allied to
the type species.
List of Named Forms
Subgenus Dactvlomvs, Geoffroy
1. DACTYLOMYS D.\CTYIJNUS DACTYLINUS, Desmarest
1S17. Nouv. Diet. d'Hist. Nat. 2nd Ed. X, p. 57.
No locality in original description.
Synonym: typtis, CJeoffrny, 1S38, Ann. Sci. Nat. Paris, 2, X, p. 127.
Brazil (?)
DACTYLOMYS— KANNABATEOMYS 137
2. DACTYLOMYS DACTYI.INL'S CANESCENS, Thomas
1912. Ann. Mag. Nat. Hist. 8, XI, p. 87.
Itacoatiara, Middle .Amazons, Brazil (below Manaos).
3. DACTVLO.MYS D.^CTYLINU-S MODESTUS, Lonnbcrg
1921. Archiv. fiir Zool. XIV, no. 4, p. 38.
Banks of Rio Curaray, Ecuador (Prov. del Oriente).
4. DACTYLO.MYS B(:)LIVIEN.SI.S, Anthony
1920. Journ. Mamm. Baltimore, I, p. 82.
Mission San Antonio, Cochabamba, Bolivia.
Subgenus Lachnomys, Thomas
5. DACTYLOMYS PERUANUS, Allen
1900. Bull. Amer. Mus. Nat. Hist. XIII, p. 220.
Juliaca, Peru.
Genus 3. KANNABATEOMYS, Jentink
i8gi. Kannabateomys, Jentink, Notes Leyden Mus. XIII, p. 109.
Type Specie.?. — Dactylomys amblyonyx, Wagner.
Range. — Paraguay and South-eastern Brazil.
Number of Forms. — ^Tvvo.
Char.^cters. — The palate very slightly constricted anteriorly; the check-
teeth not completely divided into lobes, the lobes being
connected bv a small bridge; the enamel folds more nearly perpendicular to the
molar series than in Dactylomxs. The lower cheekteeth with an anterior
V-shaped fold, and a posterior elongated one, as in Dactylomxs, but the lobes
thus formed united by a small bridge. Lower premolar like that of Dactvlomys,
but anterior lobe larger.
Skull much like that of Dactvlomys; apparently a sagittal ridge is not formed ;
the paroccipital processes curve under the bullae.
Externally rather smaller than typical Dactylomys; the fur thick, soft.
Forefeet much as in Dactylomys; hindfoot relatively broad, essential digit
arrangement as in Dactylomxs. Tail very long, relatively well haired.
Remarks. — Very closely allied to Dactylomys. The character of the palate is
perhaps the most important in keeping the two genera separate.
Forms seen : amblyonyx, pallidior.
List of Named Forms
1. KANNABATEOMYS AMBLYOViTC AMBLYONY'X, Wagner
1845. Archiv. fiir N'aturg. i, p. 146.
Ypanema, Province of Sao Paulo, Brazil.
2. KANNAB.VnCOMYS AMBLYONV.X I'ALI.IDIOR. Thomas
1903. Ann. Mag. Nat. Hist. 7, XI, p. 489.
Sapucay, Paraguay.
Fic. 17. Kannabatf.omys amblyonyx amblyonyx, Wagner.
B.M. No. 1.6.6.67, V; ■ li.
Fig. 18. Kannab.^teomys amblyonyx amblyonyx, Wagner.
li.M. No. 1.6.(1.67, V; X li.
MYOCASTORINAE
139
Fig. 19. Kannabateomys amblyon^-x amblyonyx, Wagner.
Cheekteeth: B.M. No. 1.6.6.67, 9; X5-
Subfamily MYOCASTORINAE
Geographical Distribution.
Number of Genera. — One.
-Southern South America.
Characters. — The external form robust and heavy, the size larger than in
other members of the family; the genus is quite one of the
giants of the Order. The external characters show strong specialization towards
aquatic life; the hindfeet have four of the toes webbed; D.5 is free, and perhaps
used for combing the fur. The hindfeet are much larger than the forefeet,
which bear a rather rudimentary pollex, and four well-developed main digits;
all the digits are armed with sharp large claws.
The skull is more heavily ridged for attachment of muscles than in other
Neotropical Echimyidae, and is the only member of the familv which tends in
this character to approach the .\frican Thryonomys. The paroccipital processes
are greatly elongated; the lateral process of each stands well apart from the
main downwardly pointing bone.
The cheekteeth decrease markedly in size from M.3 forwards; thev are
semi-rooted, and broadened, with strong inner and outer re-entrant folds, which
are long retained; the palate is strongly constricted anteriorly.
I40 MYOCASTOR
(;cnus I. MYOCASTOR, Kerr.
1792. Mvoc.'VSTOU, Kerr, Anini. KiiiRd., p. 225.
1S05. MvopOTAMUs, (ieoffroy, Ann. Mus. d'Hist. Nat. VI, p. S2. (Myopotoiinis
boiiarieiisis, Geoff roy.)
Type Species. — Mas Kjypus, .Molina.
R.'\NGE. — Southern South America; Holiister in a review of races repre-
sented in the American Museum quotes as localities: Chile, the
Straits of Magellan, Buenos Ayres, Santa Fe and Paraguay, Parana River; Rio
Negro and Rio Salados, Patagonia. Whether the genus ranges farther north
than any of these has not been ascertained; quoted by Waterhouse from
Peru.
Number of Forms. — Three.
Cn.\R.\CTERS. — The nasals are somew hat arched, the frontals broad and flat,
the parietals deeplv depressed, and in adults a very strong
sagittal ridge is present. There is a sharply pointed but short squamosal process
and a small postorbital process to the frontals. The anterior zygomatic root is
placed farther back than normal, over the middle of the toothrow. The occipital
region is high and prominent. The bullae tend to spread sidewavs, with the
neck pointing outwards and upwards, approaching the type found in Castor,
though much less developed than in that genus. The hamular processes are
thick, the palate very narrow anteriorly, broad posteriorly. Jugal thick, broader
posteriorly, with an upwardly projecting process on posterior border. There is
no special canal for nerve transmission in the infraorbital foramen.
The mandible is immensely heavily ridged and distorted outwards, the
angular process sharply drawn backwards. The coronoid process is obsolete.
The cheekteeth are extremely hvpsodont ; the fundamental pattern of the
upper series, judging by a young specimen, appears to be two external re-entrant
folds, the front one placed far forwards, the second one about in the middle of
the tooth, and two internal folds, the first almost meeting the second outer one,
the second placed posteriorly, rapidly extending across the tooth and cutting
off the posterior part altogether. The enamel surrounding the folds is wide, the
general effect of the dental pattern rather complex, probably not changing much
tiuring the animal's life. In the lower series, there are three inner and one outer
re-entrant folds; P. 4 has one small extra inner fold. M.3 is in both jaws con-
siderably the largest tooth, in the adult; M.2 is markedly larger than the anterior
two teeth, which tend to wear down in old age. In these front teeth, the folds
tend to isolate, but the effect is considerably different from such types as
Eiirvzvgiiiiiiitdmys in which as the folds isolate the pattern tends to become
simpler.
The general effect of the teeth is reminiscent to a degree of that of
Castor, perhaps owing to the similar life which these two unrelated animals
lead.
The incis(jrs are broad and powertvd.
The essential external characters are described above; the fur is soft and
MYOCASTOR 141
thick, and of some commercial value ("Nutria"). The tail is moderate in
length, scalv and poorly haired.
The largest of a small series of skins at the British Museum is 586 mm. head
and body; whether this would represent about the extreme development for
the genus I do not know.
Forms seen : coy pus.
Fig. 20. Myocastor corpus santaecruz.\e, HoUister.
B.M. No. 16. 10.3. 85; X {.
List of N.mvied Forms
(References and type localities by Mr. G. \V. C. Holt.)
I. MYOCASTOR COVPLS COYPLS, Molina
1782. Sagg. Stor. Natur. Chile, p. 287.
Chile.
Synonym: popelairi, Wesmael, 1841. Hull. .Ac. Roy. Brux. \III, 2
p. 61.
chilensis. Lesson, 1842, Nouv. Tabl. Ri-gn. Anim. p. 126.
Fig. 21. Myocastor coypus s.\ntaecrl'z.\e, Hollister.
B.M. No. 16.10.3.85; X J.
Fio. 22. Myocastor coy'pus santaecruzae, Hollister.
Mandible from below: B.M. No. 16. 10.3. 85 ; • *.
Fio. 23. Myocastor coypus santaecruzae, Hollister
B.M. No. 16.10.3.8s; X i.
Cheekteeth: B.M. No. 16.10.3.85; ,< 2.
144 THRYOXOMYINAE: THRYONOMYS
2. MYOCASTOR COVI'LS BONARIHNSIS, Geoffroy
iSof) (1805). Ann. Mus. d'Hist. Nat. VI, p. 82.
Paraguay.
SjTionym: castorides, Barrow. 1815, Trans. Linn. .Soc. London, XI,
p. 167. Brazil (?)
3. MYOCASTOR COYPUS SANTAECRUZAE, Hullister
IQ14. Proc. Biol. Soc. Washington XXVII, p. 57.
Rio Salado, near Los Palmares, Santa Cniz, Arcentina.
Subfamily THRYONOMYINAE
Gfogr.\phral Distribution. — Africa, widely distributed south of the
Sahara. "Central and East Africa from
Uahr-el-Ghazal and Uganda to Eastern Cape Province" (St. Leger); Nigeria;
Angola; T. swindcritiiuis group; Kenya, Uganda, North Congo and Nyasaland,
r. i^regoriamis group.
Number of Genera. — One.
Characters. — Skull massive, excessively prominently ridged; cheekteeth
rooted, similar in pattern to some of the genera of Echimv-
inae; incisors powerful, the upper ones heavily three-grooved; paroccipital
processes elongated; occipital region of skull extremely powerfully developed.
Arrangement of digits of fore and hindtoot perissodactyle; hallux entirely
suppressed ; D.5 of manus vestigial ; claws thick and heavy, more or less fossorial.
The shoulder-blade as described by Tullberg is apparently peculiar, and not like
that of the other Echimyidae examined by him.
Remarks. — As indicated above, \yithout comparing the shoulder-blade of
this animal with all other genera included here, it is not wise to
base a separate family on this alone. The digit reduction, unique in the present
family, is too uncertain a character in other groups to base family characters on.
Nevertheless I am not sure that this animal is rightly referred to the present
family, or if it is an entirely distinct otfshoot; it seems to stand alone rather in
the Ilvstricoid group, though having no very striking characters to separate it
off from the remainder of the more normal genera.
Genus i. THRYONOMYS, Fitzinger
1827. AuL.'iCODUS, Temminck, Mon. Mam. Tab. ML'th., p. xxvi. (Not of Eschscholtz.)
{AulacodiiS S7iinderiaitus, Temminck.)
1867. Thryonomvs, Fitzmger, Sitz.-B. K. Akad. Wiss.Wien, Math. Nat. CI., 56, p. 141.
1922. Choeromys, Thomas, .Ann. Mag. Nat. Hist. 9, IX, p. 390. {Thryonottiys gre-
goriamts, Thomas.)
Type Species. — Auhicodus scniipdlnuitus, Hcuglin.
Range. — As in the subfamily Thrvonomyinae.
Number of Forms. — Ten.
Characters. — Skull very prominently ridged; rostrum high, broad, rather
reminiscent of Pedetes except that the nasals are less arched
and the zygomatic plate is not specially projected forwards; jugal and zygoma
THRYONOMYS 145
thick, not markedly angular, the jugal nearly in contact with the iachryjnal, the
zygomatic region bearing some resemblance to that of Pedetes. Frontals broad,
with sharp angular depression immediately in front of the suture formed bv the
frontals and parietals each side, in adult. Infraorbital foramen very large, with
well marked canal for nerve transmission. Parietals converging into an exces-
sively high sagittal ridge; occipital region high and prominent; bullae moderate
in size; paroccipital processes considerably lengthened (less so than Myocastor,
probably more so than other Echimyidae). Bony palate e.xtending slightly
behind M.3; the palate straight, broad; palatal foramina verv broad and large.
Mandible with moderate coronoid process, angular portion low, drawn back-
wards to a degree, the mandible very heavily ridged and distorted outwards.
Cheekteeth semi-hypsodont, broad and hea\'y; the enamel surrounding the
folds thick, the folds broad originally, tending to become narrower with wear,
evidently not isolating on crown surface to any degree. Upper cheekteeth with
two outer, one inner folds; lower teeth reversing the pattern, P. 4 with small
extra inner fold. In old age, the pattern wears out.
Incisors very broad and powerful, probably more so than in any other
Rodent, the upper ones three-grooved, the main groove normally placed
centrally, the second and third placed between this and the inner edge of the
teeth.
Externally, size rather large (perhaps approaching 600 mm. head and
body); form heavy; fur harsh and bristly. Tail not long, comparatively well
haired. Forefoot with three main digits, the centre one longest, a minute
pollex, and D.5 so reduced that it must be almost functionless, though the claw
is about as well developed as those of digits 2, 3 and 4. Hindfoot lacking hallux;
the digits otherwise like those of forefoot, but longer; D.5 greatly reduced. I
am told that a specimen kept at the London Zoological Gardens "shed its tail"
when picked up, thus recalling a feature which is common in the Echimvinae.
In the su'inderianus group, the skull is much arched anteriorly; the gregorianiis
group was given the generic name " Choeromys" by Thomas on account of the
"almost complete absence of the large frontal sinuses present in Thrxonomxs,
and so developed as to produce a totally different shape of the opening that
leads from the cerebral to the olfactory fossa of the skull. The opening is
narrow below, broad above in Choeromys, broad below, narrow above in
Thryonomys, where its upper corners have been compressed by the large frontal
sinuses; owing to this absence of sinuses the frontal area is flat instead of
convex."
(The tail was also stated to be more reduced in "Choeromys," but in this
character T. sclateri is intermediate, having the tail nearly as long as in the
typical group. But in any case the tail is strongly reduced comparatively in the
whole genus.)
A cranial character such as this, though clearly marked, does not seem of
generic importance when one takes into account the differences to be found
in the skull of other Hvstricoid genera, for instance, Coendou, in which closely
related forms (as laemitutn and mexuanum) may ha\e the skull, in the one case
arched, in the other flat; or llystrLx, in which the nasals vary extremely, even in
10 — Living Kotlents — I
Fig. 25. iThryonomys gregorianus gregorianus, Thomas.
B.M. No. 34.6.2.68, ?: .-, I.
Fig. 26. Thryonomys gregorianus gregorianus, Thomas.
B.M. No. 34.6.2.68, V; ■ I-
Fig. 27. Thrvonomys cregorianus gregorianus, Thomas.
B.M. No. 34.6.2.68, ?; X i.
Fig. 28. Thrvo.nomys grecori.\nus cregorianus, Thomas.
Cheektteth: B.M. No. 34.6.2.68, 9; x 4.
148 THRYONOMYS
the closely allied African Crested Porcupines {cristatu compared with ajricae-
atistralis); these two groups of Thryonomys are so essentially similar in all other
characters that I do not think Choeromys is worth retaining even as a subgenus.
Forms seen: aiigolac, congicus, grcguiiaiiiis, harrisoiii, raptoriim, sclateri,
su'iihhriauus, TCiriigaiiis.
List ok Named Forms
(References and type localities hy Mr. G. W. C. Holt.)
szcindcriainis ( j roup
I. THRYONOMYS SWINDERL-WUS SWINDKRLANUS. TLiiimmck
1827. Monogr. Mamm. i, p. 248.
Sierra Leone. (For full range of specific group see "Subfamily
Thr\'onomyinae," page 144.)
z. THRYONOMYS SWINDERIANUS VARIKGATUS, Peters
1S52. Reise nach Mozambique, Zool. Saug, p. 13S.
Africa. Was first mentioned in Manuscript, Peters, 1S45, •''nd recorded
from Tette, Macanga, Sena and Boror.
Synonym: calamophagus, de Beerst, iSgy, Pousargues. Bull. Mus.
Paris, p. 160. Nyasa, Central Africa.
seiiiipahiialiis. Heuglin, 1864, Nov. Act. Acad. Leop. Dres-
den, XXXL P- 6- Central Africa.
3. THRYONOMYS SWINDERIANUS RAPTORUM, Thomas
1922. Ann. Mag. Nat. Hist. 9, IX, p. 392.
Nigeria, Lagos.
4. THRYONOMYS SWINDERIANUS ANGOLAE, Thomas
1922. Ann. Mag. Nat. Hist, g, IX, p. 392.
Angola, junction of Luandu and Cuje Rivers.
grcgoriatius Group
5. THRYONOMYS RUTSHURICUS, Lunnbcrg
1918. Stockholm. Vet, Ak. Handl. 58, no. 2, p. 78.
Central Africa, Rutshuru, east of Rutshuru River, half-way between
Lake Albert Edward and Lake Kivu.
h. THRYONOMYS C;REGORIANLS GREGORIANUS. Thomas
1894. Ann. Mag. Nat. Hist. 6, XIII, p. 202.
Luiji Reru River, Kiroyo, Kenya,
7. THRYONOMYS (;REC;ORIANUS PUSILLUS, Heller
191 2. Smiths. Misc. Coll. LIX, no. 16, p. 17.
Ndi, Taita Hills, Kenya.
S. THRYONOMYS HARRISONI HARRISON!, Thomas & Wroughton
1907. .Ann. Mag. Nat. Hist. 7, XIX, p. 384.
Lado (Anglo-Egyptian Sudan), Loka, 60 miles S.-\V. of Fort Berkeley.
9. THRYONOMYS HARRISONI CONGICUS, Thomas
1922. Ann. Mag. Nat. Hist. 9, IX, p. 390.
Uele River, Belgian Congo.
10 THRYONOMYS SCLATERI, Thomas
1897. Proc. Zoo!. See. London, p. 432.
Nvika Plateau, Nvasaland.
PETROMYINAE: PETROMUS i49
Addenda (gregorianus group):
THKYONOMYS I.OGONENSIS, Jeannin.
1936. Manim. Sauvagus du Camcroun, Encycl. Biol. 16, p. 178.
IJorders of Logone, Chad district, French Equatorial Africa.
J', nitshuricus, not seen, is described as a very short-tailed form, probably
nearest the gregorianus group. T. harrisoni has a narrower skull than in allies.
Subfamily PETROMYINAE
Geographical Distribution. — South-west Africa.
Number of Genera. — One.
Characters. — Cheekteeth rooted, but showing considerable simplification
of pattern; only one fold on each side in the upper series; the
internal side of the upper series and the external side of the lower series marked
by two elevations, the teeth strongly hypsodont. External form small, general-
ized except for the bushy tail. Bullae much inflated; skull flattened; mandible
tvpically Ilystricoid in formation.
Genus i. PETROMUS, Smith
1831. Petromus, Smith, South Afr. Quart. Journ. i, no. 5, p. 10.
Type Svzci^.— Petromus tvpicus. Smith.
Range. — As in the subfamily.
Number of Forms. — Four.
Characters. — Skull broad and flat, without any constriction in the inter-
orbital region ; infraorbital foramen with canal for nerve
transmission; bullae considerably inflated, the paroccipital processes joining
them; palatal foramina deep and long, well open, extending to toothrow; palate
extending slightly behind the toothrows, relatively narrow. Angular portion of
mandible slanting downwards posteriorly. Incisors opisthodont.
Cheekteeth as described above, the elevations clear and well marked, the
teeth set obliquely, the folds broad. Lower teeth with one fold on each side in
adult, as in the upper series. Pattern ultimately obliterated with wear.
External form more or less Rat-like except for the tail, which is bushy, and
not verv' much shorter than the head and body. Feet narrow, with short claws;
four main digits well developed on both fore and hindfeet, D.5 nearly as long
as the others; pollex vestigial, hallux short. Some stiff bristle hairs present on
hindtoes, as in Octodontinae.
The zygoma is relatively broad, sometimes with weak process on the lower
border. The mandible is clearlv distorted outwards in the angular process, like
typical llystricoids, but unlike Ctenodactylidae, with which this genus has been
associated; the coronoid is low, the angular portion drawn backwards.
Forms seen : tvpicus, trupicidis, cunealis.
cunealis was described as a species, but is probably best regarded as a race
as there seems very little essential difference between it and tvpicus.
Fig. 29. Petromus typicus typicus, Smith.
B.M. No. 12.4.25. 12 <S; • 2.
Fit;. 30. Petromus typiccs typicus, Smith.
B.M. No. 12.4.25. 12 d'; 2.
PETROMUS— ABROCOMINAE
151
Fiu. 31. Petromus typicus typiccs, Smith.
Mandible from below x 2; cheekteeth X 7; B.M. No. 12. 4. 25. 12, cJ-
1831
1923.
1935-
4-
1926.
List of Named Forms
(References and type localities by Mr. G. W. C. Holt.)
PETROMUS TYPICUS TYPICUS, Smith
South .■\fr. Quart. Joum. i, no. 5, p. 11.
Mouth of Orange River, South Africa.
PETROMUS TYPICUS TROPlC.'\LIS, Thomas & Hinton
Proc. Zool. Soc. London, p. 241.
Karibib, S.-\V. Africa.
PETROMUS TYPICUS MARJORI.A.E, Bradfield
Descr. new races of Kalahari Birds and Mammals, 2 pp. 1935.
Khan River, S.-W. .-Vfrica.
PETROMUS TYPICUS CUNEALIS, Thomas
Proc. Zool. Soc. London, p. 307.
CjLinene River Falls, S.-W. Ovamboland, S.-W. Africa.
Subfamily ABROCOMINAE
Geogr.^phical Distribution. — Northern Chile and Argentina.
Number of Genera. — One.
Ch.\r.\cters. — Cheekteeth evergrowing, the upper teeth simplified, each
tooth cut into two lobes by one wide re-entrant fold on each
side; lower teeth complex, with one outer, two inner deep folds, the spaces
152
ABROCOMINAE : ABROCOMA
caused by these folds sharply angular. Part of lachrymal canal open on side of
rostrum in front of orbit. Bullae greatly inflated. External form not highly
modified; tail haired, relatively short.
Rem.\rks. — Miller & Gidley refer this form to a distinct family; for discus-
sion ot the retention of the genus in the lichimyidae see page 103.
(Jenus I. ABROCOMA, VVaterhouse
1837. Abrocom.\, Waterhouse, Proc. Zool. Soc. London, p. 30.
Type Species. — Abrocoma bennetti, Waterhouse.
Range. — As in the subfamily Abrocominae.
Number of Forms. — Seven.
Ch.\r.'\cters. — Rostrum long and narrow, braincase round, not ridged,
frontals considerably constricted (tor a member of this
group). No canal for transmission of nerve in the intraorbital foramen. Palate
straight, short and relatively narrow; palatal foramina very long and narrow,
totally different from that of Octodontinae, hut reminiscent of the Chinchillidae.
Zygoma simple. Anterior ascending maxillary portion of zygoma extremely
narrow; jugal widely separated from the lachrymal. Bullae very large indeed.
Fig. 32. Abrocoma bennetti bennetti, Waterhouse.
B.M. No. 4.1.7.7, S\ y ■*■
Fic. 33. AsROCOMA BENNETTi BENNETTi, Waterhouse.
B.M. No. 4.1.7.7-, <J; ■' I*-
Fig. 34. Abrocom.\ bennetti bennetti, Waterhouse.
C'hecktiith ; B.M. No. 4.1.7.7., (J; X 7.
154 ABROCOMA
the mastoids appearing to a degree in superior aspect of skull. Incisors narrow.
Mandible with very narrow angular process drawn sharply backwards, a wide
curved space separating the condvle from the angular process; coronoid
low.
Lobes of upper cheekteeth united by a narrow bridge; M.3 with backwardlv
projecting heel caused bv a small extra outer fold. The folds are strong, and
rather wide. Lower teeth quite different in appearance from the upper series
(a rare feature in the Order); two inner, one outer folds; general pattern not far
removed from that present in Capromys, but effect very different, folds widely
open, not filled.
Externally, fur verv soft as a rule; tail usually short, but well haired, ear
relatively large. Forefoot short, with four digits, the claws small; hindfoot with
a reduced hallux, D.5 shorter than the three central digits. Claws weak. Some
stiff bristle-hairs present on the central digits of the hindfoot, as in Chinchillidae
and Octodontinae.
According to Waterhuuse the skeleton of A. bennetti bears a greater number
of ribs than is usual; the number quoted is seventeen pairs, which he compares
with Ciipromvs and Coendoii. in which genera sixteen are said to be present.
Two specific groups may be recognized, according to London material, the
members of which do not seem to me to be more than racially distinct from each
other.
The bennetti group contains relativelv larger forms (hindfoot 31-38), colour
more brown, posterior palatal foramina fused and conspicuous between tooth-
rows.
The cinerea group contains relatively smaller forms (hindfoot 31 or usually
less), colour grey, posterior palatal foramina not conspicuous between toothrows,
vestigial.
Forms seen : bennetti, biidini, cinerea, faiiuitina, inurrayi, schistacea, vaccarum.
List of Named Forms
(References and type localities by Mr. G. W. C. Holt.)
bennetti Group
1. .ABROCOMA BENNETTI BENNETTI, Watfrhouse
1837. Proc. Zool. Soc. London, p. 31.
Flanks of Cordillera, Aconcagua, Chile.
Synonym: cmieri. Waterhouse, 1S37, Proc. Zool. Soc. London, p. 32.
Valparaiso, Chile.
Iielviiia, Wagner, 1842. .Arch. Naturg. i, p. 7. Chile.
2. .ABROCOMA BENNETTI MURRAYI, Wolffsohn
igi6. Rev. Chilena, p. 6.
Vallenar, Province Atacama, Chile.
cinerea Group
3. ABROCO.AEA CINEREA CINEREA. Thomas
igig. .Ann. Mas. Nat. Hist. 9. IV, p. 132.
Casabindo Volcano, lujuy, North Argentina.
OCTODONTINAE 155
4. ABROCOMA CINEKEA BUDINI, Thomas
IQ20. Ann. Mag. Nat. Hist. 9, V, p. 475.
Otro Cerro, 18 kilometres north-west of Chumbicha, Catamarca,
Argentina.
5. ABROCOMA CINEREA FAM.ATINA, Thomas
1920. Ann. Mag. Nat. Hist. 9, VI, p. 419.
La Invcrnada. Rioja, Argentina (18 kilometres north-west of Nevada de
Famatina).
6. ABROCOMA CINEREA SCHISTACEA, Thomas
1921. Ann. Mag. Nat. Hist. 9, VIII, p. 216.
Los Sombreros, Sierra Tontal, San Juan, Argentina.
7. ABROCOM.\ CINEREA V.ACCARUM, Thomas
1921. Ann. Mag. Nat. Hist. 9, VIII, p. 217.
Punta de Vacas, Mendoza, Argentina.
Subfamily OCTODONTINAE
Geogr.\phic.\l Distribution. — Neotropical: Peru, Bolivia and Matte
Grosso southwards to Southern Patagonia.
Number of Genera. — Si.\.
Ch.\r.\cters. — Cheekteeth, both upper and lower, completely simplified,
with, in the upper series, one fold each side in less simplified
forms, general effect eight-shaped or "kidney-shaped." Bullae normally much
inflated. Xo part of lachrymal canal open on side of rostrum. E.xternally
generalized, or modified for subfossorial life. Some stiff bristle-hairs present
on toes of hindfeet.
The generalized forms include Octomys (tail thickly bushy, bullae largest in
subfamily, cheekteeth eight-shaped); Octodontomys (similar externally, but
bullae slightly smaller, and cheekteeth completely simple); and Octodon (tail
less bushy, bullae smaller than the above genera, and cheekteeth more or less
kidney-shaped in adult). The subfossorial types include Aconaemys (cheek-
teeth like Octomys, upper incisors not abnormal, bullae moderate in size);
Spalacopus (small forms with the upper incisor extending backwards and almost
overlapping the toothrows, bullae relatively small); and Ctenomvs (differing from
the above two genera in the much larger foreclaws, more heavily ridged skull,
particularly the zygoma, kidney-shaped cheekteeth, with vestigial M.3 ; the
bullae are relatively large; the incisors are not so extreme as in Spalacopus).
In this group, there is always an upwardly pointing process on the posterior
border of the jugal, which, however, varies in development, being most extreme
usually in Clenomys.
Key to the Gener.\ of Octodonti.v.ve
E.xternal form considerably modified for subfossorial life. Tail strongly
reduced, not much longer than hindfoot.
156 OCTODONTINAE: OCTOMYS
Cheekteeth simpler, kidney-shaped; skull more hea\ ilv ridfjed for
muscle attachment; foreclavvs strongly lengthened. (Incisor
root in upper scries not tending to overlap toothrow; bullae
relatively large.) Ctenomvs
Cheekteeth more complex, more or less eight-shaped, or completely
so; skull less heavily ridged for muscle attachment; foreclaws
less lengthened.
Re-entrant folds of upper molars not meeting in middle of the
teeth; upper incisorsstronglypro-odont.theirrootsextending
backwards, forming a projection by the side of and almost
overlapping the upper toothrow; bullae small (for the sub-
family). Spalacopus
Re-entrant folds of upper molars meeting in middle of the teeth ;
upper incisors less pro-odont, not abnormal. Bullae larger.
ACONAF.MYS
External form not modified for subfossorial life. Tail not much reduced,
little shorter than head and body (or may be longer than this
measurement). (Bullae strongly inflated.)
Re-entrant folds of upper molars meeting in middle of the teeth;
general dental effect clearly eight-shaped. (I'ail bushy; bullae
at maximum for subfamily.) Octomys
Re-entrant folds of upper molars not meeting in middle of the teeth ;
general dental effect not eight-shaped.
Cheekteeth completely simple, the folds obsolete. (Tail bushy;
bullae relatively larger.) Octodontomys
Cheekteeth not completely simple, the effect becoming kidney-
shaped, the folds not obsolete. (Tail less bushy; bullae
relatively smaller.) Octodon
Genus I. OCTOMYS, Thomas
1920. OcTO.MYS, Thomas, Ann. Mag. Nat. Hist. 9, VI. p. 117.
Type Species. — Octomys mimax, Thomas.
Range. — Argentina (Catamarca, San Juan).
Number of Forms. — Two.
Characters. — Skull without constriction in interorbital region, and not
prominently ridged for muscle attachment. Occipital region
relatively weak. Bullae large and much inflated, largest of subfamily, mastoids
visible in superior aspect of skull ; palate nearly straight, narrow, V-shaped
posteriorly, relatively short. Palatal foramina short, broad, with broad median
septum. Rostrum pointed, not shortened. Infraorbital foramen with a canal
OCTOMYS— ACONAEMYS
157
for nerve transmission. A capsule on mandible at root of M.2. Coronoid
process low.
Cheekteeth complex for the group; clearly eight-shaped, the folds meeting
in the middle of the tooth; .M.3 the smallest tooth.
External form not modified for fossorial life; fur very soft; tail about as long
as head and body or slightly longer, heavily haired. Claws small; forefoot with
four well developed digits and rudimentary pollex; hindfoot with hallux short,
D.5 shorter than the three central digits.
Forms seen : mimax, joanniiis.
I am listing "jotmnit4s" provisionally as a race, though I think it is very
probable that the two forms are based on the same animal.
Fig. 35. OcTOMYS mimax mimax, Thomas.
B.M. No. 20.5.1 1.32, ?; ■ 8.
List of N.\med Forms
(References and tvpe localities for all species of Octodontinae have been
collected by Mr. G. W. C. Holt.)
1. OCTOMYS M1M.\X .MIM.-\X, Thomas
1920. .\nn. Mag. Nat. Hist, q, VI, p. 118.
La Puntilla, Tinogasta, Catamarca, Argentina.
2. OCTO.MY.S MIM.XX JOAXNTLS, Thomas
1921. .Ann. Mag. Nat. Hist, g, VHI, p. 217.
Pedemal, San Juan, Argentina.
Genus 2. ACONAEMYS, Ameghino
1 89 1. AcoNAEMYS, Ameghino, Revista Argent, de Hist. Nat. i, p. 245.
1841. SCHIZODON, Waterhouse, Proc. Zool. Soc. London, p. 89. (Not of .Agassiz.)
isS ACONAEMYS— OCTODON
Type Sfecies. — Schizodoii Juscus, Waterhouse.
Range. — Southern Chile and Argentina.
Number of Forms. — Two.
Ch.\r.\cters. — Essentially like Oclomys dentally; cranially differing in the
rather shortened rostrum; the palatal foramina short, small
(infraorbital foramen with canal for nerve transmission); the bullae much smaller,
of medium size. Coronoid process moderate. Incisors rather broad. M.3
tends to have the posterior lobe reduced, both above and below.
Externally modified for fossorial life; arrangement of digits about as in
Octomys; claws enlarged to a degree; ear moderate-sized, not strongly reduced;
tail haired, not much longer than hindfoot.
Remarks. — This genus is very much the most generalized of the three
fossorial Octodontinae, both in external and dental characters.
The two species are closelv allied, and differ chiefly in the quality of the fur.
Forms seen : fiiscus, porteri.
List of Named Forms
1. ACONAF.MYS FUSCUS, Waterhouse
1841. Proc. Zool. Soc. London, p. gi.
Valle de las Cuevas, eastern slope of .Andes, .Argentina.
2. ACONAEMYS PORTERI, Thomas
1917. .Ann. Mag. Nat. Hist. S, XIX, p. 281.
Osomo, Southern Chile.
Genus 3. OCTODON, Bennett
1832. OcTODON, Bennett, Proc. Zool. Soc. London, p. 46.
Type Species. — Octudoii cinniin^ii, Bennett.
Range. — Chile and Peru.
Number of Forms. — Six.
Ch.\racters. — Skull essentially like that of Octomys; the weak parietal
ridges mav come together in old age; bullae relatively smaller
than in either Octomvs or Octodontoinxs; infraorbital foramen with canal for
nerve transmission.
Cheekteeth becoming modified and transitionary towards those of Cteiiomys;
anterior part in upper series projecting outwards; lower teeth with posterior
part pointing inwards. The upper teeth have a small inner fold retained; the
lower teeth are more eight-shaped than in Ctenomys, but much less so than
in Octomvs and Aconaemys. When cut, though simple, they are nearer the
Octomys type; the folds of the upper series nearly meet, though the adult
pattern is suggested already.
External form somewhat Rat-like; fur typically much less soft than in
Octodoutom\s and Octomys. Ear large: tail rather shorter than head and body,
OCTODON— OCTODONTOMYS 159
less wfll haired than in Octomys, scales traceable; moderately haired except at
end, which is slightly bushy. O. bridgesii is a softer-furred form.
Forms seen : bridgesii, ciivorum, degus.
Waterhouse synonymized pallidus and cumingii with degus; they are pro-
visionally listed here as subspecies, though I have seen neither, and they may
prove either synonymous or valid.
List oh Named Forms
1. OCTODON" DEGUS DEGUS, Molina
1782. Sagg. Storr. Nat. Chili, ist Ed., p. 303.
Chile.
2. OCTODON DEGUS CMVORUM, Thomas
1927. .-Xnn. Mag. Nat. Hist. 9, XIX, p. 556.
Puente Alto, Santiago, Chile.
3. OCTODON DEGUS PERUANA, Waterhouse
1848. Nat. Hist. Mammalia, ii, p. 257.
San Juan de Matucana, Lima, Peru.
4. OCTODON DEGUS CUMINGII, Bennett
1832. Proc. Zool. Soc. London, p. 47.
Between Valparaiso and Santiago, Chile.
5. OCTODON DEGUS PALLIDUS, Wagner
1S45. Arch. Naturg. 2, p. 33.
Chile.
6. OCTODON BRIDGESII, Waterhouse
1844. Proc. Zool. Soc. London, p. 155.
Chile.
The name franzitisi listed by Trouessart in this genus (Cat. Mamm. viv.
foss. 1904, Suppl., p. 500), is according to Tate a Geomyid.
Genus 4. OCTODONTOMYS, Palmer
1902. Neoctoix)n, Thomas, Proc. Zool. Soc. London, i, p. 114 (pre-occupied).
1903. OCTODONTOMVS, Palmer, Science, 2, XVH, p. 873.
Type Species. — Xeoctodon simonsi, Thomas = Octodon gliroides, Ger\ais &
D'Orbignv.
R.\NGE. — Bolivia.
Number of Forms. — One.
Char.'^cters. — Cheekteeth simpler than in Octodon, the folds obsolete, a
slight concavitTr- on outer side of upper molars; lower molars
with slight median constriction each side. Skull essentially as in Octomys e.vcept
the relatively smaller bullae, which, however, are larger than those of Octodon.
Infraorbital foramen with canal for nerve transmission.
Essential external characters as Octomys; tail thickly bushy, fur very soft.
Forms seen : sliroides.
i6o
OCTODONTOMYS— SPALACOPUS
List of Named Forms
1. (K-TDDONTOMYS GLIROIDES, Gen-ais & D'Orhigny
1S44. Bull. Soc. Philom. p. 22.
Bolivian .Andes, near La Paz.
Synonym: simoiisi, Thomas, 1902, Prnc. Zool. Soc. London, i, p. 115.
Potosi, Bolivia.
Genus 5. SPALACOPUS, Wagler
1S32. Sp.\l.\copl's, Wagler, Isis, XXV, p. 1219.
Type Species. — Spalacupiis poeppigi, Wagler.
Range. — Chile.
Number of Forms. — Two.
Characters. — Skull with the same essential characters as Octomvs, but
frontals appear narrower, and upper incisors strongly pro-
odont, and much lengthened, extending backwards to a level of about M.i
and forming a projection by the side of and almost overlapping the toothrow.
Fk;. 36. Spalacopus cvanus, Molina.
B.M. No. 1. 3. 21. 14, o; 2J.
Bullae smaller than in all other Octodontinae, not much inflated. Palatal
foramina small. Intraorbital foramen with no separate canal tor nerve trans-
mission. Coronoid process prominent.
Cheekteeth eight-shaped, but the folds not meeting in the middle of the
SPALACOPUS— CTENOMYS i6i
tooth, the general effect rather simpler than Oclomys; M.3 above and below
smaller, simpler.
Kxtcrnally typically smaller than other genera, about the smallest living
Hystricoid Rodent. Colour very dark. Considerably modified for fossorial
life; tail short, hairy, little longer than hindfoot; claws not greatly enlarged.
Arrangement of digits about as Octomys. Ear small.
Remarks. — The broad abnormally lengthened upper incisors differentiate
this genus clearly from all allies, and notwithstanding its small
size it may be considered one of the most specialized of the group. The teeth
too are more simplified than in Octomys and Aconaemys, but the smaller bullae
suggest a more generalized character.
■S\ tabanus appears to represent a larger form than the type, but is not well
known.
Forms seen : cvanus, tabanus.
List of Named Forms
1. SPALACOPUS CYANUS, Molina
1782. Sagg. Stor. Nat. Chili, ist Ed. p. 300.
Chile.
S>Tionym: poeppigi, Wagler, 1832, Isis, XXV, p. 1219. Quintero, Rio
Aconcagua, Chile.
ater, Cuvier, 1834, Ann. Sci. Nat. i, p. 323. Coquimbo.
noctivagus, Poeppig, 1835, Arch. Naturg. i, p. 252. Quin-
tero, Rio Aconcagua, Chile.
2. SPALACOPUS TABANUS, Thomas
1925. Ann. Mag. Nat. Hist. 0, XV, p. 585.
South Chile.
Genus 6. CTENOMYS, Blainville
1826. Ctenomys, Blainville, Bull. Soc. Philom. p. 62.
1916. Haptomys, Thomas, Ann. Mag. Nat. Hist. 8, XVUI, p. 305; subgenus for C.
leucodon, Waterhouse.
Type Species. — Ctenomys brasiliensis, Blainville.
Range. — South Brazil (Matto Grosso), Bolivia, Paraguay, Argentina (Buenos
Ayres region, Jujuy, Salta, Tucuman, Catamarca, San Juan,
Cordoba, Mendoza, etc.), Patagonia south to Tierra del Fuego; Chile.
Number of Forms. — Approximately sixty-one are named.
Characters. — Skull with broad rostrum, postorbital process usually present
to frontals, their development variable; parietals well ridged,
though evidently most often a sagittal crest is not formed; lambdoid crest
prominent; bullae large, pear-shaped, spread sideways; paroccipital processes
large, curved under them (the bullae show prominently on each side when skull
is viewed from behind). Palate essentially as in other Octodontinae; palatal
foramina usually short; jugal with extremely prominent upwardly projecting
process in larger forms; this process always present, usually well developed.
II — Living Kodents — i
Fig. 37. Ctenomys tuconax, Thomas.
B.j\l. No. 25.3,1.19, 'i; X iJ.
Fig. 3S. Ctenomy,s tucona.\, Thomas.
B.M. No. 25.3.1.19, 5; X iS.
CTENOMYS 163
Infraorbital foramen with no canal for nerve transmission. Upper incisor root
extending far backwards, and showing on inner border of infraorbital foramen,
though not so extremely as in Spalacopiis. Mandible with angular processes
widely spreading, sharply distorted outwards; coronoid process moderate.
Incisors much thickened, usually not pro-odont, except in leucodon and
lewisi. Cheekteeth like Octodon, but simpler, the small inner fold obsolete in
the upper molars. iM. I vestigial.
Fig. 39. Ctenomys tuconax, Thomas.
Mandible from below, X ij; Cheekteeth, X 6: B.M. No. 25. 3.1. 19, §.
Externally much modified for subfossorial life; eyes and ears reduced;
forefoot with extremely large claws (pollex less reduced than is normal) ; hindfoot
with moderate claws; hallux rather less reduced than in other Octodontinae,
otherwise general arrangement of digits like allied genera ; tail strongly shortened
though not vestigial, moderately or poorly haired.
Forms seen : antonii, azarae, barbariis, hergi, boliviensis, budini, coludo,
dorsalis, emiliamis, fochi, fodax, frater, fuegiiius, fiimosiis, fulviis, goodfellowi,
liaigi,johamiis, juris, knighti, latru, lentulus, leucodon, leivisi, luteolus, niagelhmicus,
mendocina, mordosus, nigriceps, occidtus, opimiis, perrensi, pontifex, porteousi,
recessus, saltarius, sericeiis, steinbuchi, syhamis, talariim, torquatus, tuconax,
tucutnanus, tulduco, utibilis, viperinus.
'I'his genus is undoubtedly in great need of revision. The forms seem ex-
tremely closely allied to each other, generally speaking, though most of them are
1 64 CTENOMYS
standing at present as distinct "species." There is great difference in size be-
tween some of the forms, emilianus, tuconax, and iiigiiceps having a hindfoot
measurement of 3S mm. and terms like recessiis and occuliiis only 26 mm. But
intermediate forms exist between both e.xtremes, so that all hindfoot measure-
ment figures exist within the genus between the figures 38 mm. and 26 mm.
leucodon and lewisi, the latter described as semi-aquatic, have more pro-odont
upper incisors than the others. Three large Bolivian types, boliviensis, good-
fellowi and steinbachi, appear to have a skull which is broader than normal,
particularly in the region of the muzzle.
The genus has been reviewed by Rusconi, 1928 (Anal. Soc. Arg. Geogr.
"Gaea," III, p. 235), who shows the subgenus " Haptomys" to be no longer
retainable.
I propose lor the purposes of the present work to divide the genus into
sections. Xo attempt is made to reduce forms to subspecies, the genus being
far too big for a revision to be attempted in the present work; undoubtedly very
many "species" now standing will ultimately be regarded as races. There
are many forms not represented in the British Museum, though in the case of
those that have been seen, except in verv few cases, a large and representative
series of skins have been examined.
So far as Patagonia is concerned, on British Museum material, there are two
well-marked groups, very small types like magellaiiiciis, and very large types like
fodax present only. But elsewhere, there are the "small," "medium," and
" large " sections living apparently more or less side by side, the measurements of
which grade into each other.
The sections here recognized are as follows, though it must be borne in mind
that the plan followed here is no more than provisional, and an attempt to get
some order out of considerable chaos.
1. magellaiiiciis section: small forms, smallest of genus; hindfoot usually
under 30, rarelv exceeding this measurement, never more than 32;
often 24, 25, 26.
2. torquatus section: moderate-sized forms, not becoming very large; hind-
foot rarely under 30, never less than 28, usually measurement 31-35 ;
never more than 37.
3. opimus section: like the last, but becoming large, approaching maximum
for the genus; hindfoot usually over 36, not under 35 excepting one
race of opimus {luteolus), which agrees with the larger members of
section 2. At maximum, hindfoot up to 48 (fodax); in others as a
rule not more than 39.
4. boliviensis section : agreeing in measurement with the last, but skull
unusually broadened, particularly in the muzzle region. (Bolivia:
boliviensis, goodfellowi, steinbachi.)
5. leucodon section : incisors strongly pro-odont ; hindfoot measurement about
30 (not many seen).
6. /f««/ section : incisors also pro-odont. hindfoot measurement 32-37;
water-side dwelling type. Thomas suggested that this was not a near
ally of leucodon.
CTENOMYS 1 6s
List of Named Forms
magellanicus section
1. CTENOMYS HAIGI HAIGI, Thomas
1919. Ann. Mag. Nat. Hist. 9, III, p. 210.
Maiten, Western Chubut, Argentina.
2. CTENOMYS HAIGI LENTLI.US, Thomas
1919. Ann. Mag. Nat. Hist. 9, III, p. 211.
Pilcaneu, Upper Rio Negro, Argentina.
3. CTENOMYS SERICEUS, Allen
1903. Bull. Amer. Mus. XIX, p. 187.
Cordilleras, upper Rio Chico de Santa Cruz, Patagonia.
4. CTENOMYS MAGELL.-VNICUS, Bennett
1835. Proc. Zool. Soc. London, p. 190.
Port Gregory, Straits of Magellan.
Synonym: neglectus, Nehring, 1900, Zool. Anz. XXIII, p. 535. Pata-
gonia.
5. CTENOMYS TALARUM TALARUM, Thomas
1898. Ann. Mag. Nat. Hist. 7, I, p. 2S5.
Los Talas, Ensenada, La Plata, Argentina.
6. CTENOMYS TALARUM ANTOMI, Thomas
1910. Ann. Mag. Nat. Hist. 8, V, p. 242.
Los Yngleses ranch, Ajo, eastern Buenos Ayres, Argentina.
7. CTENOMYS TALARUM RECESSUS, Thomas
1912. Ann. Mag. Nat. Hist. 8, IX, p. 241.
Bahia Blanca, .Argentina.
8. CTENOMYS MENDOCINA, Phihppi
1869. Arch, fiir Naturg. p. 38.
Mendoza, Argentina.
9. CTENOMYS PONTIFEX, Thomas
1918. .-Xnn. Mag. Nat. Hist. 9, I, p. 39.
East side of Andes, Province of Mendoza, Argentina (near Fort San
Rafael).
10. CTENOIVrVS BERGI, Thomas
1902. -Ann. Mag. Nat. Hist. 7, IX, p. 241.
Cruz de Eje, Cordova, .Argentina.
11. CTE.NOMYS FOCHI, Thomas
1919. .Ann. Mag. Nat. Hist. 9, III, p. 117.
Chumbicha, Catamarca, Argentina.
12. CTENOMYS TUCUMANUS, Thomas
1900. .Ann. Mag. Nat. Hist. 7, VI, p. 301.
Tucuman, Argentina.
13. CTENOMYS L.ATRO, Thomas
1918. Ann. Mag. Nat. Hist. 9, I, p. 38.
Tapia, Tucuman, .Argentina.
i66 CTENOMYS
14. CTENOMYS ()CCL:i/rL'S, Thomas
1920. Ann. Mat;. Nat. Hist, g, VI, p. 243.
Monteagudo, 80 kilometres south-east of Tucunum City, Argentina.
15. CTENOMYS SALTARIUS, Thomas
1912. .Ann. Mag. Nat. Hist. 8, X, p. 639.
Salta. Northern Argentina.
ih. CTENOMYS JURIS, Thomas
1920. Ann. Mag. Nat. Hist. 9, V, p. 194.
El Chaguaral, Jujuy, Argentina, 20 kilometres east of San Pedro de
Jujuy, between San Pedro and Villa Carolina.
17. CTENOMYS DORSALIS, Thomas
igoo. Ann. Mag. Nat. Hist. 7, VI, p. 385.
Northern Chaco, Paraguay.
torqiiatiis section
18. CTENOMYS PERRENSI. Thomas
1896. Ann. Mag. Nat. Hist. 6, XVIII, p. 311.
Goya, Corrientes, Argentina.
ig. CTENOMYS AZAR.'^E, Thomas
1903. .Ann. Mag. Nat. Hist. 7, XI, p. 228.
37 45' S., 65' W., 780 kiloinetres south-west of Buenos Ayres, Buenos
.Ayres Province, Argentina.
20. CTENOMYS PORTEOUSI PORTEOUSI, Thomas
191 6. .Ann. Mag. Nat. Hist. 8, XVIII, p. 304.
Bonifacio, South-west Buenos .Ayres, Argentina.
21. CTENOMYS PORTEOUSI AUSTRALIS, Ruscom
1934. Rev. Chili. Nat. Hist. 38, p. 108.
Province Buenos Ayres, Argentina.
22. CTENOMYS TULDUCO, Thomas
1921. .Ann. Mag. Nat. Hist. 9, VIII, p. 218.
Los Sombreros, Sierra Tontal, .San Juan, Argentina.
23. CTENOMYS FAMOSUS, Thomas
1920. Ann. Mag. Nat. Hist. 9, VI, p. 420.
Potrerillo, Rioja, Argentina.
24. CTENOMYS COLUDO COLUDO, Thomas
1920. Ann. Mag. Nat. Hist, g, VI, p. 119.
La Puntilla, Tinogasta, Catamarca, Argentina.
25. CTENOMYS COLUDO JOHANNIS, Thomas
1921. .Ann. Mag. Nat. Hist. g. VII, p. 523.
Caiiada Honda, San Juan, Argentina.
26. CTENOMYS VIPERINUS, Thomas
ig26. Ann. Mag. Nat. Hist, g, XVII, p. 605.
Tablelands above Norco, Vipos, Dept. of Trancas, Tucuman, Argen-
tina.
27. CTENOMYS SYLVANUS SYLVANUS, Thomas
igig. .Ann. Mag. Nat. Hist. g. IV, p. 155.
Tartagal, Pro\ince Salta, .Argentina.
CTENOMYS 167
28. CTENOMYS SYLVANUS UTIBILIS, Thomas
1920. Ann. Mag. Nat. Hist. 9, V, p. 193.
Yuto, Rio San Francisco, Argentina, 20 kilometres east of San Pedro
de Jujuy.
29. CTENOMYS SYLVANUS MORDOSUS, Thomas
1926. .'Xnn. Mag. Nat. Hist. 9, XVH, p. 325.
Tambo, 75 kilometres cast of Tarija, Bolivia.
30. CTENOMYS DUDINI BUDINI, Thomas
1913. .\nn. Mag. Nat. Hist. 8, XI, p. 141.
Cerro de Lagunita, Jujuy, Argentina.
31. CTENOMYS BUDINI BAKBARUS, Thomas
1 92 1. .Ann. Mag. Nat. Hist. 9, VII, p. 185.
Sunchal, Jujuy, ;\rgentina.
3=. CTENOMYS FRATER, Thomas
1902. Ann. Mag. Nat. Hist. 7, IX, p. 228.
Potosi, Bolivia.
33 CTENOMYS TORQUATUS, Lichtenstein
1830. Darstell. Saugethiere, text of PI. XXXI.
Southern Provinces of Brazil and banks of Uruguay River.
leucodon section
34. CTENOMYS LEUCODON, Waterhouse
1848. Nat. Hist. Mammalia, II, p. 281.
San Andres de Machaca, Bolivia (Dept. of La Paz).
lewisi section
35. CTENOMYS LEWISI, Thomas
1926. Ann. Mag. Nat. Hist. 9, XVII, p. 323.
Sama, 50 kilometres west of Tarija, Bolivia.
opimiis section
36. CTENOMYS EMILIANUS, Thomas & St. Leger
1926. Ann. Mag. Nat. Hist. 9, XVIII, p. 637.
Chos Malal, Neuquen, Argentina.
37- CTENOMYS FODAX, Thomas
1910. Ann. Mag. Nat. Hist. 8, V, p. 243.
Valle de Lago Blanco, Chubut, Patagonia.
3S. CTENOMYS FUEGINUS, Philippi
1880. Arch, fiir Naturg. p. 276.
Eastern Island of Tierra Del Fuego.
39. CTENOMYS FULVUS, Philippi
i860. Reise. .Atacama Halle, p. 157.
Desert of .Atacama, Chile.
40. CTENOMYS KNIGHTI, Thomas
1919- Ann. Mag. Nat. Hist. 9, III, p. 498.
Otro Cerro, 45 kilometres west of Chumbicha, Catamarca, Argentina.
1 68 CTENOMYS
41. CTENOMYS TLCONAX, Thomas
1925. Ann. Mag. Nat. Hist, g, XV, p. 583-
Concepcion, Tucuman, Argentina.
42. CTENOMYS OPIMUS OPIMUS, Wagner
1S4S. Archiv. fur Naturg. i, p. 75-
Bolivia.
43. CTENOMYS OPIMUS NIGRICEPS, Thomas
1900. .Ann. Mag. Nat. Hist. 7, VI, p. 383.
Tetiri, Puno Moquegua Road, South Peru.
44. CTENOMYS OPIMUS LUTEOLUS, Thomas
1900. Ann. Mag. Nat. Hist. 7. VI, p. 384-
Cordilleras of Jujuy. .Argentina.
bolivieiisis section
45. CTENOMYS BOLIVIENSIS, Waterhouse
1848. Nat. Hist. Mammalia, ii, p. 278.
Plains of Santa Cruz de la Sierra, Bolivia.
46. CTENOMYS STEINB.ACHI. Thomas
1907. .Ann. Mag. Nat. Hist. 7, XX, p. 164.
Campo of Province Sara, Bolivia.
47. CTENOMYS GOODFELLOWI, Thomas
1921. .Ann. Mag. Nat. Hist. 9, VII, p. 136.
Esperanza, Concepcion, Eastern Bolivia.
\ot seen, and not allocated to section
48. CTENOMYS BRASILIENSIS, BlainvillL-
1826. Bull. Soc. Philom. p. 62.
Minas Geraes, Brazil. (Waterhouse treats torquatus, number 33, as a
synonym of this species.)
49. CTENOMYS OSGOODI, Allen
1905 Report Princetown Univ. Exped. to Patagonia, p. 191-
Rio Chico de Santa Cruz, Patagonia. „ „ ,^ x- . u- .
Svnonvm: robustus, Allen, not of Philippi, 1903. Bull. Mus. Nat. H,st.
XIX, p. 185. Patagonia. (According to measurements
from description, this species will belong in the moRel-
lanictis section.)
50. CTENOMYS COLBURNI, Allen
IQ03. Bull. Amer. Mus. Nat. Hist. XIX, p. 18S. „ . •
' ^ Arroyo Aike, 50 miles south-east of Lake Buenos Ayres, Patagonia.
(According to measurements from description this species probably
belongs in magcUanicus section.)
51. CTENOMYS MINUTCS, Nchring
1 887. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 47. , j c 1 n 1
"Campos," East of Mundo Novo, Rio Grande do bul, Brazil.
52. CTENOMYS RONDONI, Ribeiro
1914. Comm. Linhas, Tel. Annexo, 5, p. 39.
Juruena, Matto Grosso. Brazil.
CTENOMYS 169
53. CTENOMYS BICOLOR, Kibtiro
1914. Comm. Linhas. Tel. Annexo, 5, p. 41.
Matto Grosso, Brazil.
54. CTENOMYS NATTERERI, Wagner
1848. Archiv. fiir. Naturg. i, p. 72.
Caissora, Matto Grosso, Brazil.
55. CTENOMYS PUNDTI, NL-hrinc
1900. Zool. Anz. XXIII, p. 420.
Alejo Ledensa, Cordova, Argentina.
56. CTENOMYS ATACAMENSIS, Philippi
i860. Reise. Atacama Halle, p. 157.
Desert of .Atacama, Chile.
57. CTENOMYS ROBUSTUS, Philippi
1896. An. Mus. Nac. Chile, no. 13, p. 11.
Canchones, near Pica, Tarapaca, Chile.
58. CTENOMYS PALLIDUS, Philippi
1896. An. Mus. Nac. Chile, no. 13, p. 13.
Breas, desert of Atacama, Chile.
59. CTENOM\'S PERNIX, Philippi
1896. An. Mus. Nac. Chile, 13, p. 15.
Near Aguas Calientes, Chile.
60. CTENOMYS CHILENSIS, Philippi
1896. An. Mus. Nac. Chile, 13, p. 16.
Linares, Chile.
61. CTENOMYS MAULINUS, Philippi
1872. Zeitschr. f. ges. Naturw. XL, p. 442.
High Andes of Province of Maule, Chile.
Tate lists also a "cinerea" Thomas, which is evidently a mistake for
Abrocoma cinerea, Thomas.
The family Echimyidae contains according to Miller & Gidley very many
Neotropical fossil genera. The Octodontinae are quoted from the Oligocene;
one of the genera, Cephalomys, had a deciduous P. 4 (Gregory, Orders of
Mammals, 1910), a character not known in living Hystricoids; the Echimyinae
(with which Miller & Gidlev include Capromyinae and Dactviomyinae) are
quoted from the Miocene; some of the genera, as Isobolodoti (Porto Rico),
Brotomys (Dominican Republic), and Boromys (Cuba), are thought to have
existed recently. The Thryonomyinae have been described from the Miocene
of India.
ECHIMYIDAE:
GENERAL IVORKS OF REFERENCE
VVaterhouse, 1848, Natural Histor>- of Mammalia: Rodentia. General review of all
forms then known.
T.^TE, 1935, Ta.\onomv of Neotropical Hystricoid Rodents, BiJI. Amer. Mus. Nat. Hist.
LXVIII, p. 295.
PococK, Proc. Zool. Soc. London, 1922, p. 365; external characters of Hystricomorph
Rodents (notes on Octodon, Capromys, Myocastor, Daclylomys, Ctenomys, Thryo-
nomys).
I70 DINOMYIDAE
TuLLBERc, Nova Acta Reg. Soc. Sci. Upsaliensis, XVIII, ser. 3, no. i, iSgg.
Ch.\pm.^,\', iqoi, Bull. Amer. Mus. Nat. Hist. vol. XIV, p. 313. Revision of Hutias
(Caproniyinae).
RuscoNi, Review of Ctenomys, 1928, An. Soc. Arg. Geogr. "Gaea," III, p. 235.
Thom.\s, races of Thryoiiomys siLinderiiDuis, Ann. Mag. Nat. Hist., 9, IX, p. 392. 1922.
Miller, Proc. U.S. Nat. Mus, LXXII, no. 16, p. 4, 1927 (Plaaiodoiitui hylaeum).
Jentink, Notes Leyden Museum, XIII, 1891, p. 105. On Dactylomys dactyUnus and
Kannahateomys amhlyouyx.
And numerous papers by Oldficld Thomas (Echimyinae, Octodontinae).
Family DINOMYIDAE
1896. Thomas: Hystricomorpha; Family Dinomyidae.
1899. TuUberg; Hystricomorpha; (?) Family Dinomyidae.
1918. Miller S: Gidley: Hystricoidae; Family Dinomyidae.
1924. Winge: Family Hystricidae; Dasyproctini, part, group Dinomyes.
1928. Weber: Hy'stricoidea; Family Caviidae, part, subfamily Dinomyinae.
Geogr.^phical Distribution. South America; Peru, Colomhia, Ecuador
and Western Amazonia.
Number of Genera. — One.
Characters. — Cheekteeth extremely hypsodont, or probably evergrowing, a
series of transverse plates. External form heavy, terrestrial;
forefeet and hindfeet with four digits, the feet broad, the claws long and power-
ful. Limbs not lengthened. Palate constricted anteriorly. Zygomasseteric
structure typically Hystricoid, as regards the formation of the lower jaw.
Remarks. — Except by those authors who merge Cunicidus and Dasyprocta
with the Caviidae and who have regarded this genus also as a
member of that family, Diiiomys has usually been regarded as an isolated type
among Hystricoidae. There is not the slightest reason to suppose that the
animal is near the Caviidae, the lower jaw being typically Hystricoid in forma-
tion, and therefore differing from that family; nor does the genus seem closely
connected either with Ciinicnhis or Dnsyprocta, differing from botli in tooth
formation as well as the feet and digits. The palate and cheekteeth are similar
to those of the Chinchillidae, but from these Dinomys differs by its typically
ridged and distorted angular portion of the mandible, the general external form,
the absence of part of the lachrymal canal open on the side of the rostrum, as
well as by no tendency to great inflation of bullae.
Goeldi in a paper on some captivity specimens states that the animals are
slow-moving, unlike Dusvpioctii and the Chinchillidae. He mentions the fact
that like Dasyprocta but unlike Cunicuhis they will sit up on their haunches
and use the front paws when feeding. The claws on dried skins of Dinomys
appear to be fossorial in type, but Goeldi states that he has not seen the captivity
specimens use the claws for digging.
The breadth of the manubrium has been used as a character to distinguish
this genus as a family or subfamily from Dasyproctidae or Caviidae (Winge and
others); it should be noted that this character, according to Tullberg's notes,
may vary within some of the other families.
DINOMYIDAE: DINOMYS 171
Long and Narrow Broad
Laiiostomus (Chinchillidac) Chinchilla (Chinchillidae)
Dolichotis (Caviidac) Cavia (Ca\iidae)
It is stated to be long and narrow in Dasyprocta and Citnictdus, broad in
Dinomys.
'l"he clavicles in Dinomys are stated to be complete.
Genus i. DINOMYS, Peters
1873. Dinomys, Peters, Mon. Ber. Ak. Wiss. Berlin, p. 551.
TvPH Species. — Dinomys branickii, Peters.
Range. — As in the family Dinomyidae.
Number of Forms. — One only is now recognized; revised by Sanborn,
1931, Field. Mus. N.H., zool. ser. XVIII, p. 149.
Characters. — Skull heavy and broad, with long broad frontals; the parietals
are depressed for muscular attachment, but a sagittal crest
is not formed in any of the few skulls examined. No separate canal in infra-
orbital foramen for nerve transmission. Bullae medium sized; paroccipital
processes not lengthened. Jugal long, broad, but evidently simple. Palate of a
similar type to that found in Chinchillidae, but mesopterygoid fossa much
broader; the palate is continued farther backwards, to slightly behind the tooth-
rows. Palatal foramina small. Lachrymal large.
Incisors broad and heavy; cheekteeth a series of transverse plates; four of
these in each upper tooth ; four evidently in the lower teeth, but the anterior one
vestigial.
Externally large, heavy, bearing a superficial resemblance to Cunicubis; but
tail longer than hindfoot (fully haired). Hindlimbs not lengthened; the feet
broad, the claws long and heavy; no great discrepancy between the lengths of the
(four) digits; forefoot with four digits, the claws large and powerful, though
apparently narrower than in Cuniculus.
The genus is not well represented at the British Museum.
Forms seen: branickii, " occidenialis."
List of Named Forms
(The references and type localities are the work of Mr. G. W. C. Holt.)
I. DINOMYS BRANICKII, Peters
1873. Mon. Ber. .\kad. Berlin, p. 552.
Central Peru; Montana de Vitoc, Colonia .Amable Maria.
Synonym: branickii occidentalis, Lonnberg, 1921, .■\rk. Zool. XIV, no. 4,
p. 49. Ilambo, near Gualea, Ecuador.
gigas, .Ajithony, 1921, .Amer. Mus. Nov. no. 19. p. 6.
Colombia.
pacarana, Ribeiro, 1919, Arch. Escola Sup. Agric. Med.
Vet. 2, p. 13. .\mazon, Brazil.
Fig. 40. DiNOMVS br^nickii, Peters.
B.M. No. 34. 9. 10. 191, o; • 1-
Fig. 41. DiNOMYS branickii, Peters.
B.M. No. 34.9.10.191, cJ; •, i'.
ERETHIZONTIDAE 173
The family Dinomyidae as defined by Miller & Gidley (Lachr^-mal canal
closed in front of orbit; like the Echimyidae, but cheekteeth combining a
multilaminar structure with excessive hypsodonty), is known fossil from the
Miocene of South America and the Greater Antilles; many extinct genera are
quoted by these authors.
Family ERETHIZONTIDAE
iSq6. Thomas: HvsTRiroMORPHA, part; Family Erethizontidae, with subfamilies
Erethizontinae and Chaetomyinae.
1899. TullbtTg: HvsTRicoMOBPHA, part; Family Erethizontidae.
1Q18. Miller & Gidley: Superfamily HvsTRicom.^E; Family Erethizontidae. Family
Echimyidae, subfamily Echimyinae, part (Chaetomys).
1 924. Winge: Family Hystricidae; Hystricini, part; "Sphinguri."
1928. Weber: HvsTRlcomEA, part; Family Erethizontidae.
Geographical Distribution. — America; Canada, Western United
States; Mexico, Central America, and
the greater part of Tropical South America.
Number of Genera. — Four.
Characters. — Not essentially different from the Echimyidae, but externally
more highly specialized ; feet becoming abnormally modified
for arboreal life ; function of hallux being taken over in specialized forms by a
broad movable pad, the sole becoming abnormally wide; body hair modified
partly or completely into short sharp spines. Bullae prominent, but paroccipital
processes not lengthened. Cheekteeth rooted, typically with the re-entrant folds
extremely wide; external form thickset, heax-y.
Remarks. — Presumably because these animals are also known as "Porcu-
pines," or because their fur is spiny, most earlier authors placed
them in the family Hystricidae. Thomas very properly formed a distinct family
for them, and most subsequent authors have retained the distinction. TuUberg
states that there is hardly a single common feature between the Old World
and New World Porcupines except the spines, and even these are of a consider-
ably different structure. The two families differ entirely in the structure of the
feet, the structure of the cheekteeth, tlie formation of the bullae, the structure
of the tail; even in the essential arrangement of spiny covering. They agree in
zygomasseteric structure, which proclaims them both members of the Hystri-
coidea, but this seems about all they have in common. In fact, it would seem
from cranial and dental characters, at least, that the American representatives of
the Hvstricidae (if that family has American representatives, and the resemblance
is not due to convergence) are the Dasyproctidae; certainly not the present
group.
In Erethizontidae the paroccipital processes are less lengthened and evidently
of a more generalized structure than in Echimyidae. The zygoma is simpler
than in that family. The cheekteeth of the typical subfamily, which contains
Erethizon, Echinoprocta, and Coendou, and which has been incorrectly split into
174 ERETHIZONTIDAE: CHAETOMYINAE
two subfamilies bv some authors (see notes on Echinoprocta below), are remari<-
able for the width of their reentrant folds, paralleling in this formation certain
Squirrels as Fiiiiischinis, also the Anomaluridae, and to a degree reminiscent of
some of the more complex-toothed Neotropical Cricetinae.
Chcietomys, on the other hand, has teeth more like those of Echimys. This
is an isolated type, the relationships of which are by no means clear, so that
it might be quite correct to refer it to a distinct family Chaetomyidae. Agreeing
with most specialized Erethizontinae in the structure of the feet, it differs to a
very wide degree from them in cranial and dental characters. The orbit is
almost completely surrounded by bone, a very rare feature in the Order; and
in no member of the Order which I have seen is this specialization so nearly
complete. Moreover, the teeth are not in the least like those of Erethizon and
Coendou. It is a rare genus, the exact locality of which I have so far been unable
to trace, and evidently little is known about it. The spiny covering of the body
is very poorly developed compared with other members of the family. Miller
& Gidley transferred it to the Echimyidae, but it seems not to belong there
in cranial characters, and the feet are as highly specialized as in Coendou, and
evidently in exactly the same manner.
Thomas expressed the opinion that mainly on this accoimt it might be re-
tained in this family, and formed a subfamily for its reception. This view is
here adopted.
Key to the Subfamilies of F-Irethizontidae
Orbit almost surrounded by extremely thickened jugal and short post-
orbital process of frontals. Cheekteeth with narrow re-entrant
folds, the structure of the upper series not far removed from
laminate. Subfamily Chaetomyinae
CI>aeto7nys
Orbit large; frontal w-ithout postorbital process; jugal not specially
thickened. Cheekteeth with wide re-entrant folds.
Subfamily Eretiiizontin.\e
Erethizon, Eclivio[>ioctii, Coendou
The mandible in this family is characterized by the length of the symphysis;
the angular process is not so conspicuously distorted as in the Echimyidae, and
a weak ridge below the condyle similar to that sometimes found in Chinchillidae,
and presumably for the attachment of masseter medialis, foreshadowing that
which is so much lengthened and such an important feature of the jaw in
Caviidae, can be present. In Erethizon the lower border of the angular process
is conspicuously broadened.
Subfamily CHAETOMYINAE
Geographical Distribution. — ? Brazil.
Number of Genera. — One.
Characters. — As indicated in the above key.
CHAETOMYS 175
Genus i. CHAETOMYS, Gray
1843. CHAETOMYS, Gray, List Specimens Mamm. in Coll. Brit. Mus. p. 123.
Type Species. — Hystrix subspinosa, Kuhl.
Range. — Brazil; exact locality apparently not known.
Number of Forms.— One.
Characters.- — Frontals extremely broad, but some narrowing present in
front of the well-marked postorbital process. Parietals
strongly ridged, but the posterior part of the skull broad, and parietal ridges
showing no signs of coming together. Palate relatively narrow; short. Palatal
foramina very short, far in front of toothrows.
Bullae relatively large, the meatus produced sharply sideways, forming sharp
angle. Paroccipital processes short. Nasal chamber appears less open than is
usual in Erethizontinae. Jugal with anterior part immensely broadened, nearly
in contact with the postorbital process; the jugal nearly extending to the lachry-
mal. No canal for nerve transmission in infraorbital foramen. Mandible with
low coronoid; angular process relatively small, the lower border not specially
widened, but this part of the jaw clearly distorted outwards.
Fig. 42. CtMETOMVs subspinosus, Kuhl.
B.M. No. 3.9.4-86, S; X 1.
Fig. 43. Chaetomys subspinosus, Kuhl.
B.M. No. 3.9-4-86. ?; ;' >■
Fig. 44. Ch.-\f,to\iy.s sub.spinosis, Kuhl.
Cheekteeth: B. M- No. 3. 9.4.S(,, V; 3.
CHAETOMYS— ERETHIZONTINAE 177
Upper cheekteeth are divided into three lobes, the middle one being simple,
straight, separated from the front and the hind ones, which are each subdivided
by a well-marked outer re-entrant fold. The lower cheekteeth are not unlike
those of Kchimvs; there are one outer and two inner folds present. Incisors
narrow.
The spiny covering of the back is rudimentary, consisting of long wavy
bristles only; the head is densely covered with sharp spines, which extend over
the neck and forelimb. Feet as specialized as in any member of the family;
four digits present on fore and hindfeet, these bearing long curved claws, the
poUcx and hallux minute, replaced by a broad pad. Tail relatively long, scaly,
and moderately haired, the underside, near the body, clothed with stiff bristles,
as in Coendou.
Forms seen : suhspinosiis.
List of Named Forms
(The references and type localities of all Erethizontidae are the work of
Mr. G. W. C. Hoh.)
I. CHAETOMYS SUBSPINOSUS, Kuhl.
1820. Beitr. Zool. Manim. p. 71.
Brazil (?).
Synonym: tortilis, Olfers, 1820, Neue Bibl. Reis. XV, p. 211. Brazil.
moricandi, Pictet, 1843, Rev. Zool. p. 227. Brazil.
Subfamily ERETHIZONTINAE
Geographical Distribution. — As in the subfamily.
Number of Genera. — Three.
Characters. — Differing from the Chaetomyinae in the large orbit, the lack
of postorbital process, the lack of extreme thickening of the
jugal, the pattern of the cheekteeth, which are with wide inner and outer re-
entrant folds (three outer, one inner in the upper series), and the greater develop-
ment of spiny covering of the back. The feet may be highly specialized, or in
Eretliison less so. The tail may be short {Erethizon, Echinoprocta), or long and
prehensile [Coendou).
Remarks. — Pocock in 1922 proposed to divide this group into two sub-
families, Erethizontinae and Coendinae; he does not include in
his key the genus Ecliiiiopructa which is precisely intermediate in the main
character (the tail), between Pocock's two "subfamilies."
Key to the Genera of Erethizontinae
Hallux well developed, and no well-marked pad taking its place on the
hindfoot; inner side of forefoot not or less expanded; tail short,
non-prehensile. Erethizon
llalhiv vestigial or absent, its function taken over by a broad movable
pad; inner side of forefoot more expanded.
12 — Living Rodents — 1
178 ERETHIZONTINAE: ERETHIZON
Tail short, non-prehensile, httle longer than hindfoot. Echinoprocta
Tail long, prehensile (as far as known), much longer than hindfoot.
CoENDOi;
Genus i. ERETHIZON, Cuvier
1S22. ERETHIZON, Cuvier, Mem. Mus. Hist. Nat. IX, p. 425.
Type Species. — Hvstrix duisatu, Linnaeus.
Range. — North America; "Most of forested North America north of 40°
and south in the Rocky Mountains almost to Mexican boundary"
(Anthony). Forms named from Labrador, Nebraska, California, Arizona,
British Columbia, Alaska.
Number of Forms. — Seven.
Characters. — Nasals wide, widely open anteriorly; frontals broad, strongly
ridged, these ridges extending backwards to form a sharp
sagittal crest. Palate narrow anteriorly, very broad behind, and short. Zygoma
simple, jugal broader anteriorly. Bullae large, the external meatus produced
slightly sideways. Palatal foramina medium in size. Incisors relatively thin.
Mandible with low coronoid, relatively low condylar process, this thickened;
the area beside the condyle with noticeable ridge presumably for attachment of
masseter medialis; this short, not so pronounced as in some Chinchillidae.
Angular portion distorted outwards fairly strongly, the lower border abnormally
thickened.
LIpper cheekteeth with one external and one internal main persistent folds,
the other two outer folds (anterior and posterior) tending to isolate, and to take
up most of the lobes formed by the central folds. L'sually a trace of a small
posterior fold in the hack of each tooth.
The lower teeth reverse the pattern of the upper series.
The infraorbital foramen has no separate canal for transmission of nerve, in
this respect agreeing with all other members of the subfamily.
Entire body, limbs, head, tail, and sides of feet covered with thick hair which
completely conceals the highly effective spiny covering below it. The spines
are short, with barbed tip, and detach very easily; once sticking in an object
they are sometimes quite difficult to take out (this feature common to all
Erethizontinae).
Tail short and bushy, covered with spines more or less throughout. Hind-
foot lacking the inner pad characteristic of Coendoii, and with a well-developed
hallux, which is, however, shorter than the remaining four digits; claws curved,
powerful. Forefoot broad, with four functional digits. Mammae 4 (Anthony).
Size relatively large; up to about 34 inches head and body.
The genus is noteworthy as being the only Hystricoid adapted ior life in
cold climates.
Two closely allied species are admitted.
Forms seen: dorsatum, epixtwthum, iiivops.
Fig. 45. Erethizon epixanthum myops, Merriam.
B.M. No. 4. II. 30.1; X I.
Fig. 46. Erethizon epix.\nthvm mvops, Merriam.
B.M. No. 4.1 1.30. 1 ■ ■; I.
Fig. 47. Erethizon epixanthum myops, Merriam.
B.M. No. 4. II. 30.1 ; I.
Fig. 48. Erethizon epixanthum mvops, Merriam.
(.'heckteeth: 4. 11.30. 1 ; zk.
ERETHIZON— ECHINOPROCTA i8i
List of Named Forms
1. ERETHIZON DORSATUM DORSATUM, Linnaeus
1758. Syst. Nat. i, p. 57.
Eastern Canada.
Synonjin: hudsonis, Brisson, 1756, Regn. Anim. Quadr. p. 128. America.
Trouessart quotes as synonym : pilosus americanus, Catesby,
1 73 1, Nat. Hist. Carolina, i, xxx.
2. ERETHIZON DORSATUM PICINUM, Bangs
1900. Proc. New Engl. Zool. Club, II, p. 37.
L'Anse au Loup, Strait of Belle Isle, Labrador.
3. ERETHIZON EPIXANTHUM EPIXANTHUM, Brandt
1835. Mem. Acad. St. Petersb. pi. i, p. 390.
California.
4. ERETHIZON EPIX.\NTHUM BRUNERI, Swenk
1916. Univ. Studies Lincoln Nebr. vol. XVI, p. 3.
3 miles east of Mitchell, Scotsbluff County, Nebraska.
5. ERETHIZON EPIXANTHUM COUESI, Mearns
1897. Proc. U.S. Nat. Mus. XIX, p. 723.
Fort Whipple, Yavapai County, Arizona.
6. ERETHIZON EPIX.\NTHUM NIGRESCENS, .-Mien
1903. Bull. Amer. Mus. Nat. Hist. XIX, p. 558.
Shesley River, British Columbia, Canada.
7. ERETHIZON EPIXANTHUM MYOPS, Merriam
1900. Proc. Washington Acad. Sci. II, p. 27.
Portage Bay, Alaska Peninsula, Alaska.
Genus 2. ECHINOPROCTA. Gray
1865. ECHINOPROCTA, Gray, Proc. Zool. Soc. London, p. 321.
Type Species. — Erethizon rufescens. Gray.
Range. — Colombia.
Number of Forms. — One.
Characters. — Cranially and dentally not essentially different from small
species of Coendou (next to be described); a sagittal crest
evidently formed; f rentals not specially inflated; zygoma simple; a little con-
striction noticeable in interorbital region.
Size smaller than is normal in Coendou; the spines of the back long and
bristly, gradually becoming thicker and stronger as they approach the rump,
on which they are as strong as in Coendou. Head covered with sharp spines.
Feet, including the bone formation of the specialized pad of the hindfoot
(as figured by Trouessart), similar to Coendou; hallux suppressed. Tail short,
little longer than hindfoot, hairy, non-prehensile.
Forms seen : rufescens.
List of Named Forms
1. ECHINOPROCTA RUFESCENS, Gray
1865. Proc. Zool. Soc. London, p. 322.
Colombia.
1 82 COENDOU
Genus 3. COENDOU, Lacepede
1 799. CoENDOU, Lacepede, Tabl. des Divisions des Mamm. p. 11.
1S25. Sphiggurus, Cuvier, Dents. Mamm., p. 256. (Sphiggure, 1S22, Mem. Mus.
Nat. Hist. Paris, IX, p. 427.) (Sphiggurus spinosus, Cuvier.) V.'vlid .\s a SrB(;ENUS.
1825. SlNOETHEBis, Cuvier, Dents Mamm. p. 256. (Hystrix preheiisilis, Linnaeus.)
1835. Cercol.\bes, Brandt. Mem. Acad. St. Petersburg, C>, iii, p. 301. New name for
Coendou, Lacepede.
Type Species. — Hystrix preheiisilis, Linnaeus.
Range. — Mexico (through Central America.'), to Panama; Venezuela,
Colombia, Ecuador, Peru, Bolivia, Brazil south to Parana and Rio
Grande do Sul. One form named from Chile, and one "said to be from the
West Indies"; there seems reason to doubt both these localities.
Number of Forms. — Twenty-nine.
CHAR.'iCTERS. — Skull broad, sometimes characterized by somewhat extreme
inflation of frontals (this most developed in preJiensilis group,
also to a certain degree in bicolor and mexicanum); the skull in these species
sloping sharply downwards in front, and more gradually so behind; in smaller
species as paragayensis, the portion of the skull over the posterior zygomatic
root is the highest part ; between these extremes exist intermediate forms. Nasals
well open anteriorly, usually short. Parietals ridged, and a sagittal crest may
be formed (this evidently a variable character). Palate wide, especially pos-
teriorly; hamulars thick, usually joining the bullae, which are prominent.
Palatal foramina usually relatively short. Paroccipital processes not lengthened.
Jugal rather long; zygoma simple. Mandible like Eretliizon except that the
lower border is usually less extremely broadened; there is a tendency in this
group for the degree of distortion outwards of the angular process to be weak.
Cheekteeth essentially as Eretliizon.
Externally the body is covered in short thick spines, which probably do not
much exceed four inches in length at highest development. Tail prehensile,
so far as known; its length variable in the different species; sometimes slightly
longer than the head and body, but usually rather shorter. The lower part
at the end is naked, curling upwards when grasping an object. The underside
near the body is covered with stiff sharp bristles, which it has been suggested
perform a similar function to the caudal scales of the Anomaluridae, to assist
the animal's balance when resting on a branch.
The upper part of the tail near the body is spiny. The feet are very highly
specialized, the pad on the hindfoot at its highest development; the claws are
long and curved; both fore and hindfeet with four functional digits only; there
is no very marked discrepancy in their lengths. The pads of the hindtoot are
supported by a bony structure, which is well described and figured bv Water-
house, 1848, Nat. Hist. Mamm., p. 405, and pi. 18, tig. 4.
Some forms have the spiny covering of the back mixed with or covered by
long thick fur; the hair of the chest and belly is usually in these forms less
bristly, or soft. For these the subgeneric name Sphiggurus is used by Tate;
it is here retained.
Fig. 49. CoENDOu prehensilis boliviensis. Gray.
B.M. No. 50.6.5.2; X I.
Fig. 50. CoENDOu prehensilis boliviensis. Gray.
B.M. No. so.6.5.2; X I.
Fig. 51. CoENDOU prehensilis boliviensis, Gray.
B.M. No. 50.6.5.2; , I.
Fic. 52. CoENDOU PREHENSILIS BOLIVIENSIS, Gray.
Cheekteeth: B.M. No. 50.6.5.2; x 3*.
COENDOU i8s
Forms seen: bicolur, holiviensis, centralis, "couiy" [—paragayemis), insidiosus,
laetiatufn, melanurus, mexicanum, pallidus, prehensilis, pruinosus, quichua, roberii,
rothschildi, simonsi, tricolor, vestitiis, villosiis, yucataniae.
The genus is in need of revision; it appears to be in a more chaotic state
even than is usual among these Neotropical Rodents, many "species" being
apparently based on only one skin, with exact locality unknown.
It appears to me to divide, broadly speaking, into four or possibly five groups.
Subgenus Coendou (the spines not mixed with hairy covering) :
prehensilis group : large animals (largest of genus), with frontals normally at
maximum inflation for the genus; general effect silvery as regards colour,
the spines white terminally (evidently main spines black terminally in
all others). Long-tailed types.
With prehensilis, holiviensis, which is probably a synonym, or at most
a race of prehensilis, centralis, described as near brandti; brandti, which
seems very near prehensilis; and tricolor, the status of which is doubtful,
the type skull (broken) appears to be less arched in the region of the
frontals than in allies, and the colour of the one skin seen, on the identi-
fication of which there is some doubt, rather different. Perhaps this
species should be placed incertae sedis. C. sanctaemartae, not seen,
is described as a member of the group.
bicolor group : presenting typically the following features : spines black
terminally, general effect of animal dark; head and shoulders covered by
a profuse mantle of moderately long thick bristle-like spines (not sharp,
nor effective as weapons of defence). Relatively large; frontals markedly
inflated, but less extremely than in prehensilis group. C. simonsi is
evidently not more than a subspecies of bicolor. C. quichua, a smaller
form, with less developed mantle on head and shoulders; the mantle-
spines white-tipped in the type skin. Skull not arched in frontal region.
C. rothschildi, near quichua, differing in colour.
Incertae sedis species: platycentrotus, near prehensilis according to
Waterhouse, but placed in neighbourhood of bicolor group by Tate;
and nycthemera, stated by Waterhouse to be synonymous with bicolor,
but this identification questioned by Thomas; and listed by Tate as a
member of subgenus Sphiggurus.
Subgenus Sphiggurus (the spines mixed with and typically covered by
long woolly hair).
mexicanum group. Larger, very dark types, from Central America. Typi-
cally the skull considerablv inflated in the frontal region (about as in
bicolor). Includes laenatuin, in which the frontals are Hat, with no trace
of inflation. Goldman (Mammals Panama, Smiths. Misc. Coll. 69, 5,
p. 133, 1920) states that intergradation may take place here, and refers
laenatum to mexicanum as a subspecies; this indicates that too much
attention should not be paid to cranial characters in this group, as the
skulls of laenatum and the mexicanum (with yucataniae) examined are
verv distinct from each other.
i86 COENDOU
paragayensis group: normally smaller lighter types. (It is not easy to give
exact measurements of these species, as comparatively few of the skins
examined bear measurements.) So far as seen the frontals never inflated.
paragayensis (or the skins bearing the name " coniy" which accordini,'
to I'ate must be regarded as a synonym of paragayensis), if identified
rightly, are remarkable for the fact that the spines of the head and
shoulders are exceptionally strong, and not covered by any hair, which is
present, however, on the lower part of the back. The type skin of
roberti is similar, but even less hairy on head and shoulders.
insidiosiis (or skins bearing this name) have the head and back normally
hairy; the skulls of these three last-mentioned species are very similar.
Tate suggests that spinosus of Cuvier is probably a synonym of paragay-
ensis. C. villosiis is probably a synonym of insidiosiis according to Water-
house; it appears to be treated as such at the British Museum. C.
nigricans, not seen, is considered near villosus bv Waterhouse. C. mela-
niirus is a type much like the above-mentioned, but with a jet-black tail;
the skull is flat. C. pallidiis, based on a young animal "said to be from
the West Indies," is a similar type of animal, but much lighter coloured
(albinistic?); rather short-tailed.
A rather distinct section is seen in vestitus and pruiiiosus, which differ
from each other in colour; both have no inflation of the frontals; the
spines are of two kinds, the normal mixed with longer "bristle-spines";
covered as usual in the subgenus with thick hair. In vestitus the tail
appears shorter than in any other; it is still, however, considerably longer
than Echinoprocta, and partly naked, as in Coendou.
Finally eliilensis, ajfinis, and sericeus are not represented in London;
the first-named was said to come from Chile, but Tate, p. 299, states:
"It seems improbable that any Porcupine exists in the wild state in
Chile."
List of Named Forms
Subgenus Coendou, Lacepede
prehensilis Group
1. COENDOl' PREHENSILIS PREHENSILIS, Linnaeus
1758. Syst. Xat. 10th Ed. p. 57.
Brazil. (Probably near Purnainbuco.)
Synoin-m; ciiiiiu/ii, Desmarest. 1S22, Ency. Mc'th. (Mamni.). 2, p. 346.
Hrazil.
longicaudatus, Lacepede, 1709. Tahl. des Div. dcs Mamm.
p. I. Cayenne.
2. COENDOU PREHENSILIS BOLIVIENSIS. Gray
1850. .Ann. Nat. Hist. V. p. 3S0,
Bolivia.
3. COENDOU CENTRALIS, Thomas
1903. Proc. ZooL Soc. London, p. 240.
Chapada, Matto Grosso, Brazil.
COENDOU 187
4. COENDOU BRANDTII, Jentink
1879. Notes Leydcn Mus. i, p. 96.
Matto Grosso, Brazil (?).
5. COENDOU SANCTAI•:^L■\RTAE, Allen
1904. Bull. Arner. Mus. Nat. Hist. XX, p. 441.
Bonda, Santa Marta district, Colombia.
6. COENDOU TRICOLOR, Gray
1850. Ann. Nat. Hist. V, p. 381.
Bolivia (?).
bicolor Group
7. COENDOU BICOLOR BICOLOR, Tschudi
1845. Fauna Peruana, p. 186.
Woods between Rivers Tullamayo and Chanchamayo, Peru.
8. COENDOU BICOLOR SIMONSI, Thomas
1902. Ann. Mag. Nat. Hist. 7, IX, p. 141.
Charuplaya, Secure River, Yungas, Bolivia,
y. COENDOU QUICHUA QUICHUA, Thomas
1899. Ann. Mag. Nat. Hist. 7, IV, p. 283.
Puembo, Pichincha, Ecuador.
10. COENDOU QUICHUA RICH-^RDSONI, Allen
1913. Bull. Amer. Mus. Nat. Hist. XXXII, p. 478.
Esmeraldas, Ecuador.
11. COENDOU ROTHSCHILDI, Thomas
ig02. .Ann. Mag. Nat. Hist. 7, X, p. 169.
Sevilla Island, off Chiriqui, Panama.
incertae sedis
12. COENDOU PL.ATYCENTROTUS, Brandt
1835. Mem. Acad. St. Petersb. p. 399.
"Ainerica australis."
13. COENDOU NYCTHEMERA, Kuhl
1820. Beitr. Zool. Mamm. p. 71.
No locality. Brazil (?).
Subgenus Sphiggurus, Cuvier
mexicanum Group
14. COENDOU MEXICANUM MEXICANUM, Kerr
1792. Anim. Kingd. p. 214.
Mountains of Mexico.
Synonym: novaehispaniae, Brisson, 1756, Reg. -Anim. p. 127. Me.iico.
(For status of Brisson's specific names see Tate, Bull.
Amer. Mus. N.H. LXVIII, 1935, p. 297.)
liebmanni, Reinhart. 1844, Arch. Naturg. p. 241. Me.vico.
15. COENDOU MEXICANUM YUC.\TANL\E, Thomas
1902. .Ann. Mag. Nat. Hist. 7, X, p. 249.
Yucatan, Mexico (probably near Izamal).
16. COENDOU L.\EN".VrUM. Thomas
1903. Ann. Mag. Nat. Hist. 7, XI, p. 381.
Boquete, Chiriqui, Panama.
i88 COENDOU
paragayensis Group
(Typical Section)
17. COENDOU PARAGAYENSIS, Oken
1S16. Lchrbuch der ZooIokIc. p. S70.
Paraguay.
Synon>'in: couiy, Desmarest, 1S22, Mammalogic, ii, p. 345. Brazil.
18. COENDOU SI'INOSUS, Cuvier
1822. Mem. Mus. Hist. Nat., IX, p. 433.
No locality (? Brazil) (? Based on paragayensis, Oken).
19. COENDOU ROBERTI, Thomas
1902. .-^nn. Mag. Nat. Hist. 7, IX, p. 63.
Rofa Nova, Parana, Brazil.
20. COENDOU INSIDIOSUS, Kuhl
1S20. Beitr. Zool. Mamm. p. 71.
No locality (? Brazil).
21. COENDOU VILLOSUS Cuviir
1822. M^m. Mus. Hist. Nat. IX, p. 434.
Brazil.
22. COENDOU NIGRICANS, Brandt
1835. Mem. .Acad. St. Pctersb. p. 403.
Brazil.
23. COENDOU .MELANURUS, Wagner
1S42. .Archiv. fur Naturg. i, p. 360.
Barra, Rio Negro, Brazil.
24. COENDOU PALLIDUM, Waterhouse
1848. Nat. Hist. Mamm. ii, p. 434.
"Said to be the West Indies."
(restittis section)
25. COENDOU VESTITUS, Thomas
1899. .Ann. Mag. Nat. Hist. 7, IV, p. 284.
Colombia.
26. COENDOU PRUINOSUS, Thomas
1905. Ann. Mag. Nat. Hist. 7, XVI, p. 310.
Montafias de la Pedregosa, Merida, Venezuela.
Not allocated to group ; not seen
27. COENDOU AFFINIS, Brandt
1835. Mem. Acad. St. Petersb. p. 412.
Brazil.
28. COENDOU SERICEUS, Cope
1889. Amer. Naturalist, XXIII, p. 136.
Sao Joao do Monte Negro, Rio Grande do Sul, Brazil.
29. COENDOU CHILENSIS, Molina
1S09. Geogr. Nat. and Civil Hist, of Chile, p. 242.
Chile.
Numbers 27, 28, 29 have not been seen; they are listed by Tate as members
of the subgenus Sphiggunis.
DASYPROCTIDAE 189
ERETHIZONTIDAE:
SPECIAL WORKS OF REFERENCE
VV'ATEBHOiSE, 1 848, Natural History Mammalia, Rodentia.
Tatf, 1935, Taxonomy of Neotropical Hvstricoid Rodents, Bull. Amer. Mus. Nat. Hist.
LXVIH, p. 295. '
I'ocoiK, Proc. Zool. Soc. London, 1922, p. 365. External Characters of some Hystrico-
morph Rodents (Coendou, Erethizon).
'i"ROl'ESS.\RT, Echiiwprocla, Bull. Mus. Hist. Nat. 1920, no. 6, p. 44S.
Al-LEN, North American Rodentia, p. 385, 1876. " Hystricidae " (Erethizon).
TuLLBERG, Nova Acta Reg. Soc. Sci. Upsaliensis, XVHI, 3, i, 1899.
The family is known fossil from the Oligocene from America.
Family DASYPROCTIDAE
1896. Thomas: Hystricomorph.\ : Family Dasyproctidae, part, included Cuniculus
(^" Coelogenys").
1899. Tullberg: Hystricomorpha : Family Caviidae, part.
1918. Miller & Gidley: Hystricoid.\e : Family Dasyproctidae.
1924. Winge; Family Hystricidae, part, Dasyproctini, part, Dasyproctae, part (included
Cufiiculus).
1928. Weber: Hystricoidea : Family Caviidae, part, subfamily Dasyproctinae, part
(included Citniculus).
Geogr.\phical Distribution. — Tropical America, from Mexico through
Central America to Bolivia and Paraguay,
Ecuador and Peru. Trinidad. Lesser Antilles.
Number of Genera. — Two.
Characters. — E.xternal form much modified for cursorial life; hindlimbs
lengthened; hindfeet with three digits, forefoot with four
functional digits, the claws hooflike. Cheekteeth semi-rooted, extremely
hypsodont, closely paralleling the structure present in the Hystricidae. Clavicles
undeveloped.
Remarks. — This family has (together with Cuniculus) often been united with
the Caviidae. This association appears most unnatural. The
lower jaw is totally distinct in the two groups, Dasyprocta being typically
Hystricoid in this formation; the cheekteeth show an entirely different pattern
in the two groups.
The similarities between such genera as Dasyprocta and Dolichotis in the
arrangement of digits and parts of the skeleton for swift running appear to be
parallel evolution brought about by a similar mode of life, comparable to the
similarities between such types as Bathyergus and, say, Geomys, which resemble
each other externally to a large degree and yet in which the zygomasseteric
structure and the cheekteeth are totally different. One of the reasons which has
been advocated for classing Dasyprocta with the Caviidae is the formation of the
penis, which is said to be armed with a pair of horny spikes in these genera;
it is therefore interesting to note that according to Pocock the penis of Dolichotis
and ot Hydrochoerus, both members of Caviidae, lack these spikes, disagreeing
in this character from Cazia and other members of the Caviidae, as well as from
iQO DASVPROCTIDAE: DASYPROCTA
Dasypructti and Cuiiiciiliis. But in any case the penis does not furnish a suffi-
ciently rehable character on which to base family distinctions.
Key to the Genera of Dasyproctidae
Tail not obsolete, approaching half length of hindleg; toothrow reduced,
"teeth smaller both relatively and absolutely than in any species
oi Dasyprocta" (Thomas). ^IY0PR0CTA
Tail obsolete; teeth relatively larger, and toothrow less reduced. Dasyprocta
These two genera are not well marked, and might be regarded as subgenera
of one genus. Mxoprocta contains much smaller forms than is normal in
Dasyprocta.
Ccnus I. DASYPROCTA, Illiger
iSii. Dasyproct.a, Illiger, Prodr. Syst. Mamm. et Avium, p. 03.
Type Species. — Mus aguti, Linnaeus.
Range. — As in the familv Dasyproctidae; south to South Brazil.
Number of Forms. — About forty-six are named.
Char.\CTERS. — The skull is less ridged than in other Hystricoids of a similar
size; the nasals shorter than the frontals, which are broad,
flat, and with a well-marked though short postorbital process at suture of frontals
and parietals. A weak short sagittal crest is developed in the adult. The bullae
are moderately large ; the paroccipital processes prominent, though not so length-
ened as in CiDiiciiliis. The palate is straight, and extends back to level of M.3;
the hinder part being formed much as in the Hystricidae. Palatal foramina
short, far in front of toothrow. Lachr\'mal much enlarged, forming most of
upper zygomatic root, and part of the lachr\-mal canal is open on the side of the
rostrum, immediately in front of the anterior part of toothrow. There is no
canal for nerve transmission in the infraorbital foramen, which is of medium
size. Zygoma generally simple. Mandible with angular portion powerfully
distorted outwards, and its lower border slightly drawn backwards; coronoid
process low; condylar process rather broad.
Cheekteeth strongly hypsodont; like those of the Hvstricidae in essential
pattern; one more or less persistent narrow inner fold in the upper series; the
outer folds soon isolate as islands, and there is a tendency tor the islands to divide
on the surface of the tooth, so that there may be seven or eight or more minute
islands in a worn tooth. Lower cheekteeth like the upper series, but with the
pattern reversed. Incisors relatively thin, compressed.
Externally the form is slender, cursorial, the hindlimbs lengthened, the
hindfoot very long and narrow, with three digits which bear sharp hoot-like
claws; the central digit is the longest, D.4 is a little shorter than D.2; the sole
is naked; in the skeleton of the foot the metatarsal bones for the outer digits
are absent or vestigial. The forefoot is less elongated than the hindtoot; the
digits are four, but the appearance of the foot is perissodactyie owing to D.5
being considerably reduced; the pollex is represented by a knob. The fur on
DASYPROCTA
191
the hinder part of the body is very long and thick. The ears are of medium size.
The tail is obsolete. The head and body measurement may approach 580 mm.
Forms seen : aguti, azarae, boliviae, catrinae, cayennae, coibae, cristata,
croconota, flavescens, fuliginosa, isthmica, lucifer, liinaris, maraxica, nigra, para-
guayensis, pandora, prymnolopha, punctata, ruatanica, rubrata, variegata,
yungarum.
FlC. 53. D.\SYPROCTA PUNCTATA ISTHMICA, Alston.
B.M. No. 98. 1 1.6. 10; X I.
List of Named Forms
(References and type localities for all forms of Dasyproctidae are the work of
Mr. G. W. C. Holt.)
Tate divides the genus into three sections: " Eastern or red-rumped Agoutis,"
"Central American Agoutis," and "Dark-grey Agoutis." The material ex-
amined does not support these divisions; moreover, I have been quite unable to
Fig. 54. Dasyprocta punctata isthmica, .Alston.
B.M. No. 98.11. 6.10; y I.
Fig. 55. D.^YPROCTA punctata isthmica, Alston.
ChecktL-eth: B.M. No. 98.11.6.10; • 4.
(In the lower jaw the anterior tooth is the much worn milk molar — seen in profile in fig. 5 4 ;
in the upper jaw the anterior tooth is the newly cut premolar, and the larce inner root of the
shed milk molar is seen to the right or inner side of this tooth.)
I
DASYPROCTA 193
get this large and unwieldy genus into any definite order, and therefore list
geographically.
The real dark-grey types like colombiana or fuliginos a grade quickly into dark
unicolorous types, which in turn grade into reddish-rumped types, which
seem to grade into the red types like aguti. Sometimes a blackish middorsal
area may be present, as mprymnolopha.
How many species should be recognized I am not prepared to say, but it
seems clear that far too many are at present standing, and many could be reduced
to the rank of subspecies. It is to be hoped that someone will attempt a revision
of this genus, which is much needed.
1. DASYPROCTA NOBLEI, M\en
1914. Proc. New. Engl. Zool. Club. V. p. 6g.
Goyave, Guadeloupe, Lesser Antilles.
2. DASYPROCTA ALBIDA, Gray
1842. Ann. Mag. Nat. Hist, i, X, p. 264.
St. Vincent, Lesser Antilles.
3. DASYPROCTA ANTILLENSIS, Sdater
1874. Proc. Zool. See. London, p. 666.
St. Lucia, Lesser Antilles.
4. DASYPROCTA RUBR.ATA RUBRATA, Thomas
1898. Ann. Mag. Nat. Hist. 7, H, p. 272.
Savannah Grande, Trinidad.
5. DASYPROCTA RUBR.^TA FL.WESCENS, Thomas
1898. Ann. Mag. Nat. Hist. 7, H. p. 273.
Caripe, Cumana, Venezuela.
6. DASYPROCTA LUCIFER LUCIFER, Thomas
1903. .-^nn. Mag. Nat. Hist. 7, XI, p. 491.
Caicara, Rio Orinoco, Venezuela.
7. DASYPROCTA LUCIFER C.^YEN.NAE, Thomas
1903. .\nn. Mag. Nat. Hist. 7, XI, p. 492.
Approvague, Cayenne.
8. DASYPROCTA CAY'ANUS, Lacep^de
1802. Tabl. des Div. des Mamm. p. 78.
Cayenne.
9. DASYPROCTA PRYMNOLOPHA, Wagler
1831. Isis, XXIV, p. 619.
Guiana.
10. DASYPROCTA NTGRICLUNIS, Osgood
1915. Field. Mus. Nat. Hist. Publ, Zool. ser. X^ p. 192.
Sao Marcello, upper Rio Preto, Bahia, Brazil.
11. DASYPROCTA CROCONOTA, Wagler
1831. Isis, XXIV, p. 618.
Amazon River, Brazil (mouth of Rio Madeira).
12. DASYPROCTA AGUTI AGUTI, Linnaeus
1766. Syst. Nat. 12th. ed. p. 80.
Brazil.
13 — Living Rodents — I
194 DASYl'ROCTA
(D. aguti aguti) Synonym: (?) leporina. Linnaeus, 1758, Syst. Nat. 10th. ed. p. 59.
Unknown (probably unidentifiable, according to Tate.)
bicolor, Boddaert, 1785, Elenchus Anun. p. 103.
13. DASYPROCTA AGUTI MARAXICA, Thomas
1923. Ann. Mag. Nat. Hist. 9, XII, p. 341.
Marajo Island, Amazon River, Brazil.
14. DASYPROCTA AGUTI LUNARIS, Thomas
1917. Ann. Mag. Nat. Hist. 8, XX, p. 259.
Moon Mountains, British Guiana.
15. DASYPROCTA AZARAE AZARAE, Lichtenstein
1823. Doubl. Zool. Mus. Berlin, p. 3.
Sao Paulo, Brazil.
16. DASYPROCTA AZARAE C.ATRINAF.. Thomas
1917. Ann. Mag. Nat. Hist. 8, XX, p. 311.
Santa Catharina, Southern Brazil.
17. DASYPROCTA AUREA, Cope
1889. Amer. Naturalist, p. 13S.
Chapada, Matto Grosso, Brazil.
18. DASYPROCTA CAUDATA, Lund
1841. Afh. K. Danske. Vid. Selsk. 4, viii, p. 286.
Rio das Velhas, Minas Geraes, Brazil.
ig. DASYPROCTA PARAGUAYENSIS, Liais
1872. Climats, Geologic. Faune et Geographic Botanique du Bresil, p. 536.
Paraguay.
Synonym: felicia, Thomas, 1917, Ann. Mag. Nat. Hist. 8, XX, p. 310.
Near Concepcion, Paraguay.
20. DASYPROCTA MEXICANA, Saussure
i860. Rev. Mag. Zool. 2, XII, p. 53.
"Hot zone of Mexico," probably in State of Vera Cruz.
21. DASYPROCTA PUNCTATA PUNCTATA, Gray
1842. Ann. Mag. Nat. Hist, i, X, p. 264.
Realejo, west coast of Nicaragua.
22. DASYPROCTA PUNCTATA RICHMONDI, Goldman
191 7. Proc. Bio!. Soc. Washington XXX, p. 114.
Escondido River, 50 miles above Bluefields, Nicaragua.
23. DASYPROCTA PUNCTATA CHIAPENSIS, GoIdni;in
1913. Smiths. Misc. Coll. LX, no. 22, p. 13.
Huehuetan, Chiapas, Mexico.
24. DASYPROCTA PUNCTATA YUCATANICA, Goldman
1913. Smiths. Misc. Coll. LX, no. 22, p. 12.
Apazote, Campeche, Mexico.
25. DASYPROCTA PUNCTATA UNDERWOODI, Goldman
1931. Journ. Washington Acad. Sci. XXI, p. 481.
San Geronimo, district of Pirris, West Costa Rica.
2fa. DASYPROCTA PUNCTATA DARIENSIS, Goldman
1913. Smiths. Misc. Coll. LX, no. 22, p. 11.
Near head of Rio Limon, Mt. Pirri, Eastern Panama.
DASYPROCTA 195
27. DASYPROCTA PUNCTATA NL'CHALIS, Goldman
1917. Proc. Biol. Soc. Washington XXX, p. 113.
Divala, Chiriqui, Panama.
28. DASYPROCTA PUNCTATA ISTHMICA, Alston
1876. Proc. Zool. Soc. London, p. 347.
Colon, Panama.
29. DASYPROCTA RU.^TANICA, Thomas
1901. Ann. Mag. Nat. Hist. 7, VIII, p. 272.
Ruatan Island, Bay of Honduras.
30. DASYPROCTA CALLIDA, Bangs
1901. Amer. Naturalist, XXXV, p. 635.
San Miguel Island, Panama.
31. DASYPROCTA COIBAE, Thomas
1902. Nov. Zool. IX, p. 136.
Coiba Island, Panama.
32. DASYPROCTA PANDORA. Thomas
1917. Ann. Mag. Nat. Hist. 8, XX, p. 313.
Gorgona Island, off Colombia.
33. DASYPROCTA VARIEGATA VARIEG.\TA, Tschudi
1845. Fauna Peruana, p. 190.
Chanchamayo region. Eastern Peru.
34. DASYPROCTA VARIEGATA ZAMORAE, Allen
1915. Bull. Amer. Mus. Nat. Hist., XXXIV, p. 627.
Zamora, Eastern Eucador.
35- DASYPROCTA VARIEGATA CHOCOENSIS, .\llen
1915. Bull. Amer. Mus. Nat. Hist. XXXIV, p. 627.
Los Cisneros, Choco district, Colombia.
36. DASYPROCTA VARIEGATA COLOMBIANA, Bangs
1898. Proc. Biol. Soc. Washington XII, p. 163.
Santa Marta, Colombia.
37. DASYPROCTA VARIEG.A.TA VUNGARUM, Thomas
19:0. Ann. Mag. Nat. Hist. 8, VI, p. 505.
Chimosi, Vungas, Bolivia.
38. DASYPROCT.-\ VARIEG.VPA nOI.IVI.\E, Thomas
1917. Ann. Mag. Nat. Hist. 8, XX, p. 312.
Charuplaya, Bolivia.
39. DASYPROCTA VARIEG.'^TA URUCUMA, Allen
1915. Bull. Amer. Mus. Nat. Hist. XXXIV, p. 634.
Urucum, near Curumba, Matto Grosso, Brazil.
40. DASYPROCTA FULIGINOSA FULIGINOSA, Wagler
1832. Isis, XXV, p. 1220.
Near .Amazon River, Brazil. (Borba, Rio Madeira.)
Synonym: nigra, Gray, .^nn. Mag. Nat. Hist, i, X, p. 264, 1842.
nigricans, Wagner, 1842, .Archiv. fur Naturg. i, p. 362.
Borba, R. .Madeira, Brazil.
caroliniensis, Cuvier, Ger\ais, Mamm. i, 1854, p. 329.
196 DASYPROCTA— MVOPROCTA
^i. DASYPROCTA FULIGINOSA CANDELENSIS, Allen
IQ15. Bull. Amer. Mus. Nat. Hist. XXXIV, p. 625.
La Candcia, Huila, Colombia.
42. DASYPROCTA FULIGINOSA MESATIA, Cabrera
1917. Madrid Trab. Mus. Nac. Ci. Nat. 31, p. 53.
Tarapote, Ecuador.
43. DASYPROCTA CRISTATA, Desmarest
1816. Nouv. Diet. d'Hist. Nat. 2d. Ed. i, p. 213.
Surinam, Dutch Guiana.
44. DASYPROCTA KALINOWSKII, Thomas
1897. Ann. Mag. Nat. Hist. 6, XX, p. 219.
Idma, valley of Santa ."Vna, Cuzco, Peru.
Genus 2. MYOPROCTA, Thomas
1903. Myoprocta, Thomas, Ann. Mag. Nat. Hist. 7, XH, p. 464.
Type Species. — Cavia acoucliw Erxleben.
Range. — Guianas, Brazil, Colombia, Ecuador, Peru.
Number of Forms. — Ten.
Ch.\r.\cters. — Like Dasxprocta, but smaller (about 380 or less, head and
body) ; the tail less reduced, approaching about half the length
of the hindleg, slender, hairy. Essential cranial and dental characters as in
Dasyprocta, but toothrow reduced, the teeth "smaller both relatively and
absolutely than in any species of Dasyprocta" (Thomas); the sagittal crest
appears to be in old individuals rather longer than in Dasyprocta. There may
be a small backwardly directed process on anterior portion of jugal, just behind
its point of junction with zygomatic process of maxillary.
Forms seen: acoitcliv, cavmainiin, lepttira, liiiuiniis, milleri, pratti, puralis.
There appears some doubt on the status of exilis. Apart from this the forms
divide into two groups, the type species, reddish above, and the pratti group,
duller greenish types.
List of N.\med Forms
(acoucliv Section)
1. MYOPROCTA ACOLXHY, Erxleben
1777- Syst. Regn. Anim. p. 354.
Cayenne.
2. MYOPROCTA LICPTURA, Wapner
1844. Schreber Saug. .Suppl. IV, p. 49.
Rio Negro, Brazil.
(pratti Section)
3. MYOPROCTA PRATTI PRATTI, Pocock
1913. Ann. Mag. Nat. Hist. 8, XII, p. no.
Pongo de Rentema, Rio Marafion, Peru.
MYOPROCTA— HYSTRICIDAE 197
4. MYOPROCTA PRATTI ARCHIDONAE, Lonnberg
1925. Journ. Manim. Baltimore, VI, p. 274.
Archidona, Province Oriente, Ecuador.
5. MYOPROCTA PRATTI PLKALIS, Thomas
1926. Ann. Mag. Nat. Hist, g, XVII, p. 639.
.■\yapua, about 300 kilometres south-west of Manaos, Brazil.
6. MYOPROCTA PRATTI CAY.MAXLM, Thomas
1926. .^nn. Mag. Nat. Hist. 9, XVII, p. 638.
Canabouca, Parana de Jacare, 120 kilometres south-west of Manaos,
Brazil.
7. MYOPROCTA PR.-\TTI LIMANUS, Thomas
1920. .\nn. Mag. Nat. Hist. 9, VI, p. 279.
Acajutuba, Rio Negro, Brazil.
8. MYOPROCTA MILLERI, .'^llen
1913. Bull. .\mer. Mus. Nat. Hist. XXXII, p. 477.
La Murelia, Caqueta, Colombia.
Not allocated to section ; not seen
9. MYOPROCTA EXILIS EXILIS, Wagler
1831. Isis, XXIV, p. 621.
.\mazon, Brazil.
10. MYOPROCTA EXILl.S PARVA, Lonnberg
1921. Ark. Zool. XIV, no. 4, p. 41.
Rio Curaray, Prov. Oriente, Ecuador.
DASYPROCTIDAE:
SPECIAL WORKS OF REFERENCE
Waterhovse, 1848, Natural History' Mammalia, vol. II, Rodentia.
Tate, 1935, Bull. .\mer. Mus. Nat. Hist. LXVIII, p. 295. Ta.\onomy of Neotropical
Hystricoid Rodents.
PococK, Proc. Zool. Soc. London, p. 365, 1922. External Characters of some Hystrico-
niorph Rodents.
The Dasyproctidae are known fossil from the Miocene, from the Neotropical
region only.
Family HYSTRICIDAE
1896. Thomas: Hvstricomorpha : Family Hystricidae.
1899. Tullberg: Hvstricomorpha: Family Hystricidae.
1918. Miller & Gidley: Superfamily Hystricoidae: Family Hystricidae, with sub-
families Hystricinae and Atherurinae.
1924. Winge: Family Hvstricid.ae, part, Hystricini, part, Hystrices.
1928. Weber; Hvstricoidea: Family Hystricidae.
Geographical Distribution. — Tropical parts of the Old World ; the greater
part of the African Continent; Italy and
Sicily; Transcaucasia; Southern Palaearctic Asia (Arabia, Syria, Persia, Sleso-
potamia, .\fghanistan, Russian Turkestan); Peninsular India and Cevlon;
igS HYSTRICIDAE
South China (south of the Yangtsekiang), Hainan, Assam, Burma southwards to
Alalacca, Sumatra, Java and islands to the east of it (Sumbawa, Flores); Borneo;
represented in the Phihppine Islands.
Number of Genera. — Four.
Ch.\r.\cters. — E.xternal form heavy, terrestrial-fossorial; modification of
hair into spiny covering always well developed, at extreme
development reaching a grade of specialization not seen elsewhere in the Order;
tail always bearing a group of modified quills or bristles; digits of hindfoot five.
Cheekteeth moderatelv to e.xtremely hypsodont, semi-rooted in progressive
genera, the re-entrant folds isolating early on crown surface as narrow enamel
islands; bullae relatively small, and paroccipital processes not lengthened.
Occipital region of skull strongly ridged, prominent; zygoma simple; a tendency
present towards extreme inflation and lengthening of nasals; clavicles imperfect;
habits entirely terrestrial.
According to Tullberg, the carpus has no free centrale [Atherurus, Hystrix),
unique in the Order among those examined by him except in Cuniculidae.
According also to this author, the lungs are abnormal, being divided into a
number of small lobes.
Rem.\rks. — Lyon in 1907 proposed to divide the family into two subfamilies,
the Hystricinae and the Atherurinae, on account of the differ-
ences of the length and structure of the tail between the two groups, the fact
that there are only three sacral vertebrae present in Atherurinae as against four
in the Hystricinae (at the same time remarking that there is apparently some
variation in the number of the vertebrae, especially lumbar, sacral, and caudal).
But although the Atherurus group is much less specialized than the Hxstrix
group both in spinv covering, reduction of tail, and in the more brachyodont
cheekteeth, there appear to be too manv essential characters common to both
groups for this division to be maintained; the pattern of the cheekteeth and the
structure of the feet, for example, are essentiallv similar in both groups; and
the cranial characters of Thecurus, the lowest member of the Hvsirix group,
are very similar to those of Atherurus, including the rather important character
of length of nasals.
Cheekteeth. — The cheekteeth vary individually, but the essential pattern
throughout the family is, in the upper series, one inner and
three outer folds, the folds isolating as islands almost immediately on the flat
crown surface ; like the Dasvproctidae, there is a strong tendency for the isolated
folds to divide, particularly the posterior one, so that on the outer side of the
tooth there are usuallv at least four isolated islands in the adult. The lower
cheekteeth reverse the pattern of the upper series. The milk premolars are shed
comparatively late in life.
Gener.\l Extern.al Ch.ar.acters of principal species, as regards the
development of spiny covering.
For note on details of formation of hollow "rattling-quills" (caudal quills)
of fivstn'x group, see p. 208.
HYSTRICIDAE 199
Trichvs lipura is the most primitive species in development of spiny cover-
ing, in all respects. No true quills are developed; the body is covered
with relatively short flattened weakly developed spines; the tail is more
or less naked, and bearing a brush of unmodified bristles at the end; the
head is hairy.
Atherurus macrourus presents the next stage of development; the body is
covered with spines of a similar nature, but sharper, longer, and evidently
more effective as weapons of defence; the head is similar to Trichvs;
the tail long, less naked, covered with spinv short hairs, and the end bears
a cluster of much more specialized bristles, these being alternatively
expanded and contracted; but, as far as seen, no quills are as yet developed.
Atherurus africanus and related African forms present a higher stage of
development in that among the spines of the back there are present a
few thick circular quills, of the type found in all higher Porcupines;
these var\' in their development; thev mav be quite strong or, in some
skins I have seen, verv weak, so that perhaps if a really large series of
skins came to hand from Africa it might be that some "quill-less"
ones would be among them. Otherwise the external characters are as in
the Asiatic Atherurus.
It is interesting that, judging bv a specimen of Atherurus at the
London Zoological Gardens, verv much the same noise can be made by
the rattling of the tail bristles as that produced bv the smaller species of
Hystrix.
Thecurus pumilis, from the Philippines, to which T. sumatrae evidently
bears a close resemblance in general spine characters, appears to be the
lowest true Porcupine; in this species, as in all the Hystrix group, the
tail has become strongly reduced and its end bears a cluster of small and
very poorly developed hollow "rattling-quills" which reach such a high
stage of specialization in the higher Porcupines ; some of these tend to
be closed at the tip. A certain number of short thick true quills are
developed; the spines of the body e.xtend to the rump from the upper
part of the back ; the head is hairy, much as in the Brush-tailed Porcupines.
Hystrix {Acanthion) jaraniann represents the next stage; the caudal quills
are very weakly de\eIoped, essentially as in Thecurus pumilis; the head
remains haip*-; the quills of the back are perhaps slightly more developed
than in Thecurus pumilis; judging by the few skins seen they appear, as
in Thecurus, to be tightly wedged in the body, so that I should imagine
they are very infrequently shed, but I have not seen this animal in cap-
tivity; on dried skins they do not give to the touch as do the quills of most
higher Porcupines.
Thecurus crassispinis from Borneo stands rather alone in development of
spines, and presents a mixture of generalization mixed with extremely
high specialization. The quills of the back are enormously thick, rela-
tively as thick as those even of the most highlv developed forms of
Hystrix, or so it seems to me. But no long thin quills are developed to
cover them, as they are in Hystrix cristata and leucura groups; the head
!oo HYSTRICIDAE
remains hairy, with no trace of a crest; and the caudal quills remain at
their lowest development.
Hystrix (Acniitliioii) hodgsoiu represents a stage of development typically
not very much higher than in jiivaniciim; the quills, though not as well
developed as in Thecurus crassispiiiis, are profuse and well developed,
less tightly wedged in the body apparently than m javanicum; there is a
certain growth of long hair-like quills on the back, not met with in those
described heretofore; a vestigial crest may be present (or suggested) on
the head ; but the caudal quills remain very poorly developed. Whether
certain intergradation takes place between this and such forms as klossi
from the same area I do not know; there seems to be a rather distinct
difference as a rule between skins seen of Iiodgsoni, as compared with
klossi, as regards crest, caudal quills, etc. ; comparable to the difference
seen between jai-iiniciim and biachvunis.
Hystrix {Acanthiou) bracliyurus (with lungicauda, miilleri) reaches a rather
higher state of specialization; the caudal quills are as a rule larger than
in hodgsoni, more open, and apparently less primitive; the quills of the
body are powerful and profuse, though not attaining any great length.
Hvxtrix (Acanthiou) klossi is very similar in external characters to hrachyurus,
though on cranial characters belonging to a different group; there are a
few long liair-like quills present, as in hodgsoni; the crest tends to become
less abortive, and quite well marked; this tvpe, I believe, leads on to the
Chinese Porcupine sidKiistcitiis, in which the crest is said to be quite well
developed, but skins of which I have not seen unless a very small juvenile
labelled '' viiniuiiioisis," from Annam, which has for its size surprisingly
developed caudal quills and quite conspicuous crest, represents this
species, as from these characters I suspect it may do.
Hvstrix leuciira (with hirsutirostris) marks the highest development to be
attained in a Porcupine; a long crest of hairs is present on the head;
the quills are exceedingly profuse on the back; the short ones found in
the preceding species being more or less covered by an outgrowth
of long thin quills, each with several rings instead of only one as in
the above species. The caudal quills are large, well open, and at their
highest development ; there are many short white (ordinary) quills in
the neighbourhood of the tail. Sometimes the long quills tend to be
narrower than in cristata and other African Porcupines. The bodily
size is usually larger than in other .Asiatic Porcupines.
Hystrix cristata, H. galeata, H. africaeaiistralis are indistinguishable from
one another externally; the quills may tend to be thicker, perhaps longer
and more powerful than in leucura, otherwise the external covering is
essentially similar, including the long crest and powerful tail-quills; the
size, in galeata, becomes the largest in the genus.
For the last thirteen vears these animals have been a special hobby ot the
author, in the London Zoological Gardens, and a tew words on their captivity
habits may not be amiss.
The temperament of //. cristata compared with the smaller //. hrachyurus
HYSTRICIDAE 201
type of animal is as different as that of a dog from a cat. The smaller Porcupines
never display, so far as I have observed them, the slightest nervousness, and
generally seem to tame down and come to hand almost on arrival, and to be
the most friendly of animals, though occasionally exhibiting an unpleasant
streak in their character which I have only once observed in cristata (an old
specimen newly imported from aiiroad which might have been ill-treated).
On the other hand all cristata Porcupines I have ever seen are most abnor-
mally nervy animals, extremely hard to tame; it is nothing, for instance, for a
specimen of this kind to take sixteen weeks before ever feeding from hand.
But once they get over their first nerves, and brace their courage sufficiently to
come to hand, they are most engaging animals, with an excellent temperament,
and, I think, with a good memory for people; although it has been my experience
that they never completely lose their distrust, so that the least thing outside
routine, such, for instance, as a sudden movement, or a sneeze, will send them
scurrying for shelter in a panic, and if it is an animal which does not know one
well, it will be some time before the animal can be coaxed back. One cristata
only I have known who allowed himself to be stroked. This was an individual
who seemed to delight in being scratched and rubbed all over; on this animal,
the belly, shoulders, and limbs are covered with bristles which are not harsh to
the touch; the skin of the back when reached through the mass of quills is
completely naked, flesh-coloured, soft and velvety. This nakedness is probably
not a constant character; I have noticed a certain growth of hair under the quills
of the back on other Crested Porcupines, which is also present in //. hodgsoni.
H. cristata definitely sheds the quills much more freely and frequently than the
hodgsum-brachviirus type of animal; and on occasion can do very much more
damage with them; but any Porcupine will draw blood immediately (even
accidentally), or if touched or handled when not in the mood, and speaking from
personal experience it can be agonizingly painful.
Moreover, they can attack with their spiny covering by running sideways
or backwards into their enemy, usually leaving some of the quills in anything
they run into. The quills in cristata may reach extreme length; I once had one
in my possession measuring twenty-one inches. But it is not these, but the
short thick quills which do the damage. The young have sharp little bristles at
birth; and these will develop into sufficiently sharp spines within ten days to
make handling impossible. In captivity there are one or at most two young
in a litter, so far as my experience goes. H. hodgsoni, which has the caudal
rattling quills poorly developed, does not use them except in moments of great
excitement, and even then the noise produced is feeble; but in cristata, in which
these hollow quills are at maximum development, they are normally used con-
stantly, though I have known more than one specimen which appeared quite
unable to make any sound with them at all. The sound produced by cristata is
entirely different from that produced by hodgsoni, and in moments of anger or
excitement it can be terrific; a noisy motor-bicycle is the only thing to which
I can compare it; but I have never been able to ascertain whether all this noise
is caused by the quills, or whether the animal roars at the same time. It must
be stated, not for the first time, that the belief that these animals shoot their
202 HYSTRICIDAE
quills is a mvth. But the origin of that story may be explained as follows: a
Porcupine will normally be carrying a few loose quills on the body (sometimes
these may even be picked out in very tame specimens, as in "Joe," a famous
hodgsoni {or javcviicum;) which lived in the London Zoo for about twelve years
or more). When the animal wakes up suddenly, he shakes himself, and on rare
occasions one of these loose quills is shot out and hurtles across the cage, giving
the effect that the animal has shot it.
The captivity lite is good; best of all Rodents, according to Flower's valuable
paper on the subject; in this paper a specimen is mentioned which attained the
great age of twenty years; but normally I suppose twelve or fifteen would be
the absolute limit. It may be added that their gnawing powers are prodigious
and that they use extreme ingenuity on certain occasions; for instance, I once
saw one trying to shake open a door; after some ineffective pawing attempts,
he ran backwards and then took a run at the door, just as a human being might
do, which I thought showed high reasoning power.
Key to Generic Groups
Tail relatively long, its end bearing a tuft of bristles; cheekteeth rooted,
more brachyodont; spiny covering not highly developed (thick
circular quills on back most often absent); anterior zygomatic root
over P.4. .Atherurus Group. Atheruri
Trichys ; Atheruriis
Tail short, bearing a cluster of highly modified hollow quills; cheekteeth
usually strongly hypsodont, semi-rooted; spiny covering highly
developed, always with thick circular quills present on back;
anterior zygomatic root over middle of toothrow.
Hystrix Group. Hystrices
Tlieciirus, Hystrix
The Alhenirus Group
Ch.\r.^cters. — .\s indicated in the above key. The size is relatively smaller
usually than in the Hvstrix group; the spiny covering less
specialized. Skull with no inflation of nasals, and usually no inflation of frontals.
The thick quills on the back, characteristic of true Porcupines, are not developed
in Trichys nor in the Asiatic section of Atherurus; in the African section of the
latter genus they are usually present. This indicates a higher grade of specializa-
tion for the African forms of this genus, which is paralleled by the Hystrix
group, w^hich are at their lowest in the Indo-Malayan region, and at their
highest in Africa.
Key to the Gener.\ of the Atherurus Group
Skull with well-marked postorbital process, and strong interorbital con-
striction; lower incisors more compressed; tail long, scaly, its tip
bearing a cluster of parallel-sided bristles; body clothed with
flexible spines; a prominent horizontal groove on surface of jugal.
Trichys
TRICHYS 203
Skull with scarcely marked postorbital process (or this absent), and little
interorbital constriction; lower incisors not compressed; tail long,
less naked, its tip bearing a cluster of bristles which are alternately
expanded and contracted ; body clothed with sharper bristles, some-
times mixed with quills; no groove on surface of jugal. Atherurus
Genus i. TRICHYS, Gunther
1876. Trichys, Gunther, Proc. Zool. Soc. London, p. 739.
Type Species. — Trichys lipura, Gunther.
R.\NGE. — Sumatra, Borneo, and southern Malay Peninsula (Malacca,
Perak, etc.).
Number of Forms. — Two.
Characters. — Skull rather long and narrow; a prominent sagittal ridge
present in adult; well-marked postorbital processes present,
behind which skull is considerably constricted; nasals short, narrow, shorter
than the frontals, extending back to anterior zygomatic root. Rostrum slender.
Occipital region high, strong; paroccipital processes not lengthened; bullae
relatively small. Palate straight, moderately wide; hamular process long;
palatal foramina very short, situated far in front of toothrow.
Infraorbital foramen relatively small for a Hystricoid Rodent; no canal for
nerve transmission. Zygoma simple; jugal long, though not extending to
lachrv'mal, a prominent groove running along it, on exterior border, throughout
most of its length. Mandible with angular portion well distorted outwards; the
hinder part of the jaw flattened, as in all Hystricidae (i.e. no backward pro-
longation of angular process). Coronoid high, nearly as high as condyle in type
skull.
Cheekteeth as already described, rooted; toothrow short.
Externally covered with fiat grooved spines scarcely more developed than
in some Neotropical Echimyidae; head and underparts hairy. Tail moderately
long, poorly haired, scaly, its end bearing a cluster of straight bristles. There
is a tendency present for this animal to lose the tail during life. Forefoot rather
broad, with four well-developed digits bearing thick claws, the two central
digits slightly longer than the outer ones. PoUex rudimentar}-. Hindfeet
similar to the forefeet, but longer, and the hallux more developed than the
pollex ; otherwise digits like those of forefoot.
Size smaller than is usual in the family.
Forms seen : lipura, macrotis.
The two species are verv' closely allied, and might be considered as not
more than races; their chief difference lies in the size of the ear.
List of Named Forms
(References and type localities of all Hystricidae are the work of Mr.
G. W. C. Holt.)
Fig. 56. Trichys macrotis, Miller.
B.M. No. 16. II. 16-2; >'• I-
Fig. 57. Trichys m.acrotis, Miller.
B.M. No. 16. II. 16. 2; :, I.
I
TRICHYS— ATHERURUS 205
1. TRICHYS LIPURA, Gunther
1876. Proc. Zool. Soc. London, p. 739.
Borneo.
Synonym: guentheri, Thomas, 1889, Proc. Zool. Soc. London, p. 235.
Kina Balu, Borneo.
2. TRICHYS MACROTIS, Miller
1903. Proc. U.S. Nat. Mus. XXVI, p. 469.
N.-W. Sumatra, Tapanuli Bay.
Genus 2. ATHERURUS, Cuvier
1829. .ATHERURUS, Cuvier, Diet. Sci. Nat. LIX, p. 483.
Type Species. — Hystrix macrourus, Linnaeus. (See Lyon, Proc. U.S. Nat.
Mus. XXII, 1907, p. 584).
Range. — Indo-Malayan : Southern China, Tongking, Hainan, Assam,
Tenasserim, Malay Peninsula, Sumatra. (Other Malay Islands ?)
Africa: Nigeria, Fernando Po, Sierra Leone, Senegambia, Congo, Uganda,
Kenya. (In China occurring as far north as Szechuan.)
Number of Forms. — Thirteen.
Characters. — Skull with short nasals, extending hack about to anterior
margin of infraorbital foramen; frontals long, becoming
arched and slightly inflated in africanus ; usually no postorbital process, but this
can be slightly marked, and usually ver}' little interorbital constriction. Sagittal
crest formed in adult. Zygoma simple, but jugal broader anteriorly, and
tending to be longer in African species than in Malayan ones; in manv cases
it reaches the lachrymal in African group. Bullae relatively small. Palate
essentially as Trichys. Occipital region of skull as in Trichys. Mandible with
low coronoid, rather low condyle; angular process moderately to weakly dis-
torted outwards; traces of a ridge beside the condyle, as found in Erethizontidae
and Chinchillidae, may be present. Infraorbital foramen without canal for
nerve transmission; relatively small for a Hystricoid Rodent, particularly in
africanus, in which the frontal inflation is present. Palatal foramina much
reduced. Cheekteeth as described above; rooted.
Externally covered with spines, stronger and more pow^erful than in Trichys;
head hairy. Tail moderately long, with short spiny hairs on the scales, and a
thick tuft of bristles at the end, the bristles alternately expanded and con-
tracted (up to about five times or more); in the African species these expansions
tend to be nearer to each other and rather more numerous than in the Indo-
Malayan type. Feet essentially as in Trichys. Head and body length up to
525 mm. or perhaps more.
These may be described as much faster-moving animals than members of
the genus Hystrix, judging by specimens observed in the London Zoological
Gardens.
In the African species the thick round quills, characteristic of all the mem-
bers of the Hystrix group, are present; these vary in their development from
scarcelv traceable to rather strong; it may be that with a large series, African
2o6 ATHERURUS
types might come to hand which lack tiicm. They are always absent in the
Indo-Malayan forms so far as I have examined.
Forms seen : africaiius, assomensis, biirrozvsi, centralis, macrourus, sterensi,
tionis, turiieri, zygomaticiis.
The terms are here provisionally treated as two distinct groups, based on
the presence (African species) or absence (Malayan types) of true quills.
Of the Indo-AIalayan forms, from descriptions and from those I have
examined I cannot credit that there is more than one species; trivial skull
characters have for the most part been used to separate the various forms, and
probably all are best regarded as races.
The African types divide sharply into two on skull characters, though the
form turneri seems to me to be intermediate between the two types to a certain
degree; it must also be noted as a form in which apparently the quills are weak.
In africaniis, the trontals are arched, the skull is broader, the infraorbital
foramen, due no doubt to the inflation of the frontals, seems smaller, and the
jugal appears consistently to extend up to the lachrvmal.
In centralis, the frontals are not or less arched, the skull is narrower, and
more like the Asiatic tvpes, and the jugal may or may not extend so far anteriorly.
A. Imrrowsi, based on a skull without a skin, is best regarded as a race oi centralis.
List of N.^med Forms
macrourus Group
1. ATHERURUS MACROURUS MACROURUS, Linnaeus
1758. Syst. Nat. loth Ed. no. 4, p. 57.
No exact locality'.
Synonym: {?)Jasciciilata, Shaw, 1801, Gen. Zool. ii, i, p. 11, pi. 124.
(This name used by Lyon, 1907, for the Malaccan
Trichys.)
orieiitalis, Brisson, 1756, Regn. Anini. p. 131. East Indies.
2. ATHERURUS MACROURUS PEMANGILIS, Robinson
1912. .Ann. Mag. Nat. Hist. 8, X, p. 590.
Johore Archipelago (Malaya) ; Pulau Pemanggil, between Pulau Aor and
Pulao Tioman, S. China Sea.
3. ATHERURUS I\L\CROURUS STEVENSI, Thomas
1925. Proc. Zool. Soc. London, p. 505.
Ngai-Tio, 7'ongking.
4. ATHERURUS MACROURUS ASSAMENSIS. Thomas
1921. Joum. Bombay Nat. Hist. Soc. XXVH, p. 598.
Cherrapunji, Assam.
5. .■\THERURUS M.^CROURUS H,4INANUS, Allen
1906. Bull. Amer. Mus. Nat. Hist. XXH, p. 470.
Hainan.
ft. ATHERURUS MACROURUS TERUTAUS, Lyon
1907. Proc. U.S. Nat. Mus. XXXII, p. 587.
Pulou Ten-itau (Malay Peninsula).
Fig. 58. Atherurus turneri, St. Leger.
B.M. No. 34.6.2.77, (J; X I.
L
Fig. 59. .Atheriris tlrneri, St. Lcger.
B.M. No. 34.6.2.77, (J; X I.
2o8 ATHERURUS
7. ATHERURUS MACROURUS TIONIS, Thomas
iqoiS. Joum. Fed. Malay States Mus. Vol. II, no. 3, p. 105.
Juara Bay, Tioman Island (Malay Peninsula).
8. ATHKRURUS MACROURUS ZYGOMATICUS, Miller
1903. Smiths. Misc. Coll. no. 1420, 45, p. 42.
Pulau Aor, Johore, Malay Peninsula.
africanus Group
9. ATHERURUS CENTRALIS CENTRALIS, Thomas
iSqs. Ann. Mag. Nat. Hist. 6, XV, p. 89.
Monbuttu, Congo, Central Africa.
10. ATHERURUS CENTRALIS BURROWSI, Thomas
1902. Ann. Mag. Nat. Hist. 7, IX, p. 271.
Lower Aruwimi River, Congo.
11. .-ATHERURUS TURNERI, St. Leger
1932. .Ann. \Iag. Nat. Hist. 10, X, p. 231.
West Kenya Colony; Kakamega Forest, near Kaimosi, North Kavi-
rondo.
12. .-XTHERURUS AFRICANUS, Gray
1842. .Ann. Mag, Nat. Hist. X, p. 261.
Sierra Leone.
incertae sedi's
13. ATH1:RL'RUS ARMATUS. Gervais. (Not seen.)
1S54. Nat. Hist. Mamni. i, p. 334.
Senegambia.
(Description evidently based on e.xternal characters only.)
The genus Atlierurus provides a good example of discontinuous distribution,
no form being known living between Assam and Kenya. It is interesting to
note that the genus lives side by side with llystrix throughout much of its
range; and that the primitive member of the Atheninis group {Trichys) is
restricted to the Malav Islands area, exactly as is the primitive member of the
Hystrix group (Tlieciirus).
The Hystrix Group
The Hystrix group differs from the Atheninis group primarily in the reduc-
tion of the tail, which is short, and as indicated above bears a cluster of hollow
"rattling-quills," developed to a greater or lesser degree. These quills have a
flower-like effect, being secured to the tail by a stalk, above which they open
out into the hollow terminal part. When the animal is nervous, the tail is
apparently shaken, and the quills, being lightly attached, are rattled together
to produce the warning signal. I have written at some length above on this
fact, and the habits of certain species of the genus concerning it. There are
always on the back some thick circular quills, the extremities of which are
very sharp.
The cheekteeth in this section are extremely hypsodont, usually semi-rooted.
THECURUS 209
Key to the Genera of the Ilystrix Group
Nasals narrower, shorter, confined to rostrum (essentially Atherurus-likc
in appearance), extending back to about anterior margin of infra-
orbital foramen, shorter than the frontals. Thecurus
Nasals broader, longer, not confined to rostrum, extending back about
to the level of the lachrymal in primitive species, or in progressive
species tending to reach level of posterior zygomatic root; frontals
shorter than the nasals. Hystrix
The nasals of all II\strix seen are considerably broadened, even in primitive
forms \i]i.e javanictim. So far as I can trace, and from our skulls, the percentage
of the nasal length against the occipitonasal length does not exceed 33 per cent
in Thecurus, and averages 31 in our series, 32 in a series of sumatrae the measure-
ments (taken against the "total length" of skull) published by Lyon. In
Hystrix, its lowest adult appears to be a specimen from Flores with the per-
centage 394; two specimens from Java, juvenile or subadult have the percentage
39-6 and 39'8; others measured are over 40 per cent. Some measurements
included below.
Genus 3. THECURUS, Lyon
1907. Thecurus, Lyon, Proc. U.S. Nat. Mus. XXXII, p. 382.
Type Species. — Thecurus sumatrae, Lyon.
Range.— Sumatra, Borneo, and Paragua, Philippine Islands.
Number of Forms. — Three.
Characters. — Like smaller species of Hystrix (next to be described), but
skull transitionary towards the Atherurus type, the nasals
narrow, extending back only to about the level of anterior margin of infraorbital
foramen. Rostrum narrower; skull appearing flatter than in any Hystrix seen.
Frontals longer than nasals; interorbital constriction may be suggested. Palate
not narrowed in front of toothrows; palatal foramina as usual very small;
infraorbital foramen, as in Hystrix, with no canal for nerve transmission. Bullae
relatively small. Cheekteeth strongly hypsodont; pattern as already described
(ultimately the pattern of the teeth appears to become obliterated).
Remarks. — On account of the Athenirus-hke skull (regarding the area of the
nasals) this group may, I think, stand as a genus, though it is not
very well separated from smaller species of Hystrix like javanicum. There is,
however, an undoubted and clear difference in the skulls of these two types.
Lyon in his original genus description compared Thecurus to " Acanthion" and
remarked that the caudal rattling-quills are much smaller; but, as indicated
above, they are essentially in formation as those of H. javanicum (which Lyon
evidently did not see), and H. hodgsoni from Nepal, which Lvon did not use in
his notes for comparing the two tvpes.
He also mentioned a number of skeletal characters in which the two groups
14 — Livin;; Rodents — I
Fig. 6o, Thecurus crassispinis, Gunther.
B.M, No. 92.9.6.171 X I.
Fig. 61. Thecurus crassispinis, Gunther.
B.M. No. 92.9.6.17; X I.
THECURUS 211
differed; such as, for instance (compared with Acanthion), "instead of having
a large laterally compressed neural spine on the axis, that vertebra bears a
relatively short triprismatic spine, not compressed laterally any more than it is
anteroposteriorly," etc. But before accepting such characters as these for
generic purposes it is surely necessary to examine skeletons of all known
Hystrix-group Porcupines.
As indicated above in my notes on external characters of main species in
the family, two very well-marked groups may be recognized: the pumilis group,
in which the covering is at its lowest development among true Porcupmes, and
the crassispinis group, containing a larger animal in which the quills of the
back reach a surprising degree of thickness for a relatively generalized Porcupine
of this type.
Forms seen : crassispinis, pumilis, and a recently acquired specimen from
Sumatra which I take to represent sumatrae.
List of Named Forms
pumilis Group
1. THECURUS PUMILLS, Gunthcr
1879. Ann. Mag. Nat. Hist. 5, IV, p. 106.
Paragua, Philippine Islands.
2. THECURUS SUMATR.^E, Lyon
1907. Proc. U.S. Nat. Mus. XXXII, p. 583.
Aru Bay, east coast of Sumatra.
crassispinis Group
3. THECURUS CR.ASSISPINIS, Gunthcr
1876. Proc. Zool. Soc. London, p. 736.
Mainland of Borneo, opposite Labuan.
Nasal, Front.\l and Occipitonasal Measurements (in mm.) of
British Museum Series of Skulls (other than broken ones)
OCCIPITONASAL
SPECIES NUMBER NASAL LENGTH FRONTAL LENGTH LENGTH
T. pumilis 79-5-317 25 <^- 34 87
94.2. 1. 1 5 27 32 91
T. crassispinis 76.9.20.15 (the type) 33 38 c. no
92.9.6.17 37 40 114
92.10.1.5 34 39 III
95-5-7-7 35 37 lo?
84.5.19.7 35 36 106
Measurements for comparison, of hrachytirus Group of Hystrix in
British Museum (small species with nasal percentage of occipitonasal
length less than 50 per cent)
H. jaiamcum,
juvenile 50.1 2.2. 16 43-5 29 I09
H. brachyurus 5.9.27.1 51 36 I2Z
OCCII'ITONASAL
NASAL LHNGIH
FRONTAL LENGTH
LENGTH
63
34
130
62-5
34
128
43
31
109
45
31
109
^I'S
left
31
5
114
48 r
ght
44
31
5
104
42
32
loO
212 THECURUS— HYSTRIX
Sl'ICIFS NUMIIEU!
H. hniihvurus 3.2.6.74
H. wiillmO),
from Borneo 93. 3. 4. 8
Frost Collkction, 1938.
Not yet registered.
Flores, adult female jtn'aiiiciini
Flores, adult nva\c jmaniciim
East Java, adult male javaiiicum
East Java, old female javaniciiiii
East Java, sub-adult y«f(7/H'c»/«
All other Hystrix, so far as traced, have the percentage of nasals against
occipitonasal length more than 50 per cent e.xcept sometimes H. Icucura (^•j,fide
Lonnberg).
Genus 4. HYSTRIX, Linnaeus
1758. Hystrix, Linnaeus, Syst. Nat. loth. Ed. i, p. 56.
1822. AcANTHiON, Cuvier, Mem. Mus. Hist. Nat. IX, p. 425. (Acaiithion javaiiicum,
Cuvier.) Valid as a subgenus.
Type Species. — Hystrix cristata, Linnaeus.
Range. — The greater part of the African Continent (Morocco, Asben
(Sahara), Upper Egypt, Senegal, Gambia; Uganda, Kenya,
Somaliland, Tanganyika, Portuguese East Africa, South-west Africa, South
Africa); Italy and Sicily; Palestine, Syria, Asia Minor, Mesopotamia, South
Arabia (specimens in B.M.), Afghanistan, probably Persia; Transcaucasia
(Talysh); South-western Siberia (Turkmenia, Semirechia, Kopet-Dag moun-
tains, Karakum), (Vinogradov); Baluchistan; Peninsular India (Punjab, Rajpu-
tana. Central Provinces, Palanpur, Cutch, Kathiawar, Deccan, Mysore, Coorg,
Nilgiris, Malabar); Ceylon; Nepal, Sikkim, Bhutan, Assam; Burma, Tenas-
serim. South China (Szechuan, Y'unnan, Fukien, Anhwei); Hainan; Malay
Peninsula, Sumatra, Java, Borneo, Sumbawa, Flores.
Number of Forms. — .Approximately thirty-five.
Ch.\r.\cters. — Nasals, even in the most primitive forms, longer and broader
than in Thectirits, extending about to lachrymal level in
brachyurus group, progressively lengthened in most other species; broader to a
degree and much longer in suhcristatus group; relatively short in leucnra group,
and not wider behind than in front; relatively short in afrkaeustralis group but
enormously broadened, much wider behind than in front; considerably broader
behind, and also much lengthened in cristata group, ultimately approaching
the level of the posterior zygomatic root. In these larger African species the
skull becomes much arched. A prominent sagittal crest normally present in
adult. Occiput thick, strongly ridged, prominent. Bullae relatively small;
paroccipital processes not much lengthened. Palate broad, extending about to
end of toothrows behind, and straight; not depressed in front of toothrows;
HYSTRIX 213
palatal foramina short, far in front of toothrow. Infraorbital foramen of
moderate size for a Hystricoid Rodent, relatively small in some in which the
nasals reach their maximum inflation ; no canal for nerve transmission. Zygoma
broad but simple; jugal not approaching lachrymal as a rule; zygomatic plate
projected forwards, appearing as an anterior prolongation of zygoma; lachrymal
moderately large. Incisors broad. Mandible with hinder part flattened, the
angular portion powerfully ridged and distorted outwards. Coronoid process low'.
Externally becoming very large in progressive species ("38 inches" and
810 mm. the largest (measured) skins seen, galeata type). Forefeet broad, with
four well-developed digits bearing thick claws; hindfeet longer but essentially
similar except that the hallux is quite well developed, the two central digits
slightly longer than the outer ones.
I have already written at length on the external characters as regards arrange-
ment of head-crest, body quills and spines, and caudal rattling-quills, of the
various species, on pp. 199, 200.
The cheekteeth are essentially as already described under the heading
"Family Hystricidae" (p. 198). They are strongly hypsodont, the pattern is
long preserved though ultimately obliterated, and the premolar is shed late in life.
Remarks. — The genus is frequently divided into two, Hystrix and Acan-
thion. Great as are the differences between the highly specialized
H. cristata (type of Hystrix), and the relatively primitive H. javanicitm (type of
Acanthion), it becomes clear that so many intermediate forms exist that this
classification cannot be retained. This is made very clear in a paper by Lonn-
berg, 1923 (on the Chinese Porcupine H. subcristatus, Swinhoe with remarks
on other members of the genus, Arkiv. for Zoologi, Band 15, No. 18, pp. i-io),
and in other papers by this author.
In 1912 Aliller restricted the genus Hystrix to the European and African
species only, on account of the "inflation of facial regions of skull at maximum
for the family; nasal bones extending to glenoid level." But this appears to
include in Hystrix the Chinese subcristatus, currently referred to Acanthion, and
to exclude the African crested species africaeaustralis, which is naturally a
Hystrix. It is quite clear that on nasal structure alone this genus will not divide
into two.
In my opinion, in an animal of this description, the external characters
(development of quills, etc.) must be regarded as being just as important as any
cranial character. There is a definite break in the species between cristata,
africaeaustralis, leucura groups (Crested Porcupines) on the one hand, and
subcristatus {}) (skin not seen, but description fits in with Acanthion as here
suggested), klossi, brachyurus, javanicum types on the other. For the present I
suggest that Acanthion may be used subgenerically for the latter group (with
crest poorly developed, vestigial or absent; caudal-quills moderately to poorly
developed; body quills thick but without the profuse mantle of manv-ringed
longer quills present and covering them). If on the other hand subcristatus
proves to be an intermediate between the two groups, the name Acanthion will
have to be placed in svnonymv.
214 HYSTRIX
Subgenus ACANTHION
(With characters as just indicated; quills with one ring only.)
There are two groups here recognized, a "short-nasal" group containing
bidchxurus, of the Malay Peninsula and Islands (with loiigicauda and iniiUeri as
races or synonvms), and jai-aiiiciim, and a "long-nasal" group. H. braclivurus
group (nasals in percentage of occipitonasal length averaging 45-3 in our
specimens; 49 in five measured by Lyon (nasals against "total length" of skull).
H. javankum has an average percentage of 40-9 in the few adult skulls repre-
sented in London ; sunibazcae, Schwarz, from Sumbawa has a slightly lower
measurement, 36-2, in the figures published. This might or might not be a race
oi jaraniciim; specimens received from Flores (which is beyond Sumbawa east-
wards from Java), appear quite indistinguishable from typical javanicum. I
divide this group into two sections, the typical, and the javanicum section
(several skins of which have been seen), these sections differing from each other
in the external characters indicated on pp. 199, 200. Although between these
two sections there is quite a clear difference in the material examined, perhaps
if enough material was collected, the two types would intergrade.
//. subcristatiis group (with long nasals) contains siibcristatiis, klossi (with
mills!, based on skulls the external characters of which are unknown, pro-
visionally treated as a race), and hodgsoni. Nasals in percentage of occipitonasal
length in subcristattis S^'(^57'7 (Lonnberg), (and mesopterygoid space unusually
wide in our skulls); the percentage in hodgsoni averages 55-6 in four measured;
the same in klossi (four measured), is 53-6. This group is divided into two
sections, in precisely the same way as the braclivurus group; in hodgsoni the
external characters are more primitive than in klossi \ but probably intergradation
would take place in these external characters, if enough specimens came to
hand. These characters have been noted on p. 200.
The measurements of the skulls just quoted are:
OCCIPITONASAL
SPECIES
NUMBER
NASAL LENGTH
FRONTAL LENGTH
LENG1
H. klossi
14. 12.8.224
80
31
139
,- -, (type)
14. 12. 8. 223
75
34
145
1) 7»
I 5. 1 1.4.220
71
30
i3'3
,, ,, (nnllsi\ type
21.7. 16.4
68
Zi
128
H. hodgsoni
S3.8.16.I I
64
24
IIS
») )»
45.1.8.8
6S
20
116
)j »)
21.10.4.3s
66
23
115
i< ti
79.11. 21. 637
61
24
114
Two of the main difficulties of dealing with animals of this description are
(i) the rarity, and (2) the frequently bad condition, in which they come to hand.
But even if these notes are based on insufficient material, they do at least
give a preliminary survey of all the main species of the whole genus, and not
sections of it (for instance, Lonnberg's paper compares only suhcristatus with
the larger crested types; Lyon's paper compares only the Indo-Malayan ones.
I have seen no paper which compares klossi, hodgsoni, etc., with either of these,
or either of these groups with each other).
HYSTRIX 215
Subgenus HYSTRIX
(With more highly specialized development of external characters; crest long,
fully developed; caudal rattling-quills at maximum development; a profuse
Fig. 62. Hystrix cristata, Linnaeus.
(From Miller's Catalogue of the Mammah of Western Europe.)
xj.
growth of very long many-ringed quills covering the short thick ones of
Acanthion. For detail notes see p. 213.)
Lonnberg states that these Porcupines (with subcristattis) should divide into
three groups, as follows:
2i6 HYSTRIX
"Nasal cavity widened chiefly by means of prolongation backwards of
nasals. Proc. nasales of premaxillaries truncate behind, only little
widened, suhciistatiis."
"Nasal cavity much enlarged by extremely broad proc. nasales of pre-
maxillaries . Iciiaiiii, ga/eata."
(He suggests this group might divide into two, one for kuciiici
(Indian, Asiatic types), the other for the African galeata.)
" Nasal cavity enlarged by expansion of nasals, proc. nasales of premaxillaries
wedge-shaped behind, not or only moderately enlarged, africae-
australis, cristata."
(He suggests that this group might also divide into two, one for
cristata and seiiegalica type, one for ajricaeaiistralis.)
But there is a very profound difference betw'een leiicura and galeata. Skulls
referred to galeata in London seem to vary individually in the shape of the
nasals. I believe that all species of African Porcupines excepting africacaiistralis
would prove to be referable to one species cristata if enough of them came to
hand, and that the shape of the nasals would be found to vary individually so
that galeata would become merged into cristata. H. leuctira with hirsutirostris
seems to me to be a perfectly natural group, sharply differentiated on nasal
structure from all African Porcupines. The nasals are not or scarcely broader
posteriorly than anteriorly; the whole skull lacks that broadening characteristic
of African Crested Porcupines. Further, the nasals in percentage of occipito-
nasal length are short; (47-49 kiiciira, 48- 2-49-6 liirsiitirostris) (fide Lonnberg);
a leiicura measured for comparison with these figures has the percentage Sf".
This is markedly shorter even than in africacaiistralis.
Russian authors give hirsutirostris full specific rank, but there seems no
reason to believe that it is distinct from the Indian
leucura. I am therefore treating all named forms of
this group as subspecies of the earlier name leucura.
The africaeaustralis group, from South and Southern
Africa, appears on the material examined to be clearly
separable from the cristata group as here understood.
Compared with leucura, the nasals are much broadened,
always as far as seen considerably broader posteriorly
i^^Sy than anteriorly; compared with the cristata group, they
are short (percentage of occipitonasal-nasal length 54-
^^■2, fide Lonnberg; this percentage slightly exceeded
(55-9), in British Museum material). H. stegmanni, not
seen, appears from Lonnberg's percentage figures and
remarks to belong in this group.
The cristata group contains all other African Porcu-
pines,as here understood. The nasal lengthis maximum
for the genus, and is combined with great broadening, but the shape is very
variable both individually and between some of the species (cristata, per-
centage 58-9-65, senegalica, 69; galeata, 6i-i-66-8; galeata ambigua. 60-7,
Fig. 63. HvsTRix
CRISTATA, Linnaeus.
Cheekteeth; i^.
HYSTRIX 217
fide Lonnberg). The nasals extend much farther back than in the africae-
australis group. //. aeritla, from Ashen, is a small form allied to senegalica.
In my opinion probably all North African Porcupines are no more than races
of cristata, including uccidanea, not seen, as figured by Cabrera.
Forms seen: aeriilti, africacaustralis, " hengalensis," brachyurus, cristata,
cuneiceps, "curieri," galeata, hirsutirostris, hodgsoni, javanicum, klossi, leucura,
millsi, miilleri, senegalica, schmidti, somaliensis, subcristatus.
List of N.ameu Forms
Subgenus Acanthion, Cuvier
brachyurus Group
(javanicum section)
1. HYSTRIX JAVANICUM, Cuvier
1822. Mem. Mus. Xat. Hist. IX, p. 431.
Java. (Occurs also in Flores.)
Synonym: torquata, van der Hoev, 1836, Tijdschr. iii, p. 110. Java.
breiispinosa, Wagner, 1844, Schreber, Saug. Suppl. \\ , p. 20.
Java.
ecaudala, van der Hoev, 1836, Tijdschr. iii, p. no. Java.
2. HYSTRIX SUMBAWAE, Schwarz
191 1. Ann. Mag. Xat. Hist. 8, VH, p. 639.
Dompu, Sumbawa, East Indian Archipelago.
(typical section)
3. HYSTRIX BRACHYURUS BR.ACHYURUS, Linnaeus
1758. Syst. Nat. loth Ed. p. 57, no. 5.
Malacca.
Synonym : ^o/fi. Gray, 1866, Proc. Zool. Soc. London, pi. 31, p. 310.
flemiiigi. Gray, 1847, Proc. Zool. Soc. London, p. 103.
4. HYSTRIX BR.ACHYURUS LONGICAUDA, Marsden
181 1. Hist. Nat. Sumatra, 3rd Ed. p. u8.
Sumatra.
5. HYSTRIX BR.ACHYURUS MCLLERI, Marshall
1871. Proc. Zool. Soc. London, p. 235. (See Lvon, Proc. U.S. Nat. Mus. XXXII, 1907,
p. 580.)
Padang, Sumatra.
(An animal of this type occurs in Borneo.)
suhcristattts Group
{hodgsoni section)
6. HYSTRIX HODG.SONI, Gray
1847. Proc. Zool. Soc. London, p. loi.
Nepal, Himalayas.
Synonym: bengalensis. Blyth, 1851, Joum. .As. Soc. Bengal, XX, p. 170.
alophus, Hodgson, 1847, Joum. .As. Soc. Bengal, XVI, p. 771.
Himalayas.
2i8 HYSTRIX
(typical section)
7. HYSTRIX KLUSSI KLOSSI, Thomas
1916. Ann. Mag. Nat. Hist. 8, XVII, p. 139.
Tenasserim Town.
S. HYSTRIX KLOSSI MILLSI, Thomas
1922. Journ. Bombay Nat. Hist. Soc. XXVIII, no. 2, p. 431.
Sangrachu, Assam.
9. HYSTRIX SUBCRISTATUS SUBCRISTATUS, Swinhoe
1870. Proc. Zool. Soc. London, p. 638.
South China; Fokien Province.
10. HYSTRIX SUBCRISTATUS PAPAE, Allen
IQ27. Amer. Mus. Nov. no. 290, p. 3.
Hainan.
Not allocated to Group
11. HYSTRIX YUNNANENSIS, Anderson
1878. Anat. and. Zool. Res. Yunnan, p. 332.
West Yunnan.
(According to Thomas based on a short-nosed species allied to javanicum.)
Subgenus Hystrix, Linnaeus
leiiciira Group
12. HYSTRIX LEUCURA LEUCURA, Sykes
1 83 1. Proc. Zool. Soc. London, p. 103.
India; Sayul of Mahrattas.
Synonym: zeylonensis, Blyth, 1851, Journ. As. Soc. Bengal, XX, p. 171.
Ceylon.
malaharica, Sclater. 1865, Proc. Zool. Soc. London, p. 353.
Cochin, India.
indica, Gray & Hardwicke, 1833-34, 111. Indian Zool. ii,
pi. 14.
13. HYSTRIX LEUCURA CUNEICEPS, Wroughton
1912. Journ. Bombay Nat. Hist. Soc. XXI, p. 771.
Nokania, Cutch, India.
14. HYSTRIX LEUCURA HIRSUTIROSTRIS, Brandt
1835. Mamm. Exot. Nov. p. 39.
Afghanistan.
15. HYSTRIX LEUCURA BLANFORDI, Muller
191 1. Sitz. Ber. Ges. Nat. Freunde Berlin, p. 121.
Jalk, Baluchistan.
16. HYSTRIX LEUCURA SATUNINI, Mullei
1911. Sitz. Ber. Ges. Nat. Freunde Berlin, p. 117.
Transcaspia, Geok Tepe, east of Caspian Sea, 75^' E., 38" N.
17. HY.STRIX LEUCURA MERSINAE, Muller
191 1. Sitz. Ber. Ges. Nat. Freunde Berlin, p. 122.
Mersina, south-east of Taurus, Asia Minor.
HYSTRIX 219
i8. HYSTRIX LEUCURA AHARONII, Muller
191 1. Sitz. Ber. Ges. Xat. Freunde Berlin, p. 123.
Palestine; Emmaus, west of Jerusalem.
19. HYSTRIX LEUCL'RA SCHMITZI, Muller
191 1. Sitz. Ber. Ges. Nat. Freunde Berlin, p. 126.
Palestine; Ain Dcheier, N.-W. of Dead Sea, Jordan valley.
20. HYSTRIX LEUCURA NARYXENSIS, Muller
1919. Sitz. Ber. Ges. Nat. Freunde Berlin, p. 67.
Naryn, Turkestan.
21. HYSTRIX LEUCURA MESOPOTAMICA, Muller
1920. Zool. Anzeiger, 51, p. 198.
Jebel .-Xbdul Azir, N.-E. Syria; 40° 20' E., 36° 20' N.
africaeaustralis Group
22. HYSTRIX AFRIC.\E.AUSTR.\LIS AFRICAEAUSTRALIS, Peters
1852. Reise nach Mossambique, Saugeth. p. 170.
South-East Africa; Querimba, Tette; 11° to 17° south.
Synonym: copensis. Grill, 1858, Zool. Anteckningar af. J. F. Victorin,
p. 19.
23. HYSTRIX AFRICAEAUSTR.A.LIS PRITTWITZI, Muller
1910. Sitz. Ber. Ges. Nat. Freunde Berlin, p. 311.
Tabora, Tanganyika Territon.-.
24. HYSTRIX AFRICAEAUSTRALIS ZULUENSIS, Roberts
1936. Ann. Trans. Mus. XVIII, p. 240.
Zululand, White Umfolosi River.
25. HYSTRIX STEG.M.AXNI. Muller
1910. .\rch. f. Naturgesch. Jahrg. 76, Band i. p. 186.
Kissenji, north-east of Lake Kivu, Tanganyika.
cristata Group
26. HYSTRIX CRISTATA CRISTATA, Linnaeus
1758. Syst. Nat. i, loth Ed. p. 56.
Near Rome, Italy. (See Miller, Cat. Mamm. W. Europe, 1912, p. 543.)
Synonym: cuiieri. Gray, 1847, Proc. Zool. Soc. London, p. !02.
Locality not known.
(?) datibentoni, Cuvier, 1822, Mem. Mus. Nat. Hist. IX,
p. 431. Locality unknown; perhaps best regarded as
unidentifiable.
alba, de Selys-Longchamps, 1839, Etudes de Micro-
mamm. p. 152, nom, nud.
europaea, Kerr, 1792, Anim. Kingd. p. 213.
(Some specimens in B.M. labelled " moroccana" ; the reference to this name
has not been traced.)
27. HYSTRIX CRIST.\TA OCCIDANEA, Cabrera
1924. Bol. Soc. Esp. Hist. Nat. XXIV, p. 220.
Mogador, West Morocco.
28. HYSTRIX CRIST.ATA SEXEG.A^LICA, Cuvier
1822. Mem. Mus. Hist. Nat. IX, p. 430.
Senegal, West .\frica.
220 HYSTRIX— CUNICULIDAE
29. HYSTRIX CRISTATA AKRULA, Thomas
1925. Ann. Mae. Nat. Hist, f), XVI, p. 196.
Aouderas, Asben, Sahara.
30. HYSTRIX GALEATA GALEATA, Thomas
1S93. Ann. Mag. Nat. Hist. 6, XI, p. 230.
Lamu, Kenya.
31. HYSTRIX GALEATA SOMALENSIS, Lbnnberg
1912. Kungl. Sv. Vet. Akad. Handl. Band. 48, no. 5, p. 109.
Njoro, Guaso Nyiro, North Kenya.
32. HYSTRIX GALEATA AMBIGUA, Lbnnberg
1908. Sjost. Kilimanj. Meru. Exp. p. 29.
Kibonoto, Kihmanjaro, East Africa.
33. HYSTRIX GALEATA LADEMANNI, Muller
1910. Sitz. Ber. Ges. Nat. Freunde Berlin, p. 314.
Kondoa-Irangi, Tanganyika.
34. HYSTRIX GALI:ATA CONRADSI, Muller
IQIO. Sitz. Ber. Ges. Nat. Freunde Berlin, p. 314.
Ukerewe Island, Lake Victoria.
35. HYSTRIX GALEATA LONNBERGI, Muller
1910. Sitz. Ber. Ges. Nat Freunde Berlin, p. 315.
Kilimanjaro, East Africa.
The family Hvstricidae is known fossil from the Upper Miocene, from the
Old World only. '
HYSTRICIDAE :
SPECIAL WORKS OF REFERENCE
Waterhoi'se, 184S, Natural Historj- Mammalia; Rodentia (Vol. II).
PococK, 1922, Proc. Zool. Soc. London, 1922, p. 365. External Characters of some
Hystricomorph Rodents.
TuLLBERG, 1899, Nova Acta Reg. Soc. Sci. Upsaliensis, XVIII, 3, i.
Lyon, Porcupines of the Malay Peninsula; Proc. U.S. Nat. Mus. XXXII, 1907, pp.
575-594-
LoNNBERi;. On the Chinese Porcupine Hystrix siihcristatiis. Swinhoe, .Arkiv. for Zoologi,
Bd. 15, no. 18, p. I, 1923.
Miller, Catalogue of Mammals of Western Europe, 1912, p. 542. Hvstricidae. {Hystrix
cristata).
Family CUNICULIDAE
1S96. Thomas: Hvstricomorpha: Dasyproctidae, part.
1899. TuUberg: Hvstricomorph.a : Caviidae, part.
1918. Miller S: Gidley: Hv.sTRicoiD.y- : Family Cuniculidae.
1924. Winge: Family Hvstricidae; Dasyproctini, part, Dasyproctae, part.
1928. Weber: Hystricoidea: Caviidae, part; subfamily Dasyproctinae, part.
Geographical Distribution. — Tropical America, from Mexico to South
Brazil.
Number of Genera. — One.
Characters. — Skull highly abnormal, the greater part of the maxillary and
jugal expanded to form large bony cheek-plates, the surface
CUNICULIDAE: CUNICULUS 221
of which tends to become rugose. Cheekteeth hypsodont, semi-rooted, charac-
terized by deep re-entrant folds which isolate as long islands on crown surface.
External form heavy, terrestrial, the limbs not lengthened; feet with digits of
sub-ungulate type, the claws extremely thick; hindfoot perissodactyle, with
three main digits, but both D.5 and hallux present though strongly reduced;
forefoot artiodactyle, with four main digits.
Clavicles not suppressed, but (said to be) incomplete. According to Tull-
berg, the carpus lacks a free centrale, alone of all Rodents examined by him
excepting the Hystricidae.
Rhm.^rks. — The diflFerences between this genus and the Dasyproctidae with
which it has often been associated have been discussed at length
by Pocock (Proc. Zool. Soc. London, 1922, p. 424), who, following Miller &
Gidley, refers the animal to a distinct family. I am entirely in agreement
with this classification. The unique skull structure in the family indicates
evolutionary development along a very different line not only from that of
Dasyproctidae but from all other Rodents; the feet do not agree with those of
the Dasyproctidae, nor are the limbs lengthened; the cheekteeth do not agree
exactly with those of the Dasyproctidae. (The relationships of Dasyproctidae
as compared with Caviidae, with the latter of which Cuniculus has also been
associated, have been fully discussed when dealing with the family Dasy-
proctidae.)
As indicated already, the cranial characters of this genus difTer entirely from
those of any other Rodent; in this particular it must be looked upon as the
most aberrantly specialized member of the whole Order.
Genus i. CUNICULUS, Brisson
1762. Cuniculus, Brisson, Regn. Anim. ed. 2, p. 13.
1799. Agouti, Lacepede, Ordres et Genres Mamm. 9. Agouti paca (^Mus paca,
Linnaeus).
1807. CoELOGENUs, Cuvier, Ann. Mus. Hist. Nat. Paris, X, p. 203. (Mus paca,
Linnaeus.)
1924. Stictomys, Thomas, Ann. Mag. Nat. Hist. 9, XHI, p. 238. {Coelogenys tacza-
ncnuskii, Stolzmann.)
Type Species. — Mus paca, Linnaeus.
R.WGE. — Forms are named from Mexico, Panama, Cayenne, Ecuador,
Colombia, Brazil, Venezuela. Specimens in British Museum from
Paraguay, Peru; according to Thomas ranging to South Brazil.
Number of Forms. — Ten approximately.
Characters. — Skull with zygomatic region abnormally modified by out-
growth from maxillary and jugal, of bone forming a cheek-
plate which extends downwards and conceals a large part of the mandible, the
maxillary' part ot this plate being deeply hollowed internally. The infraorbital
foramen is smaller than in other Hystricoidae, in adults becoming strongly
reduced, being dwarfed by the cheek-plate. Infraorbital foramen with a separate
Fig. 64. CrsicuLUS paca, Linnaeus.
B.M. No. 13.10.24.61, a; ■ slightly more than 1.
CUNICULUS
233
canal for nerve transmission. Nasals broad, relatively short; frontals broad,
very long; parietals depressed, and a sagittal ridge formed in adult; well marked
postorbital processes occur at the suture of the frontals and parietals. Parocci-
pital processes thick and rather long; bullae relatively small. Palate broad, not
constricted anteriorly, the anterior part extending beyond the toothrow as a
i
Fic. 65. Cltmiculus paca, Linnaeus.
Cheekteeth: B. M. No. 13. 10. 24. 61, (J; x 2J.
narrow shelf bordered by two high longitudinal ridges which extend nearly to
the incisors, and on either side of which lie the enormous cavities caused by the
cheekplates. Palatal foramina obsolete. On account of the cheek-plate, the skull
of the typical species appears nearly as broad as long. There is a marked
tendency for the cheek-plate in old specimens to become rugose, and for the
frontals to assume a similar character, this being apparently especially marked
224 CUNICULUS
in males. The hinder part ot the mandible is more or less flattened, as in
Ilystricidae; coronoid process relatively low; degree of distortion outwards of
angidar process moderate. Cheekteeth rather complex; upper series apparently
with two inner and three outer re-entrant folds, except M,3, which is the largest
tooth of the series, and in which the number of folds appears to be reversed.
Sometimes there is a tendency for M.i to become reduced in size and elements,
with wear. Most of the folds isolate as long persistent islands almost immedi-
ately; the unworn tooth shows, as usual in this Order, an extremely complex
pattern. Some of the isolated folds become suppressed with wear. Lower teeth
with one outer, three inner folds each; the premolar may have an e.xtra inner
fold.
Incisors thin, compressed.
Externally typicallv the fur is harsh, the sides of the body with longitudinal
rows of spots. Hindtoot with D.5 much reduced, but with moderate claw
(though less strong than those of D.2, 3, 4); hallux rudimentary; three main
digits long, bearing very sharp somewhat hoof-like claws. F"orefoot with four
main digits; D.2 is longer than D.5, but shorter than the central pair; pollex
represented by a knob. Tail obsolete. Form heavy, and size large, one of the
largest members of the Order. (The largest skin examined is 6S5 mm. head and
body, but this measurement probably may be exceeded.)
It is probable that the habits of these animals are not cursorial (com-
pared with Dasyproctidae); Pacas are said to take to the water when
alarmed.
In the tdcsdiiuicskii group (Mountain Pacas), the fur is thicker, less harsh.
This group was referred to a distinct genus by Thomas, on account of "the
narrow compressed claws and much more profusely granulated soles; cranially
by the proportionately longer nasals, much smaller orbits, more anteriorly
situated postorbitals — the zygomata narrower, generally much less rugose,
though as usual there is much variation in this respect — finally the incisors are
orthodont." Mammae i — 1 =4 (sierrae), (Thomas).
But there are far too many essential characters shared by the two groups
for there to be any question of even subgeneric separation, in my opinion.
These characters indicate that the plains Pacas and mountain Pacas belong to
distinct species. It may be stated that in a skin of sierrtif, the claws seem even
thicker than a specimen oi paca with which it was compared; there is certainly
no generic difterence so far as claws are concerned; (compare, tor instance,
Chimliilla with Lagoitomiis; Ciiviti with Kerodun); when two groups have gone
so far together in specialization (cranial characters), it not only seems unneces-
sary but bad classification to give them generic names on small cranial differences
such as the above.
An interesting account of the formation of the cheek-pouches of the genus
is given by Mr. R. I. Pocock (under the name of Cotlogoiys), Proc. Zool. Soc.
London, 1922, p. 37(1.
Forms seen : paca, fiiiantii, taczaiimcskii, sierrac.
I can see no specific difference between the last two forms, the latter being
described as a distinct species.
CUNICULUS aas
List of Named Forms
(References and type localities collected by Mr. G. W. C. Holt.)
paca Group
1. CUNICULU.S PACA PACA, Linnaeus
1766. Syst. Nat. i, p. 81.
Cayenne.
.Synonym: fulvus, Cuvier, 1807, .\nn. Mus. X, p. 207.
suhniger, Cuvier, 1807, Ann. Mus. X, p. 206. Tobago.
2. CUNICULUS PACA NELSONI, Goldman
1913. Smiths. Misc. Coll. LX, no. 22, p. g.
Catemaco, Southern Vera Cruz, Mexico.
3. CUNICULUS PACA VIRGATUS, Bangs
igo2. Bull. Mus. Comp. Zoo!. Har\-ard Univ. XXXIX, p. 47.
Divala, Chiriqui, Panama.
4. CUNICULUS PACA ALBA, Kerr
1792. Anim. Kingd. p. 217.
.St. Francis River, Brazil.
5. CUNICULUS PACA MEXIANAE, Hagmann
1908. Arch. Rassenbiol. 5, p. 25.
Mexiana Island, Amazonian estuary, Brazil.
6. CUNICULUS PACA GUANTA, Lonnberg
1921. .\rk. f. Zool. Band XIV, no. 4, p. 45.
Pacto, below Gualea, Ecuador.
7. CUNICULUS PACA SUBLAEVIS, Gervais
1854. Ger%'ais Mamm. i, p. 326.
Colombia.
taczanowskii Group
8. CUNICULUS TACZANOWSKII T.-VCZANOWSKII, Stolzmann
1885. Proc. Zool. Soc. London, p. 161.
Ecuador; forests on either slope of the .Andes, between 6,000 and
10,000 ft.
9. CUNICULUS TACZANOWSKII ANDINA, Lonnberg
1913. Ark. f. Zool. Band VIII, no. 16, p. 28.
Mount Pichincha, Ecuador.
10. CUNICULUS T.A.CZ.ANOWSKII SIERRAE, Thomas
1905. .Ann. Mag. Nat. Hist. ser. 7, XV, p. 589.
I'edregosa Mountains, Sierra de Merida, Venezuela.
Tate quotes " thomasi," nom. nud. (.') ex Thomas, Bull. Amer. Mus. Nat.
Hist., LXVHI, 2, p. 314, 1935.
CUNICULIDAE:
SPECIAL WORKS OF REFERENCE
POCOCK, 1922, Proc. Zool. Soc. London, 1922, p. 365. External Characters of some
Hystricomorph Rodents; p. 376, account of the mechanics of the cheek-pouches.
15 — Living Rodents — I
226 CHINCHILLIDAE
Watebhoise, 1848, Natural History Mammalia, Rodentia.
Tate, 1935, Taxonomy Neotropical Hvstricoid Rodents, Bull. Amcr. Mus. Nat. Hist.
LXVHI, p. 295.
TvLLBERG, Nova Acta Reg. Soc. Sci. Upsaliensis, XVHI, 3, i, 1S99.
Family CHINCHILLIDAE
1896. Thomas: Hvstricomorpha: Family Chinchillidae.
1899, TullberK: Hystbitomorpha: Family Chinchillidae.
1918. Miller &: Gidley: Hvstbicoidae: Family Chinchillidae.
1924. Winge: F'amily Hystricidae, part; "Eriomyini" (= Chinchillini).
1928. Weber: Hystricoidea: Family Chinchillidae.
Geographical Distribution. — Western and Southern South America,
from Peru, Bolivia and North Argentina
southwards.
Number of Genera. — Three.
Characters. — Cheekteeth evergrowing, the pattern one of transverse plates.
Mandible with no sharply defined ridge for attachment of
masseter lateralis; the angular portion less strongly distorted outwards than is
usual in Hystricoidae; a weak ridge below condylar process presumably for
attachment for masseter medialis may be present, foreshadowing the structure
present in Cavioidae, but much shorter and less developed than in that group.
Jugal usually in contact with lachrymal; zygoma simple, but normally thickened
anteriorly. A tendency present towards great inflation of inastoids and bullae;
paroccipital processes relatively long. Palate much constricted anteriorly; palatal
foramina usually very long, narrow. Incisors relatively narrow. External form
slender, the hindfeet lengthened (semi-saltatorial or cursorial Rabbit-like types);
digits of hindfoot three or four, D.5 when present extremely reduced, perhaps
functionless.
The lachrymal is large; part of the lachrymal canal is open on side of rostrum
in front of orbit.
A skeleton has been examined in each of the three genera, and each presents
the feature that the fibula, though not fused with the tibia, is excessively reduced,
a structure rather different from that of other Hystricoid Rodents examined for
this character.
The Chinchillidae fall into two well-marked groups, one containing Lago-
stomiis only, the other ChiiicliiUa and Lagidiiiiii.
The difierences between the skulls and the digits of these groups are rather
extreme, and they have been regarded as subfamilies (Pocock, 1922). But these
difl^erences seem rather adaptive; and I have seen it stated that Chinchilla has
bred with "the much larger but related Vizcacha" (Jcnnison, 1929). They are
therefore here treated as groups only.
Key to the Groups of Chinchillidae
Paroccipital processes long, standing apart trom bullae, which are not
specially inflated; occipital region of skull strong, prominent; skull
CHINCHILLIDAE: CHINCHILLA 227
more prominently ridged for muscle attachment; digits of hind-
foot three, the claws heavy, prominent, excessively sharp; palatal
foramina shorter; cheekteeth, excepting M.3, upper series, bila-
minate. Lagostomus Group. Lagostomi
Lagostomus
Paroccipital processes closely applied to bullae, not or less elongated;
bullae considerably to extremely inflated; occipital region of skull
weak; skull not prominently ridged for muscle attachment; digits
of hindfoot four; the claws blunt and weak; palatal foramina long
and narrow; cheekteeth trilaminate. Chinchilla Group. Chinchillae
Chinchilla, Lagidium
The Chinchilla Group
Characters. — As indicated in the above key.
Key to the Genera of the Chinxhilla Group
Bullae abnormally inflated, the mastoids showing prominently in superior
aspect of skull. Jugal usually not in contact with lachrvmal.
Laminae of cheekteeth straight. Chinchilla
Bullae less abnormally inflated, the mastoids scarcely showring in superior
aspect of skull. Jugal in contact with lachrj'mal. Laminae of
cheekteeth curved. L.agidium
Genus i. CHINCHILLA, Bennett
1829. Chinchilla, Bennett, Gard. and Menagerie Zool. Soc. i, p. i.
1830. Eriomys Lichtenstein, Darstell. Saug. VI, pi. 28. (Eriomys chinchilla, Lichten-
stein).
Type Species.— Mw laniger, Molina.
Range. — Chile. ? Bolivia.
Number of Forms. — One is recognized.
Characters. — Mastoids and bullae abnormally inflated, the mastoids
showing prominently each side and at back of skull. Con-
siderable interorbital constriction evident. No canal for nerve transmission in
infraorbital foramen. Palatal foramina very long; palate narrow, considerably
so anteriorly. Jugal usuallv not extending to lachrvmal, broad. Paroccipital
processes moderate, closely applied to and dwarfed by the bullae. Mandible with
narrow angular process, which is sharply drawn backwards; the ridge outside
the condylar process weak.
Cheekteeth like Lagidium (next to be described), but the laminae straighter;
three lobes per tooth, the hinder one in M.3 pointing backwards, as a heel.
The anterior lobe of the lower teeth short, reduced.
Externally, with very soft fur; the tail long, though not as long as head and
body, heavily haired throughout. Hindfeet long and narrow; stiff bristle hairs
Fig. 66. Chinchilla laniger, Molina.
B.M. No. 1. 8. 24. 1, S; ■■ '■i-
Fig. 67. Chinchilla laniger, Molina.
B.M. No. 1.8.24.1, <i; ■ 'i-
CHINCHILLA— LAGIDIUM 229
present on inner digit; three main digits; D.5 placed high on foot, and not
nearly reaching base of D.4; extremely short. Ear large. Forefoot short; four
main digits; pollex represented by a tubercle. Rudimentary cheek-pouches
present (Pocock).
Forms seen : laniger.
List of Named Forms
(References and type localities of all Chinchillidae are the work of Mr.
G. W. C. Holt.)
1. CHINCHILLA LANIGER, Molina
1782. Sagg. Stor. Nat. Chile, p. 301.
Northern Provinces of Chile.
Synonym: chinchilla, Meyen, 1833, Nova Acta, XVI, 2, p. 586. Chile.
brevicaudata , Waterhouse, 1848, Mamm. II, p. 241. Bolivia.
velligera, Prell, 1934, Zool. Anzeig. Leipzig, Bd. 108, p. 100.
Chile.
As is well known, these animals were nearly exterminated on account of the
value of their fur; now it appears they are being farmed, and it is hoped they
will be saved from extinction.
Genus 2. LAGIDIUM, Meyen
1833. LAcmiUM, Meyen, Nova .^cta Ak. Caes. Leop. Car. XVI, 2, p. 576.
1816. VISCACCIA, Oken, Lehrb. Nat. iii, 2, p. 835.
"The rulings of the International Zoological Nomenclature have reduced Viscaccia,
Oken to a synonym of the later described Lagidium" (Tate).
1833. Lagotis, Bennett, Proc. Zool. Soc. London, V, p. 58. {Lagotis cuvieri, Bennett).
Type Species. — Lagidium peruanum, Meyen.
Range. — Peru, Bolivia, Argentina, Chile; south in Argentine to 50° S., or
nearly to Magellan.
Number of Forms. — Twenty-one are named.
Characters. — Skull narrow, with long rostrum; a tendency present for the
frontals to be depressed between the orbits; the braincase
flat, round; no sagittal ridge formed. Bullae very- large, but not distorting the
occipital region of the skull as they do in Chinchilla, and not appearing much in
superior aspect of the skull. Paroccipital processes straight, joining the bullae.
Palatal foramina long and narrow; palate much constricted anteriorly. Jugal
very broad, in contact with lachrymal anteriorly, and with small upwardly
directed process on hinder upper border. Infraorbital foramen with no canal
for nerve transmission. Mandible near that of Chinchilla; the angular process
narrow, drawn backwards to a degree, the angular process not much distorted
outwards (older specimens seem more developed in this respect), the short
ridge beside the condyle very weak as a rule.
Cheekteeth each with three laminae, the laminae curved; the upper series
with the third plate of each tooth shorter than the other ones; M.3 with a
backwardly pointing heel. In the lower scries the front lobe of each tooth is
reduced (three laminae per tooth).
230 LAGIDIUM
Externally larger than Clunchilla; fur thick and soft; usually a black mid-
dorsal stripe present. Ear large. Hindfeet narrow, with four digits; D.5 much
as in Chinchilla, e.xtremely short. Claws weak and blunt. Forefeet with four
digits. Tail shorter than head and body, but of considerable length, and heavily
hairy.
Forms seen : arequipae, boxi. ciiscus, famatiniie, iiica, luti-um, locktvoodi,
moreni, pallipes, periiamim, pciliiteiim, puiiensis, surae, saturatits, suhrosea,
tuntalis, tuciimanum, riscaccia, vulcani, viatorum, zvoljfsohni.
Mr. R. W. Hayman has looked through the considerable British Museum
material with a view to getting the twenty-one "distinct species" in this genus
into some sort of revision. He reports as follows:
"There appear to be four species in this genus, two of them containing
eighteen of the twenty-one named forms recognized here.
"In Peru, extending southwards as far as the Bolivian and Chilian borders is
a group of small forms having the following features in common : smallish size,
dorsal stripe mostly absent or indefinite, rostrum short and teeth small. Peiii-
anum is the earliest name for this group.
" Southwards from North Bolivia to Chubut in the Argentine Andes is a
second group having the following in common : larger size, dorsal stripe usually
well marked and contrasting sharply with the usually greyish pelage, long
rostrum and large teeth.
" J'iscaccia is the earliest name in this group. Both this and the preceding
group have the hindfeet usually conspicuously paler than the body colour, and
in both the proportionate ear to head and body length may range from 17 per
cent to 21 per cent.
"In the southern Argentine Andes another group occurs, closely related in
skull characters to the riscaccia group, but characterized externally by very
short ears in proportion to large overall size, the percentage being from 13 to 15,
and by the hindfeet being uniformly coloured with the body. Boxi, sarae and
zvolffsohni belong here, the latter being the most southerly species of the genus.
" Except for the three last-named, all are here listed as subspecies of viscaccia
and peruanum. Although small skulls of viilcani, one of the northern races of
viscaccia, closely approach in proportions large skulls of inca, the most removed
geographically of the peruanum group, the skins are quite distinct.
"Actually where the two species approach each other geographically they are
most distinct (compare puneiisis and arequipae of South Peru with lutea, cuscus
and perlutea of North Bolivia).
I.
V.
riscaccia.
2.
V.
lutea.
3-
V
discus.
4-
V
perlutea.
5-
V
vidcani.
6.
V
tiicumana.
7-
V
lockuoodi.
8.
I-
famatinae.
LAGIDIUM 231
9. V. tontalis.
10. V. viatorum.
11. V. moreni. (5-1 1 inclusive doubtfully separable).
12. p. peruiinum.
13. /). piiHipi's. Possibly a synonym of 12.
14. />. /«a/.
IS- p- subrosea.
16. p. saturata.
17. /). pnnensis.
18. /). arequipae. 17 and 18 doubtfully distinct.
19. 6. AOATZ'.
20. b. sarae.
21. woljfsohni."
Note. — L. zvolffsolmi differs clearly in colour pattern from all the remainder.
List of Named Forms
1. L..\GIDIUM VISCACCIA VISCACCIA, Molina
1782. Storr. Nat. Chile, p. 307.
Chile.
Synonym: luiescetis, Philippi, 1S96, .Ann. Mus. Chile, 13, p. 8. Tacapuca,
Northern Chile.
cuvieri, Bennett, 1833, Proc. Zool. Soc. London, p. 58. Peru.
aureus, Geoffroy & D'Orbigny, 1830, Ann. Sci. Nat. XXL
p. 291. Corrientes, Buenos .Ayres.
criniger. Gay, 1847, Fauna Chile, i, p. 49. Chile.
crassidens, Phihppi, 1896, Ann. Mus. Nac. Chile, 13, p. 10.
chilemis, Oken, 1816, Lehrbuch Naturgesch. ii, p. 836,
Chile.
crimgerum, Philippi, 1896, Ann. Mus. Nac. Chile, 13, p. 10.
Chile.
viscaccica, Brandis, 1786, Versuch einer Naturgesch. von
Chile, p. 272.
2. LAGIDIUM VISCACCIA LUTEA, Thomas
1907. Ann. Mag. Nat. Hist. 7, XIX, p. 443.
Esperanza, Mount Sajania, Bolivia.
3. LAGIDIUM VISCACCIA CUSCUS, Thomas
1907. Ann. Mag. Nat. Hist. 7, XIX, p. 443.
Paratani, Bolivia.
4. L.\GIDIUM VISC.A.CCIA PERLUTEA, Thomas
1907. .Ann. Mag. Nat. Hist. 7, XIX, p. 443.
Pampa .Aulliaga, Bolivia.
5. LAGIDIUM VISCACCI.A VULCA.NI, Thomas
1919. Ann. Mag. Nat. Hist. 9, IV, p. 133.
Cerro Casabindo, Jujuy, .Argentina.
6. L.AGIDIUM VISC.ACCI.A TUCUMANA, Thomas
1907. -Ann. Mag. Nat. Hist. 7, XIX, p. 444.
Cumbre de Mala-Mala, Sierra de Tucuman, .Argentina.
7. LAGIDIUM VISCACCIA LOCKWOODI, Thomas
1919. Ann. Mag. Nat. Hist. 9, III, p. 499.
Otro Cerro, Rioja, .Argentina.
232 LAGIDIUM
S, LAGIDIUM VISCACCIA KAMATINAK, Thomas
1920. Ann. Mag. Nat. Hist. 9, VI, p. 421.
La In\'ernada, Rioja, Argentina.
Q. LAGIDIUM VISCACCIA TONTALIS, Thomas
1921. Ann. Mag. Nat. Hist. 9, VHI, p. 219.
Los Sombreros, Sierra Tontal, west of San Juan, Argentina.
10. LAGIDIUM VISCACCIA VIATORUM, Thomas
1 92 1. Ann. Mag. Nat. Hist. 9, VI H, p. 220.
Punta dc Vacas, Mendoza, Argentina.
11. LAGIDIUM VISCACCIA MORLNI, Thomas
1897. Ann. Mag. Nat. Hist. 6, XIX, p. 467.
Hills near Chubut, Argentina.
12. LAGIDIUM PERUANUM PERUANUM, Meyen
1833. Nova Acta Ac. Nat. Cur. XVI, p. 57S.
Southern Peru.
13. LAGIDIUM PERUANUM PALLIPES, Bennett
1835. Proc. Zool. Soc. London, p. 67.
(Believed to be) Chilian Andes.
14. LAGIDIUM PERUANUM INCA, Thomas
1907. Ann. Mag. Nat. Hist. 7, XIX, p. 442.
Incapirca, Zezioro, Junin, Peru.
15. LAGIDIUM PERUANUM SUBROSEA, Thomas
1907. Ann. Mag. Nat. Hist. 7, XIX, p. 442.
Galera, west of Oroya, Dept. Lima, Peru,
lb. LAGIDIUM PERUANT'M SATURATA, Thomas
1907. Ann. Mag. Nat. Hist. 7, XIX, p. 442.
Limbane, Inambari, Dept. of Puno, Peru.
17. LAGIDIUM PERUANUM PUNENSIS, Thomas
1907. Ann. Mag. Nat. Hist. 7, XIX, p. 443.
Puno, Lake Titicaca, Peru.
18. LAGIDIUM PERUANUM AREQUIPAE, Thomas
1907. Ann. Mag. Nat. Hist. 7, XIX, p. 442.
Sumbay, near Arequipa, Peru.
19. LAGIDIUM BOXI BOXI, Thomas
1921. Ann. Mag. Nat. Hist. 9, VII, p. 179.
Pilcaneu, Rio Negro, Argentina.
20. LAGIDIUM BOXI SARAE, Thomas & St. Leger
1926. .\nn. Mag. Nat. Hist. 9, XVIII, p. 639.
Pino Hachado, Neuquen, Argentina.
21. LAGIDIUM WOLFFSOHNI, Thomas
1907. .Ann. Mag. Nat. Hist. 7, XIX, p. 440.
Sierra de los Bagualcs y de las Vizcachas, 50 50' S., 72 20' \V., on
boundary between Chile and Argentina.
The Lagostomus Group
Differing chiefly from the Chinchilla group in the bullae, whicli are not
greatly inflated, the paroccipital processes, whicli are lengthened and stand apart
from the bullae, the extremely sharp claws of the feet, the complete suppression
of D.^ on the hindtoot, the skull more heavily ridged for muscle attachment.
LAGOSTOMUS
233
Fig. 68. Lagostomus maximus m-wimus, Desmarest.
B.M. No. 17.5.2.18; X t-
Genus 3. LAGOSTOMUS, Brooks
1828. L.\GOSTOMUS, Brooks, Trans. Linn. See. XVI, p. 96.
1824. VizcACi.A, Schinz. Naturg. und Abbild. Sijugeth. p. 243. (This name is not to be
used as it is a homonym of Viscaccia, Okcn, (Tate).)
Type Species. — Lagostomus tricltodactylus, Brooks=£>ipui maximus, Des-
marest.
Range. — Argentina. One form, from Peru, is probably extinct.
234
LAGOSTOMUS
Number of Forms. — Four.
Characters. — Skull flat, with broad frontals, which bear quite well-marked
postorbital processes. Nasals relatively long and narrow.
Fig. 69. Lagostomus maximus m.\.ximus, Desmarest.
B.M. No. 17. 5. 2. 18; ■, J.
A well-marked sagittal crest present. Paroccipital processes lengthened (probably
about as much as in T/irvoiiomvs); bullae not much inflated compared with the
Chinchilla group, though appearing to a certain degree behind, each side of
occipital region. A prominent canal present in infraorbital foramen for nerve
Fig. 70. Lagostomls maximus maximus, Desmarest.
B.M. No. 17.5.2. 18; X t-
Fig. 71. Lacostoml's m.wimis maximus, Desmarest.
Cheekteeth: B.M. No. 17.5.2.18; X 2}.
236 LAGOSTOMUS
transmission. Palate strongly constricted anteriorly. Jugal slanting sharply
upwards anteriorly, and in contact with lachrymal. As in other members of the
family, the upper zygomatic root is placed far backwards, over the middle of
hinder part of the toothrow. Mandible with rather strong ridge beside condyle
for attachment of masseter medialis; this, however, much shorter than in
Caviidae. Coronoid low; condylar process relatively low; angular process
distorted outwards rather weakly.
Cheekteeth set at an angle, the upper series with their outer edge pointing
forw'ards; all upper and lower teeth with two laminae only except M.3 upper
series, which is the largest tooth in the series and has three laminae. Incisors
medium in width, their surfaces covered with faint longitudinal grooves.
According to Pocock the penis differs considerably from that of other Hystricoid
genera examined bv him, including ChinchiUa; see also note on breadth of
manubrium on p. 171, which indicates yet another wide distinction from the
Chinchillae.
Externally relatively large (up to 470 mm. head and body in London
collection); fur less soft than in Chinchilla group; tail not long, fully haired;
forefoot with four digits armed with stout claws; hindfoot rather long, with
three digits only, the claws in their development comparable to those of Cuiii-
culits, excessively thick and sharp. Stiff bristle-hairs present on D.3, which is
the longest digit; D.2 is shorter than D.4.
Forms seen : crassus, inunoUis, inaximus.
List of Named Forms
1. LAGOSTOMUS M.AXIMUS MAXIMUS, Desmarest
1S17. Nouv. Diet. d'Hist. Nat. xiii, p. 117.
Argentina {?). (Localirj- unknown.)
Synonym: tricliodactyhis, Brooks, Trans. Linn. Soc. XVI, p. gfi, 1828.
.■\rgentina.
diana, Griffith in Cuvier, 1827, Anim. Kingd. III. p. 170.
viscaccia, GeofFroy & D'Orbigny, 1830, .\nn. Sei. Nat. xxi,
p. 291.
aiiiiger. Lesson, 1842, Nouv. Tabl. Regne. Anim. p. 105.
poniparum, Schinz, 1825 (1S24) Naturg. und Abbild.
Saugeth. p. 244.
americana, Sehinz, 1825, Cuviers Thierreich, IV, p. 429.
2. LAGOSTOMl'S AL-VXIMUS IMMOLLIS, Thomas
1910. Ann. Mag. Nat. Hist. ser. 8, V, p. 245.
Tapia, Tucuman, Argentina.
3. LAGOSTOMUS M.'iXIMUS PETILIDENS, Holl.ster
1914. Proc. Biol. Soc. Washington XXVII, p. 58.
8 miles south of Camien de Patagones, Southern Argentina.
4. LAGOSTOMUS CRASSUS, Thomas. (Extinct?)
1910. Ann. Mag. Nat. Hist. ser. 8, V, p. 246.
Santa .^na, C'uzcn, Peru. (Known from cranial characters only.)
CAVIOIDAE: CAVIIDAE 237
CIIINCHILLIDAE:
SPECIAL WORKS OF REFERENCE
Waterhouse, 1848, Natural Historj' Mammalia, Rodentia.
Tate, 1935, Taxonomy of Neotropical Hystricoid Rodents, Bull. Amer. Mus. Nat. Hist.
LXVIII, p. 295.
PococK, Proc. Zool. Soc. London, 1922, p. 365. External Characters of some Hystri-
comorph Rodents.
TuLLBERC, Nova -Acta Reg. Soc. Sci. Upsaliensis, XVIII, 3, no. i, 1899.
The family is known fossil from the IMiocene, from the Neotropical region
only. Miller & Gidley quoted several extinct genera.
Superfamily CAVIOIDAE
1896. Thomas: Hystrromorpha, part.
1899. TuUberg: Hystricogn.^thi, Hystricomorpha, part.
1918 Miller & Gidley: Superfamily Hystricoid.\e, part, medialis series.
1924. Winge: Family Hystricidae, part.
1928. Weber: Hystricoidea, part.
This Superfamily is equal to the medialis series of the Hystricoidae of Miller
& Gidley, and contains one family, the Caviidae, containing the genera and
subgenera Cavia, Galea, Caviella, Monticavia, Nanocavia, Kerodon, Dolichotis,
Paradolichotis and Hydrochoerus only.
I have elsewhere, when dealing with the Hystricoidae (Hystricoidae,
p. 97) remarked on the desirability of removing the Caviidae from the tj'pical
Hystricoid series, on account of the different formation of the lower jaw.
Apart from this structure and the formation of the cheekteeth they appear to
agree in essential characteristics with the Hystricoidae; but although Chin-
chillidae may show a certain resemblance in mandible formation to Caviidae, I
am unable to regard the Caviidae as typical Hystricoidae now, whatever their
ancestors may have been.
FamUy CAVIIDAE
1896. Thomas: Hystricomorph.a : Family Caviidae.
1899. TuUberg: Hystricomorpha : Family Caviidae, part, included Dasyprocta and
Cuniculus ( ' ' Coelogenys ").
1918. Miller iSc Gidley: Hvstricoid.\e (Medialis series). Family Caviidae; and Family
Hydrochoeridae (Hydrochoerus and fossil allies).
1924. Winge: Family Hystricidae, Dasyproctini, part, group Caviae.
1928. Weber: Hystricoidea: Family Caviidae, part, subfamilies Caviinae {Cavia), and
Hydrochoerinae (Hydrochoerus, Dolichotis).
Geographical Distribution. — The greater part or the whole of South
America; extending north to Panama.
Number of Genera. — Six.
Ch.\racters. — Zygomasseteric structure differing from that of the Hystri-
coidae in the formation of the lower jaw, which has the
angular process drawn backwards but not distorted outwards, and possesses a
deep horizontal ridge, for the insertion of masseter medialis (according to
23S CAVIIDAE: CAVIINAE
Miller & tiidley) present on side of mandible sliglitiy below alveolar level and
extending from the level of the condylar process to about as far as the hinder
part of M.I. Infraorbital foramen very large, as in Hystricoidae, and zygomatic
plate narrow, remaining completely beneath it.
Dental formula i. J, c. ;!, p. r, ni. if = 20. Cheekteeth evergrowing, unilat-
erally hypsodont, normally comparatively simplified in structure, but with
sharp folds and angular projections, the general effect more or less prismatic.
'Fibia and fibula not fully fused.
Clavicles suppressed. External form ambulatory or cursorial; digits of
hindfoot reduced to three. Tail obsolete. Lachrymal large; part of lachrymal
canal open on side of the rostrum, except in the genus Dolicliotis.
Two well-marked subfamilies may be recognized, which are sometimes
considered as families; but which, notwithstanding the high specializations of
the Hydrochoerinae, appear to agree in very many essential features.
Key to the Subeamilies of the Cavud.^e
M.3 not greatly enlarged; pattern of cheekteeth comparatively simple;
palate short to extremely short (from before backwards); parocci-
pital processes not abnormallv elongated. Subfamily Caviinae
[Caria, Galea, Caviella, Kirodon ; Dolicliotis)
M.3 extremely enlarged (upper series); pattern of cheekteeth compara-
tively complex; palate not short (from before backwards); parocci-
pital processes abnormallv lengthened. Subfamily Hvhrochoerinae
(Hydroclioerus)
Subfamily CAVIINAE
Geographical Distribution. — As in the family, except not known from
Panama.
Number of Genera. — Five.
Remarks. — The Caviinae fall into two well-marked groups, the Cavia
group, smaller genera with short limbs, shorter ears, and
moderate claws (or in Kerodon blunt nails), and the Dolicliotis group, containing
a single genus, with larger size, long limbs, long cars, sharp hoof-like claws, the
external form more modified for cursorial life.
Doliclujtis seems to be too nearly allied to the Caviae for these groups to be
regarded as subfamilies, as has been done (Pocock, Tate).
The Cavia group was revised by Osgood in 1915 (Field. Mus. Nat. Hist.
Publ. Zool. ser. X, no. 13, pp. 194, 195), who rightly restricted the genus
Kerodon to the species nipestris, and proposed the subgenera Galea and Caviella
for those species of Cavia with more simple cheekteeth. Thomas in 1916
treated Galea and Caviella as full genera, and erected Monticaria for the species
niata (referred by Osgood to Caviella). Later Thomas erected Naiiocavia for
a new species shiptoni, allied to Caviella and Monticavia.
The Caviinae have recentlv been revised at some length by Kraglievitch,
193 1, Ann. Mus. Nac. Buenos Aires, XXXVL p. 77. He divides the subfamily
CAVIINAE 239
Caviinae into the two groups here recognized. The genera Cavia, Galea,
Caviella, Monticavia, and Kerodun are retained in the Cavia group; Nanocavia
he reduced to a subgenus of Monticavia.
On this account I retain Galea and Caviella as full genera, though I am
bound to say that I feel convinced that it would be wiser to retain Osgood's
original classification, in which these two groups are regarded as of subgeneric
value only, as the characters which separate them from each other are of very
doubtful value.
But Monticavia is so closely allied to Caviella that I cannot treat it as more
than a subgenus. The main differences between Monticavia and Caviella are
that the heel in M.3 of Monticavia is less sharply defined, and that the incisors
are in Monticavia more pro-odont. But Kraglievitch gives the measurement of
the angle of the incisors of Caviella as between 88° and 1 10°, and that of Monti-
cavia (Xanocavia) shiptoni as 1 11°, so that the difference in this respect appears
to amount to one degree between the two "genera," which is hardly sufficient
to base a generic name on !
Kerodon is a distinct genus, which cannot be confused with any of the other
members of the Cavia group, whatever their status may be.
Key to the Groups of the Caviin.\e
The limbs shorter; ears short; claws not hooflike, less broadened, or may
be blunt; nasals not narrowed and pointed anteriorly; interorbital
region narrower; paroccipital processes less lengthened.
CAVL^i Group (Caviae)
{Cavia, Galea, Caviella; Kerodon)
The limbs longer; ears long; claws powerful and hoof like; nasals markedly
narrowed and pointed anteriorlv; interorbital region very broad;
paroccipital processes more lengthened.
Dolichotis Group (Dolichotides)
{Dolichotis)
The Cavia Group
Characters as indicated in the above key. Size medium or small, not
becoming large.
In all the genera, the jugal is broad but rather short, and zygoma not angular;
incisors relatively short, narrow; palate extremely constricted anteriorlv, the
premolars almost touching; upper cheekteeth much higher on inner side than
outer side, the lower cheekteeth much higher on outer side than inner side.
Mandible with coronoid process obsolete; condylar process of medium
height; angular process drawn far backwards, but not distorted outwards.
Beside and below the condylar process and extending forwards about to level
of hinder part of M.i is an extremely deep and prominent ridge for insertion
of masseter medialis.
Key to the Ge.nera of the Cavia Group
Claws blunt. Sternum narrow and rounded (Osgood). Kerodon
240 CAVIA
Claws sharp. Sternum broad and Hat (Osgood).
Posterior lobe of upper cheekteeth with a clear and deep outer re-
entrant fold; dental pattern less simplified. Cavi.^
Posterior lobe of upper cheekteeth with no re-entrant fold; dental
pattern more simplified.
Orbital branch of maxillae completely interrupted by the lachrymal ;
incisors pigmented; skull not bowed. G.alea
Orbital branch of maxillae not completely interrupted by the
lachrymal; incisors not pigmented; skull bowed to a
greater or lesser degree. Caviella
The ditferences between Galea and Caviella are based on characters which
are in other groups very variable; for instance, in Dolichotis the interruption
of the orbital branch of the maxillae by the lachrymal may be present or absent.
The incisors may or may not be pigmented within many genera elsewhere in
the Order, for instance, Ctenoiiivs, Xenis (Geosciunis), and others. The orbit
is more circular in Caviella than in Galea.
Genus i. CAVIA, Pallas
1766. Cavia, Pallas, Misc. Zool. p. 30.
Type Species. — Cavia cubava, Pallas = Miis porcellus, Linnaeus.
Range. — South America; Brazil, the Guianas, Venezuela, Colombia, Peru,
Bolivia south to Northern Argentina.
Number of Forms. — Approximately seventeen.
Characters. — Skull with some interorbital constriction apparent, and a
sagittal crest developed in the adult. Infraorbital foramen
broader below than above; a canal present for transmission of nerve. Bullae
relatively large. Paroccipital processes noticeably elongated. Palate short, ex-
tending about to front of M.3. Palatal foramina short, narrow. Jugal medium,
not approaching the lachrymal. Incisors not pigmented.
Upper cheekteeth divided into two lobes by inner re-entrant fold in the
upper series, the hinder lobe larger than the anterior one, and with a deeply
indenting fold in its outer border. M.3 with posterior projection. Lower
cheekteeth with one deep outer fold dividing tooth into two lobes and with an
inner fold in the posterior lobe. Mandible as already described.
Externally the limbs not specially elongated, the hindfeet long, with three
digits, the central digit the longest; the claws sharp. Forefeet with four digits,
D.3 the longest, D.5 the shortest; D.4 rather longer than D.2. Ears relatively
short.
This genus is quite well differentiated from Galea and Caviella by the more
complex cheekteeth.
Forms seen : anolaiinae, apereti, azarae, festiiui, fulgiiia, niiianae, nana, pain-
parnm, pallida, porcellus, rosida, stolida, tsclnidii, umhrata.
For notes on the species of Cavia see I'homas, Ann. Mag. Nat. Hist. 8,
XIX, p. 152, 1917.
CAVIA 241
List of Named Forms
(References and type localities of all named forms for the Caviidac are the
work of I\Ir. G. W. C. Holt.)
1. CAVIA GUIANAK, Thomas
1901. Ann. Mac. Nat. Hist. 7, VIII, p. 152.
Kanuku Mountains, British Guiana.
2. CAVIA VENKZUELAIC, Allen
191 1. Bull. Amer. Mus. Nat. Hist. XXX, p. 250.
Altagracia, Immataca district, Venezuela.
Considered by Thomas as doubtfully distinguishable from guianae.
3. CAVIA .\PI;REA APEREA, Erxleben
1777. Syst. Regn. .Anim. i, p. 348.
Brazil.
Synonym: leiicopyga, Brandt, 1835, M^m. Acad. St. Petersb. 6, iii,
p. 436. Brazil.
4. CAVIA APEREA AZ-iVR-^E, Lichtenstein
1823. Doublet. Zool. Mus. Berlin, p. 3.
Ypamena, Province Sao Paulo, Brazil.
5. CAVIA ROSIDA, Thomas
1917. .Ann. Mag. Nat. Hist. 8, XIX, p. 154.
Roca Nova, East Parana, Brazil.
6. CAVIA PAMPARUM, Thomas
1901. .Ann. Mag. Nat. Hist. 7, VIII, p. 539.
Goya, Corrientes, Argentina.
7. CAVIA TSCHUDII TSCHUDII, Fitzinger
1867. Sitz.-Ber. K. Akad. Wien (Math. Nat.), LVI, p. 154.
Cit>' of Yea, 70 miles east of Pisco, Western Peru.
8. CAVIA TSCHUDII UMBRATA, Thomas
1917. .Ann. Mag. Nat. Hist. 8, XIX, p. 157.
Incapirca, Zezioro, Central Peru.
9. CAVIA TSCHUDII AREQUIPAE, Osgood
1919. Journ. Mamm. Baltimore, p. 34.
.Arequipa, Peru.
Synonym: tschudii pallidior, Thomas, 1917, Ann. Mag. Nat. Hist, 8,
XIX, p. 158. 'Sox. (niata) pallidior, Thomas. The name
arequipae was proposed in case Monticavia was regarded
as not distinguishable generically from Cavia. Perhaps
pallidior should stand in the present work, as niata
pallidior is regarded as a Caiietla.
10. CAVIA TSCHUDII STOI.IDA, Thomas
1926. .Ann. Mag. Nat. Hist. 9, XVIII, p. 166.
Rio Utcubamba, 15 miles south of Chachapoyas, Peru.
11. CAVIA TSCHUDII FESTINA, Thomas
1927. Ann. Mag. Nat. Hist. 9, XX, p. 604.
Huariaca, Junin, Peru.
12. CAVIA TSCHUDII SODALIS, Thomas
1926. Ann. Mag. Nat. Hist. 9, XVII, p. 607.
Norco, 20 kilometres north-west of Vipos, Prov. Tucuman, .Argentina.
16 — Living Rodents — 1
242 CAVIA— GALEA
13. CAVIA TSCHUDII ATAHUALPAE, Osgood
1Q13. Field Mus. Nat. Hist. Publ. Zool. ser. X, no. 13, p. gS.
Cajamarca, Peru.
14. CAVIA NANA. Thomas
191 7. Ann. Mag. Nat. Hist. !S, XIX, p. 158.
Bolivian Highlands (Chulumani, Yungas).
15. CAVIA FULGIDA, Waglcr
1 83 1. Isis, XXIV, p. 512.
.Amazonia.
Synonym: rufescens, Lund, 1841, Afh. K. Danske. Vid. Selsk. 4, VIII,
p. 284. Lagoa Santa, Brazil.
nigricans, Wagner, 1841, Schreber, Saug. Suppl. IV, p. 64.
Brazil.
16. CAVIA .ANOLAIMAE, Allen
1916. Bull. .\mer. Mus. Nat. Hist. XXXV, p. 85.
Anolainia, west of Bogota, Colombia; on a branch of River Bogota.
17. CAVIA PORCELLUS, Linnaeus. (Domestic)
1758. Syst. Nat. loth ed., i, p. 59.
Brazil.
Synonym: aperoides, Lund, Blik. Dyr. pi. 25. Brazil.
robiista, Lund, 1841, Blik. Dyr. pi. 25, fig. 16.
brasilieiisis, Linnaeus, 1754, Mus. Adolphi. Friederici, p. 9.
gracilis, Lund, 1841, Blik. Dyr. pi. 25.
ciitleri, Bennett, 1835, Proc. Zool. Soc. London p. 191.
Lima, Peru.
cobaya, Marcgrave, 1648, Hist. Nat. Bras. p. 224. Peru.
longipilis, P'itzinger, 1879, Sitz.-Ber. K. Akad. Wien
(Math. Nat.), LXXX, Ab. i , p. 43 1 . Japan.
Genus 2. GALEA, Meyen
1833. Galea, Meyen, Nova Acta Ak. Caes. Leop. XVI, 2, p. 597.
Type Species. — Galea musteloides, Meyen.
Range. — Bolivia, North Argentina, Chile and Brazil.
Number of Forms. — Ten.
Characters. — Like Cavia but cheekteeth simpler, each upper tooth cut
into two lobes by one inner fold; M.^ with weak backwardly
projecting heel. Lower teeth two-lohed; P. 4 with short anterior prolongation.
Orbital branch of maxillary completely interrupted by lachrymal. Incisors
pigmented.
Forms seen : aiiceps, bolivieiisis, copies, demissa, flavidiis, littoialis, iiegremis,
palustris, spixii.
List of Named Forms
I. GALEA MUSTELOIDES MUSTELOIDES, I\Teyen
1833. Nova Acta Ak. Caes. Leop. XVI, 2, p. 598.
Pass of Tacara and Tajori, Andes, North-west Bolivia.
Synonym: bolizicnsis, Waterhouse, 1848, Nat. Hist. Mamm. 11, p. 175.
Bolivia, highlands between Cochabamba and La Paz.
comes, Thomas, 1019, .•\nn. Mag. Nat. Hist. 9, IV, p. 134.
Maimara, Jujuy, Argentina.
GALEA— CAVIELLA 243
2. GALEA MUSTKLOIDIiS Li;UCOBLKPH.\RA, Burmeister
1861. Reise durch La Plata, H, p. 425.
Mendoza to Tucuman, Argentina.
3. GALEA MUSTELOIDES LITTORALIS, Thomas
igoi. Ann. Mag. Nat. Hist. 7, VU, p. 195.
Bahia Blanca, Argentina.
Synonym: miisteloides negrensis, Thomas, 19 19, Ann. Mag. Nat. Hist.
9, HL p. 211. Pilcaneu, Upper Rio Negro, .-Argentina.
4. GALEA MUSTELOIDES DEMISSA, Thomas
1921. Ann. Mag. Nat. Hist. 9, VIII, p. 623.
San Antonio, Parapiti, Bohvia.
5. GALEA MUSTELOIDES AUCEPS, Thomas
1911. Ann. Mag. Nat. Hist. 8, VIII, p. 255.
Guarina, Lake Titicaca, Bolivia.
6. GALEA MINIMUS, Molina
1782. Sagg. Stor. Nat. Chili, ist ed., p. 306.
Chile.
Considered a subspecies of miisteloides by Tate ; if this is so, the name
antedates miisteloides, and all races must be regarded as races of
minimus.
7. GALEA SPIXII, Wagler
1831. Isis, XXIV, p. 512.
Brazil.
Synonym: saxatilis, Lund, 1841, Afh. K. Danske Vid. Selsk. 4, VIII,
p. 286. Lagoa Santa, Brazil.
8. GALEA WELLSI, Osgood
1915. Field Mus. Nat. Hist. Publ. Zool. ser. X, no. 13, p. 196.
Sao Marcello, Bahia, Brazil.
0. GALEA PALUSTRIS, Thomas
191 1. Ann. Mag. Nat. Hist. 8, VII, p. 608.
Cameta, lower Rio Tocantins, Brazil.
10. GALEA FLAVIDENS, Brandt
1835. M^m. Acad. St. Petersb. p. 439.
Brazil.
Synonym: obscurus, Lichtenstein, 1823, Doublet. Z. Mus. Berlin, p. 3.
Brazil.
bilobidens, Lund, 1841, Afh. K. Danske Vid. Selsk. 4, VIII,
p. 286. Brazil.
Genus 3. CAVIELL.A, Osgood
1915. C.^vlELL.^, Osgood, Field. Mus. Nat. Hist. Publ. Zool. ser. X, no. 13, p. 194.
Regarded by Kraglievitch as indistinguishable from Microcavia, Genais and
Ameghino, 1880, .Mamm. Foss. Anier. Sud. p. 50, a fossil genus.
1916. MoNTiCAViA, Thomas, Ann. Mag. Nat. Hist. 8, XVIII, p. 303. Cavia niata,
Thomas. Valid as a subgenus.
1925. Nanocavia, Thomas, Ann. Mag. Nat. Hist. 9, XV, p. 41S. Manocavia shiptoni,
Thomas. Valid as a subgenus.
244 CAVIELLA
Type Speciks. — Cavia amtralis, Geoffroy & D'OrhiKiiy.
Range. — Bolivia and Argentina, south to Patagonia.
Number of Forms. — Eight.
Characters. — Si\ull with rostrum slanting downwards anteriorly, more
bowed than in allies, the highest part of the skull usually
about over posterior zygomatic root. Palatal foramina larger than in preceding
Fig. 72. Caviella australis joannia, Thomas.
B.M. No. 71. 12. 29. 12, 9; X 2.
genera, triangular, placed more closely to toothrows. Sagittal crest present in
old age. Bullae relatively larger, and orbit more circular than in Curia and
Galea. Incisors without pigment. Cheekteeth like Galea, but usually I\1.3 with
deeper posterior fold.
Monticai'ia, here regarded as a subgenus of Caviella, has more pro-odont
incisors, the angle with the line of toothrow about 1 1 s . M.3 is less complicated,
the heel a short projection, without internal notch. Skull more bowed anteriorly.
Nafiocavia, as remarked above, is intermediate between typical Caviella
and Monticavia in the angle of the incisors; the bullae are considerably smaller
than in either, the portion appearing on occipital surface of skull practically
uninflated.
Fig. 73. Caviella australis joannia, Thomas.
B.M. No. 71. 12. 29. 12, ?; X 2.
Fig. 74. C.-\vi£LL.\ australis joannia, Thomas.
Cheekteeth: B.M. No. 71. 12.29. 12, ?; x 6.
246 CAVIELLA— KERODON
Forms seen: aiistnilis, juaiinia, maenas, niata, " iiigiuinti," pa/lidiur, salinia,
shiptoni.
List of Named Forms
Subgenus Cai-ie/hi, Osgood
1. CAVIELLA AUSTRALIS AUSTRALIS, Gcoffroy S: D'Orbigny
1833. Mag. Zool. I. pi. 12.
Rio Negro, Patagonia.
Synonym: australis nigriana, Thomas, i(j2i, Ann. Mag. Nat. Hist. 9, VII,
p. 446. Neuquen, Rio Negro, Argentina.
2. CAVIELLA AUSTR.ALIS KINGII, Bennett
1835. Proc. Zool. Soc. London, p. iqo.
Port Desire, Patagonia.
3. CAVIELLA AUSTRALIS JOANNIA, Thomas
1921. Ann. Mag. Nat. Hist. 9, VII, p. 446.
Canada Honda, San Juan, Argentina.
4. C.WIELLA AUSTRALIS MAENAS, Thomas
1898. Ann. Mag. Nat. Hist. 7, I, p. 284.
Chilecito, Rioja, Argentina.
5. CAVIELL.A AUSTRALIS SALINIA, Thomas
1921. Ann. Mag. Nat. Hist. 9, VII, p. 447.
South-east Catamarca, .Argentina.
Subgenus Nanocavia, Thomas
6. CAVIELLA SHIPTONI, Thomas
1925. Ann. Mag. Nat. Hist. 9, XV, p. 419.
Laguna Blanca, Catamarca, Argentina.
Subgenus Monticavia, Thomas
7. CAVIELLA NIATA NIATA, Thomas
1898. Ann. Mag. Nat. Hist. 7, I, p. 282.
Espcranza, 50 km. from Mt. Sahama, Bolivia.
8. CAVIELLA NIATA PALLIDIOR, Thomas
1902. Ann. Mag. Nat. Hist. 7, IX, p. 229.
Pampa .Aullaga, Lake Poopo, Bolivia.
Genus 4. KERODON, F. Cuvier
1825. I'iERODON, F. Cuvier, Dents, des Mamm. p. 151.
Type .Species. — Cavia nipcstiis, Wied.
Range. — Brazil. (British Museum specimens froin Bahia.)
Number of Forms. — One.
Characters. — Much like Curia cranially. Sagittal ridge feeble or absent.
Infraorbital foramen with no canal for nerve transmission.
KERODON— DOLICHOTIS 247
Palatal foramina excessively narrowed. Rostrum relatively narrow. Bullae
moderate; paroccipital processes rather long. Upper cheekteeth two-lobed;
1VI.3 with a weak backwardly projecting heel. P.4 lower with a well-marked
e.xtra anterior projection; heel of M.3 (lower) poorly defined. Differing,
according to Osgood, in several details of the skeleton from Cavia and allies,
the chief of which is that the sternum in this genus is narrow and rounded
instead of broad and flat, the spinous processes of the lumbar vertebrae are
thick, heavy, and depressed, and the large neural spine of the axis fully overlaps
the first cervical.
Externally differing markedly from all allies in the fact that the digits are
armed only with blunt nails.
Remarks. — Whatever the status of Galea and Caviella compared with
Cavia, there is no doubt that on account even of the nails alone,
this genus is distinct from that group.
Forms seen : rupestris.
List of Named Forms
I. KERODON RUPESTRIS, Wied
1820. Isis, VI, p. 43.
Rio Grande de Belmont, Rio Pardo, etc., Brazil.
Synonym: moco, F. Cuvier, Dents, des Mamm. 1825, p. 151. Brazil.
sciureus, Geoffroy, 1826, Diet. Class. IX, p. 120. Brazil.
The Dolichotis Group
Becoming larger than the Cavia group; to very large (head and body 690,
or more?); hindlimbs lengthened, general form modified for cursorial life.
Hindfoot very long, with three digits bearing hooflike claws; arrangement of
digits perissodactyle. A rudimentary tail present. Forefoot artiodactyle, the
four digits armed with sharp claws. Frontals much broadened, and nasals
considerably specialized.
Genus 5. DOLICHOTIS, Desmarest
1819. Dolichotis, Desmarest, Joum. Phys. Paris, LXXXVIII, p. 211. (Cavia pata-
chonicha, Shaw.)
1927. WE-iT!NBERCiiiA, Kraglicvitch, Physis, VIII, p. 579. Subgenus for Dolichotis
saliniivla, Bumieister. Name preoccupied.
1927. P.ARADOLICHOTIS, Kraglievitch, Physis, VIII, p. 594. Dolichotis salinicola, Bur-
meister. Valid as a subgenus.
1927. Pediol.\cus, Marelli, Mem. Jardin Zool. la Plata, vol. Ill, p. 5. Dolichotis salini-
cola, Burmeister.
1928. Lagospedius, Marelli, Physis, IX, p. 103. Dolichotis salinicola, Burmeister.
Type Species. — Cavia patachonica, Shaw.
Range. — Patagonia and Argentina.
Number of Forms. — About five have been named. There appear to be
only two species.
Characters. — Nasals large, much pointed anteriorly, considerably excised
at the side on joining the maxillae in the typical subgenus.
24S DOLICHOTIS
Nasals not extending as tar forwards as the premaxillae. Frontals very broad,
the orbits roofed in by expansion of the frontal bone, which is deeply notched
anteriorly. Occiput relatively weak, sloping forwards; paroccipital processes
considerably elongated, much more than is normal, but not comparable to the
structure found in llvdrochoerus. Bullae moderately large. Palate very short,
extending only to about level of ]\1.2; toothrows nearly meeting anteriorly.
Palatal foramina long and narrow. Jugal broad, short; often a small upwardly
directed process on posterior border. Mandible as normally in Caviidae, the
masseteric ridge sometimes less deep than in the Cavia group. Lachrymal very
large, but apparently the canal is practically or completely closed in front of the
orbit. Cheekteeth evergrowing, unilaterally hypsodont as in Cavia group.
Upper cheekteeth each two-lobed, except M.3, which is cut into three lobes by
two re-entrant folds. Lower cheekteeth v^'ith one outer fold cutting the teeth
into two lobes; P. 4 with an extra anterior prolongation.
No separate canal for nerve transmission in the infraorbital foramen.
Ears long; essential external characters as described above.
Paradollcliotis is proposed as a subgenus for the smaller species salinicola
(head and body to about 460 in few skins seen), differing in the lower anterior
prolongation of the nasals being rudimentary or absent, and in several parts of
the skeleton (there are no skeletons of this species in London). For further
details see Kraglievitch, 1931, Anales Museo Nac. Buenos Ayres, xxxvi, p. 77.
•According to Pocock the penis of DoUchotis differs considerably from that of
members of the Cavia group.
Forms seen : magellanica, centricola, salinicola.
It appears that magellanica and patachonicha may be regarded as synonyms
of the oldest name patagoiiuiir, I have never seen any notes in which these forms
have been compared or regarded as distinct.
List of Named Forms
Subgenus DoUchotis, Desmarest
1. DOLICHOTIS P.ATAGONA PATAGON.A, Zimmermann
17S0. Geogr. Gesch. 11, p. 32S.
Patagonia.
.Synonym: 0) patachonica, Shaw, iSoi, Genl. Zoology, 2. i, p. 226.
(?) magellanica, Kerr, 1792, .Anim. Kingd. p. 220. Magellan.
2. DOLICHOTIS PATAGON.A CENTRICOLA. Thomas
1902. Ann. Mag. Nat. Hist. 7, IX, p. 242.
Cruz del Eje, Central Cordova, Argentina.
Subgenus Paradolicliotis, Kraglievitch
3. DOLICHOTIS SALINICOLA, Burmeister
1875. Proc. Zool. .Soc. London, p. 635.
Stations Totoralejo and Recreo, Central Argentine Railway. 29° S.,
6s" W.
Synonym: 0) centralis, Weyenbergh, 1877, Versk. Ak. .Amsterdam,
XI, p. 247. Cordova, Argentina. Status fide Thomas,
Trouessart.
HYDROCHOERINAE
249
Subfamily HYDROCHOERINAE
Geographical Distribution. — The warmer parts of South America, north
to Panama.
Number of Genera. — One.
Characters. — Cheekteeth more complex than in the Caviinae; M.3 enor-
mously enlarged, exceeding the combined length of the three
anterior teeth in size; paroccipital processes extremely elongated, ven,' much
more so than in any other Rodent; bodily size largest in the Order; (habits
semi-aquatic).
Fig. 75. Hydrochoeris hvdrochaeris hydrochaeris, Linnaeus.
B.M. No. 27.2.1 1.112, 9; X j.
250 HYDROCHOERUS
Genus i. HYDROCHOERUS, Brisson
1762. HYDROCHOERUS, Brisson, Regn. Anim. 2d Ed. p. 12.
Type Species. — Siis hydrochaeris, Linnaeus.
R!\^;q]7_ — As in the subfamily. Forms named from Brazil, Paraguay, and
Panama. Specimens in British Museum from British Guiana.
Fig. 76. HYDROCHOERUS HYDROCHAERIS HYDROCHAERIS, Linnaeus.
B.M. No. 27. 2. II. 112, 9; y i.
Said to occur in Venezuela, and also to extend to Peru and Bolivia; but the
exact range of this genus has not been traced.
Number of Forms. — Three.
HYDROCHOERUS
251
Characters. — Skull heavy, rather flat; nasals broad; frontals broad and
long; occiput relatively narrow, and evidently a sagittal ridge
is not formed. Lachrymal large, with part of lachrymal canal open on side of
rostrum. Bullae proportionately smaller than in Caviidae; palate much longer,
extending back to hinder part of 1\1.3; palatal foramina large; palate constricted
anteriorly; pterygoid fossae very deep; infraorbital foramen without canal for
nerve transmission. Paroccipital processes abnormally elongated. Mandible
Fig. 77. Hydrochoerus hy-droch.\eris hydrochaeris, Linnaeus.
B.M. No. 27.2.11.112, ?; X J.
typically Cavioid in formation, the masseter medialis ridge moderately to strongly
developed. Jugal broad, zygoma heavy.
Incisors broad, faintly one-grooved. Cheekteeth remarkable for the amount
of cement present. In the upper series, P. 4, M.i, and M.2 are each divided into
two lobes, the lobes united by cement, and each lobe with a further deep outer
fold; the lobes narrowed mternallv and pointing forwards. M.3 with nine or
ten narrow transverse plates joined to each other, and to an anterior and a
posterior lobe, the anterior lobe like those of the other molars, the posterior
252
HYDROCHOERUS
lobe consisting of two transverse plates joined externally. In the lower teeth,
P. 4 and M.i are each divided into three lobes; in the premolar each lobe has an
inner fold; in IM.i the two anterior lobes are with one inner fold, the posterior
lobe with an outer told. AI.2 and M.3 have each four lobes, the central two of
which are simple transverse plates, the anterior with an inner fold, the posterior
Fig. 78. Hydrochoerus hydrochaeris hy'droch.\eris, Linnaeus.
Cheekteeth: B.M. No. 27.2.1 I.I 12, $; < li.
with an outer fold. i\1.3 is the dominant tooth of the series, but is very much
less enlarged than M.3 in the upper series.
External form very large, easily the largest member of the Order, though
not of course comparing with "very large" forms of other Orders. Head
broad, ears short; fur harsh; limbs not greatly lengthened. Forefoot perisso-
dactyle; D.3 the longest digit; D.5 considerably shorter than the others.
HYDROCHOERUS— APLODONTOIDAE 253
Hindfoot perissodactyle, with three digits only, the digits webbed, but the
webbing poorly developed. Claws hea\T, thick. Tail rudimentary.
The largest specimen seen is 1 175 mm. head and body, but I should imagine
that this does not represent the extreme development for the genus.
Contrary to the opinion of some I have discussed the subject with, my very
limited experience with these animals in captivity, at the London Zoological
Gardens, indicates that they possess considerable intelligence; more so indeed
than in any Rodent 1 have ever tried to establish contact with.
Forms seen: hydrochaeris.
List of N.^med Forms
1. HYDROCHOERUS HYDROCH.^ERIS HYDROCHAERIS, Linnaeus
1766. Syst. Nat. 12th Ed. p. 103.
Brazil.
Synonym: capybara, Erxleben, 1777, Syst. Regn. Anim. p. 193.
2. HYDROCHOERUS HYDROCHAERIS NOTIALIS, Hollister
1914. Proc. Biol. Soc. Washington XXVII, p. 58.
Paraguay.
3. HYDROCHOERUS ISTHMIUS, Goldman
1912. Smiths. Misc. Coll. LX, no. 2, p. 11.
Marraganti, Rio Tuyra, Eastern Panama.
Remarks. — Notwithstanding the extreme specializations of this genus, such
as the lengthened paroccipital processes, lengthening of AL3, and
enormous bodily size, I think that there are far too many essential characteristics
shared between it and the Caviidae for Hydrochoerus to be referred to a separate
family, as has recently been advocated (Pocock, Miller & Gidley).
The Caviinae and Ilydrochoerinae are both known from the Miocene from
South America. The Hydrochoerinae are also known from the Pleistocene of
the South-eastern L'nited States.
CAVIIDAE:
SPECIAL WORKS OF REFERENCE
PococK, 1922, P.Z.S. p. 365. External characters of some Hystricomorph Rodents.
Waterhol'SE, 1848, Natural History Mammalia, vol. II (Rodentia).
Tate, 193';. Ta.\onomv of Neotropical Hystricoid Rodents, Bull. Amer. Mas. Nat.
Hist. LXVIII, 2, p. 295.
Tl'llberc, 1899, Nova Acta Reg. Soc. Sci. Upsaliensis, XVIII, 3, i.
Osgood, 1915, Field. Mus. Nat. Hist. Publ. Zool. ser. X, no. 13, p. 195.
Kraglievitch, An. Mus. Nac. Buenos .■Vires, XXXVI, 193 1, page 59.
Superfamily APLODONTOIDAE
As here understood this contains one living family.
Family APLODONTIIDAE
1896. Thomas: Aplodonti.\e : Family .Aplodontiidae.
1899. Tullberg: Sciuromorpha: Sriuroidei, part; Family Aplodontiidae.
254 ArLODONTlIDAE
igi8. Miller CV' Gidle\': Siipcrfamily Dipodoidae, part; Family Aplodontiidae.
iy24. Winge: Haplodontidae, part: Haplodontini.
ig2X. Weber: Haplodontoidea: Family Haplodontidae.
Geographical Distribution. — Nearctic; Western North America; the
Pacific side of the Rocky Mountains, from
California to extreme Southern British Columbia.
Number of Genera. — One.
Characters. — Zygomasseteric structure of a type differing from any found
elsewhere in the Order. Masseter lateralis superficialis
attached anteriorly to zygoma; but infraorbital foramen not transmitting muscle;
zygomatic plate very narrow, completely below the infraorbital foramen;
mandible with angular portion not distorted outwards by specialized portion of
lateralis muscle, but with its outer border sharply pulled inwards.
Skull flattened, greatly widened posteriorly. Bullae with neck directed
horizontally outwards. Cheekteeth evergrowing, simplified in pattern, the
dental formula i. ], c. ",, p. f, m. § = 22. Fibula not reduced nor fully fused
with the tibia, at any rate as compared with Murine or Dipodoid genera.
External form more or less modified for fossorial life.
Remarks. — The Aplodontiidae were included by Miller & Gidley in their
Superfamily Dipodoidae; but the genus is evidently very far
removed from other members of that group as understood by these authors.
The main difference as regards jaw-muscles is that in this case the infraorbital
foramen does not transmit muscle, or scarcely does so; (according to Taylor it
definitely does not do so; Coues states that it does transmit a small strand, and
from skulls examined it appears that it could do so; as figured by Tullberg, and
as stated by Winge, it does not; but the difference between this type of jaw-
muscle structure, even if it does transmit a small strand, is very widely different
from that present in such specialized families as Dipodidae, Pedetidae, and
Anomaluridae, with which this family is associated by Miller & Gidley). It
should be noted that the main difference between the Hystricoidae and the
Bathyergoidae of Miller & Gidlev is that in the latter the infraorbital foramen
does not transmit muscle, and in the former it does so; so that it would appear
that if the classification of Miller & Gidley were retained, this family should be
separated from "Dipodoidae" if only on the grounds ot consistency.
Coues, Tullberg, and other writers have come to the conclusion that
Aplodoiitia is a Sciuroid; Tullberg places the genus as a family together with the
Sciuridae in his section Sciuroidei, a section of Sciuromorpha equal in impor-
tance to his Castoroidei, and Geomyoidei. But once again it appears that the
zygomasseteric structure of Aplodontia is widely different from either Sciuridae,
Castoridae, or Geomyidae, all of which have evolved a more specialized arrange-
ment of the forepart of the skull for attachment of masseter muscles in that
the zygomatic plate is broadened, to a highly specialized degree in all but a
few genera of the Pteromys group, and even in these the difference between
Aplodontia and such primitive forms as Belomys is already quite well estab-
lished. In these families, masseter lateralis rises up the broadened zygomatic
APLODONTIIDAE: APLODONTIA 255
plate to the superior border of the rostrum, and masseter laterahs superficiahs
has become distinct from the zygoma; but in Aplodotitia this is not the case;
so that as far as zygomasseteric structure is concerned it appears that Aplodontia
is not to be considered a near ally of either Sciuridae, Castoridae, or Geomyidae.
The extraordinary inflection of the angular portion of the mandible is so
far as I have seen without parallel in the Order, though certain Sciuridae, as
Cynumys, and certain Dipodidae, as I believe Cardiocranitis, and certainly to a
degree Zapiis, approach it.
Apart from this, it would seem that Aplodontia stands nearest what one might
consider the primitive or ancestral type of zygomasseteric structure of Rodentia;
this apparently is the theory of Winge, who derives directly or indirectly all
families of Rodentia as here understood from his family "Haplodontidae."
'I'he jaw muscles, though arranged in a different manner from most members
of the Order, are according to Tullberg very strongly developed. The tem-
poralis muscles are strong and extensive.
The familv contains one living genus.
Genus i. APLODONTIA, Richardson
1829. Aplodontia, Richardson, Zool. Joum., vol. 4, p. 334.
Type Species. — Aplodontia leporina, Richardson = ^nwowy^; rufa,
Rafinesque.
Range. — As in the family. A good map of the range is published in
Anthony, Field Book of North American Mammals, 1928, p. 455.
Number of Forms. — Nine.
Characters. — As in the superfamily. The skull is abnormally broadened
posteriorly, and considerably so anteriorly; behind the
anterior zygomatic root the frontals are abruptly and considerably narrowed.
The parietal ridges are well marked, but not fused in the few skulls examined.
The zygomata are widely spreading. The skull is much flattened. The width
of the occipital region is about equal to two-thirds of the length of the skull.
The auditory bullae are flask-shaped, with the neck directed horizontally out-
wards. The posterior zygomatic root is noticeably at right angles to the supra-
occipital. Incisive foramina not large, situated far in front of the palate, which
is broad, and extends behind the toothrows.
-Mandible with the outer side of the angular process pulled inwards to an
abnormal degree; to such an extent that the posterior border is horizontal, and
the two edges of this process form the base of a triangle which has for its apex
the condylar process. This is perhaps best expressed by noting that if the two
halves ot the mandible are separated each halt may be made to stand up on a
table resting on the angular portion. Coronoid process verv high and curved
backwards.
As noted above, the infraorbital foramen appears sufficiently enlarged to
transmit a very small strand of muscle; in appearance it is round; but it is very
2s6
APLODONTIA
small comparfd with any Rodent in which the infraorbital may be said definitely
to transmit muscle, and according to all authors I have read on the point with
the exception of Coues it does not do so.
Incisors powerful.
Cheekteeth evergrowing, i ; P. 3 minute, probably functionless, the pattern
of the other teeth in the adult nearly simplified to a ring, the inner side of those
of the upper series circular, the outer side of each tooth with an externally
pointing projection on either side of which is a slight depression.
Fig. 79. .^PLODONTI.A ri'f.a rvfa. Rafinesquc.
B.M. No. 98.9.28.1, V; ■■- 1.
Lower teeth like those of the upper series, but with the pattern reversed,
and with tendency for a small re-entrant external angle to be present, this
wearing out with age.
Form thickset, heavy ; fur thick and soft ; limbs short ; eyes and ears small ;
tail more or less vestigial, much shorter than hindfoot. Claws, particularly
those of the forefoot, enlarged and powerful. Forefoot with all digits present,
but pollex very short. D.3 the longest digit, then D.4, next D.2, last D.5.
Hindfoot with the three centre digits roughly equal, the hallux and D.5 shorter
than these. Mammae 0 (Taylor).
APLODONTIA
257
Fig. 8o. Aplodontia rlfa rlfa, Rafinesque.
B.M. No. 98.9.28.1, 0; X I.
Fig. 81. Aplodontia rufa rufa, Rafinesque.
Mandible from below, X i; Cheekteeth X 4; B.M. No. 98.9.28.1, 5.
The species of Aplodontia are described as burrowing, mainly nocturnal
animals, living in colonies in dense wet forests, in which they construct numerous
tunnels, and feed on bark, leaves, and twigs. Thev are said not to hibernate.
The family Aplodontiidae is known from the Miocene of Western North
America. A closely related genus is described fossil from Eastern Asia.
Though this work is concerned with living Rodents, mention mav be made
17 — Living K^hIciUs — I
258 APLODONTIA
of an interesting fossil family from North America which appear to have had a
similar zygomasseteric structure to the Aplodontiidae, and are referred to that
family by Winge, namclv the Mylagaulidae. They appear to have developed
along lines noticeably diflerent trom any living group ot Rodents, and I feel
that no general work on Rodentia would be complete without reference to them.
"Family Mylagaulidae
"Skull excessively fossorial, occipital region obliquely truncate, with lambdoid
crest moved forward nearly to level ot zygomatic root; frontal with short post-
orbital processes; bonv horn-cores on rostrum in two genera, absent in a third;
cheekteeth highly modified from normal heptamerous structure, the grinding
function of toothrow in adult almost completely taken over by the greatly
enlarged fourth premolar. General structure of skull much as in the Aplodon-
tiidae; cheekteeth i or I, a reduced heptamerous pattern evident in slightly
worn crowns, but this giving place with wear to a system of narrow longitudinal
and oblique lakes. Molars relatively small, soon crowded out by the premolar,
an excessively hypsodont laterally compressed tooth closed at base and rapidly
increasing in crown length from the unworn surface downwards. Skeleton
highly modified for imderground life.
"Mylagaidiis, Ceidtogauliis, and Epigaulus, North American Miocene and
Pliocene" (Miller & Gidley).
APLODONTIIDAE:
SPECIAL WORKS OF REFERENCE
T.AYLOB, Univ. Calif. Publ. Zool., XVII, pp. 435-504, 1918. A full revision of the
genus.
CouES, Monograph of North American Rodents, p. 540, 1S77. Monograph of genus.
Tt'LLBERG, Nova Acta Reg. Soc. Sci. Upsaliensis, XVIII, 3rd ser., no. i, 1899.
Forms examined: ;;//(/, pacifica, olviiipicti, "major."
List of N.\med Forms
Nine forms are now recognized, all regarded as races of the type species.
Revised by Taylor, 1918, Univ. Calif. Publ. Zool., XVII, p. 435. The
references and type localities are the work of Mr. R. W. Hayman.
1. .APLODONTIA RUFA RUFA, Rafinesque
1S17. Arner. Monthly Mag. 2, p. 45.
Columbia River, Oregon.
Synonyms: leporina, Richardson, 1S29, Zool. Jouni. 4, p. 335.
grisea, Taylor, 1916, Lniv. Calif. Pub. Zool. XII, p. 497.
(Near Seattle, Washington.)
cliryseola. Kellogg, 1914, L'niv. Calif. Publ. Zool. XII, p. 295.
(Jackson Lake, California.)
2. APLODONTIA RUFA OLYMPICA, Merriam
1899. Proc. Biol. Soc. Washington, XIII, p. 20.
Quinault Lake, Chehalis County, Washington.
APLODONTIA— SCIURIDAE 259
3. APLODONTIA RUFA COLUMBIANA, Taylor
1916. Univ. Calif. Publ. Zool. XII, p. 499.
Vicinity of Hope, liritish Columbia.
4. APLODONTIA RUFA RAINIKRI, Mcrriam
1899. Proc. Biol. Soc. Washington XIII, p. 21.
Paradise Creek, south side Mount Rainier, Washington.
5. .APLODONTIA RUFA PACIKICA, Mcrriam
1899. Proc. Biol. Soc. Washington, XIII, p. ig.
Newport, Yaquina Bay, Lincoln County, Oregon.
6. APLODONTIA RUFA HUMBOLDTIANA, Taylor
1916. Proc. Biol. Soc. Washington XXIX. p. 21.
Carlotta, Humboldt County, California.
7. APLODONTIA RUFA CALIFORNICA, Peters
1864. Monatsber. k. Preuss. Akad. Wiss. Berlin, p. 179.
(Assumed to be) Sierra Nevada, California.
Synonym: major, Merriam, 1886, Science, 7, p. 219, .\nn. New York
Ac. Sci., HI, pp. 312, 316. (California, Sierra Nevada
Mountains.)
8. APLODONTIA RUFA NIGRA, Taylor
1914. Univ. Calif. Publ. Zool. XII, p. 297.
Point Arena, Mendocino County, California.
9. APLODONTIA RUFA PHAEA, Merriam
1899. Proc. Biol. Soc. Washington, XIII, p. 20.
Point Reyes, Marin County, California.
Superfamily SCIUROIDAE
Thi.s as here understood contains one family only.
Family SCIURIDAE
1896. Thomas: Sciuromorpha: Family Sciuridae (with subfamilies Sciurinae and
Nannosciurinae).
1899. Tullberg: Sciuromorph.\ : Sciuroidei (part, included Aplodontiidae) : Family
Sciuridae.
1918. Miller & Gidley: Superfamily Sciuroidae, part: Family Sciuridae, with sub-
families Sciurinae, Nannosciurinae and Pteromyinae.
1924. Winge: Family Sciuridae, part; Sciurini.
1928. Weber: Sciuroidea: Families Sciuridae, Pteromyidae, Xeridae, Tamiidae,
Marmotidae.
Geographical Distribution. — Practically cosmopolitan; absent only from
the Australasian region, Madagascar, and
Southern South America (Patagonra, Chile, most of Argentina). Also absent
from certain desert regions, as Arabia, and Egj'pt, etc.
Number of Genera. — I have retained forty-four genera.
Characters. — Zygomasseteric structure as follows: infraorbital foramen not
transmitting muscle, or scarcely so; masseter lateralis super-
ficialis with anterior head distinct from zygoma; zygomatic plate broadened,
26o SCIURIDAE
tilted upwards, forming base for masseter lateralis to rise obliquely to superior
border of rostrum, which it does to the exclusion of masseter medialis. Mandible
never highly modified; usually with the angular portion pulled inward to a
degree, as in Muscardinidae, Dipodidae, Aplodontiidae, etc.
I o 2 I ^
Dental formula i. -, c. -, p. - or -, m. - = 20 or 22.
I o ' I 1 3
Cheekteeth rooted, brachyodont or hvpsodont, their pattern usually charac-
terized by prominent cusps and ridges, the lower series most often basin-shaped,
the pattern as a rule complex. Tibia and fibula not fully fused. Jugal long,
usually in contact with the lachrymal. Skull with well-marked postorbital
processes, which may in rare cases be poorly de\'eloped (for instance Mvosciiiriis).
Tail always fully haired, usually thick and bushy. External form suited to
arboreal or terrestrial life. Digits of hindfoot five.
This is one of the largest and most widely distributed groups of living
Rodents, and possesses about thirteen hundred named forms.
Craxi.\l Characters. — The following cranial characters are general in
the group.
There are, as indicated above, always postorbital processes present. There
is very rarelv any strongly marked interorbital constriction present, though this
becomes noticeable in some members of the Pteromvs group. The jugal is long,
usually in contact with the lachrymal. The bullae are rarely excessively in-
flated, but as a rule evenly roimded and of relatively large size. The palate is
broad as a rule, and usuallv terminates slightly behind the toothrows, or about on
a level with the hinder molars. In the Xenis section it is much produced back-
wards. The incisive foramina are always situated considerably in front of the
toothrows, and are as a rule ver\- short. The incisors are usually laterally com-
pressed in the arboreal genera (with some exceptions), and less or not so in the
terrestrial forms.
Dental Characters. — The cheekteeth of all the genera seem referable to
a single original pattern, though varying consider-
ably in the various genera. In Larisciis and Rhiitlirosciurus they are almost sim-
plified. The original plan appears to be, in the upper series, that of four trans-
verse ridges, a more or less marked cusp marking the outer border of each of
these ; the ridges run across to the inner border of the tooth, w'hich is normally
formed by one large inner cusp, except in some primitive Flying-squirrels, in
which the three inner cusps are retained; each of these transverse ridges has
between them a depression; the second and third ridges are the highest; the
first and fourth mark respectively the anterior and posterior terminations of the
tooth. The lower cheekteeth are normally with a large central basin-shaped
depression, often tending to take up most of the tooth, and surrounded by cusps,
of which there is usually a main one at each corner, though sometimes the
posterointernal cusp may be hardlv developed. The anterointernal cusp is in
almost every case the highest one.
If Winge's theory', that the infraorbital foramen of this family once trans-
mitted muscle and has become secondarilv closed to muscle transmission, is
SCIURIDAE 261
correct, we have in this family and others which share its zygomasseteric struc-
ture (Castoridae, Heteroinyidac, Geomyidae), one of the most highly specialized
arrangement of jaw muscles known in the Order. But primitive genera like
Belomys appear to be so (comparativelv) near the Aplodontiidae in zygomatic
plate and infraorbital foramen formation that I very much doubt whether this
is so, and think it more probable that the arrangement has been derived from
something like the Aplodontoid type of zygomasseteric structure.
All forms belonging to this family, including the Flying-squirrels, which have
often been referred to a distinct family, appear to me to be so obviously closely
related, and so obviously offshoots of one essential stem, that I can recognize
no subfamilies. I have provisionally divided the family into two groups, the
Flying-squirrels and the Xon-flying-squirrels, though when dealing with the
Anomaluridae I have endeavoured to point out that a flying-membrane is not
a structure on which subfamilies should be based ; this membrane may have been
developed independently in the present group on more than one occasion, and
I must admit that this di\ision into groups is largely for convenience owing to
the very large size of the family.
Key to the Groups of Sciuridae
A flying-membrane present attached to sides. Cheekteeth often but not
always with tendency towards abnormal complexity. Zygomatic
plate often but not always low, weak, and little tilted upwards.
Pteromys Group (Pteromyes)
(Belomys, Trogopterus, Pteromysais, Petaurista, Aeromys,
Pteromys, Glaucomys, Eoglaucomys, Hylopetes, Petinomys,
Petaurillus, loiins, Eupetaunis.)
No flying-membrane present. Cheekteeth never with tendency towards
abnormal complexity. Zygomatic plate often but not always
strongly tilted upwards. SciURUS Group (SciURl)
(Myoschirtis, \tinnosciurus, Sciurillus, Microscinriis, Svntheo-
sciunis, Sciurus, Tumiasciunts, (Jallosciunis, Funiimbtiliis, Dremo-
mys, Ratiifa, Menetes, Luriscus, (ilyphotes, Rheithrosciurus,
RJiinosciurus, Hyosciurus, lltliosciwus, Paraxerus, Funisciurus,
Prutoxenis, Mvrsihis, Epixerus, Xerits, Atlantoxeriis, Spermo-
pliilopsis, Sciurotamias, Tamias, Citellus, Marmota, Cynomys.)
Outline of Previous Classification of F.uiily Sciuridae
1891. Flower & Lydekker, Mammals Living and Extinct.
Ten living genera were recognized in the Family Sciuridae, which was
divided into the subfamilies Sciurinae and Arctomyinae ( = Marmotinae), the
latter "so intimately connected with the. preceding subfamily that the division
into two groups is purelv a matter of convenience."
The genera were :
I . Sciurus. (All Tree-squirrels except number 2.)
262 SCIURIDAE
2. Rlieithrosciurus.
3. Xenis.
4. Tamias.
5. Pteromys (=Petaurista).
6. Sciuropterus (—-Pteromys).
7. Eupftauiiis.
8. Arctomvs {^Marmota).
9. Cynomys.
I o . Spermophihis ( = Citellus) .
Two years later Forsyth Major, published his paper on the dentition of the
Sciuridae (1893, Proc. Zool. Soc. London, p. 179) whicli has formed the basis
of most modern classifications of the family. He recognized three subfamilies,
the Sciurinae (including Marmotinae of Flower & Lydekker), the Pteromyinae,
(Flying-squirrels), and the Nannosciurinae containing the Old World Pvgmy-
squirrels. He recognized ten genera as in Flower & Lydekker except that
Nannosciurus was raised to generic rank, and that Tamias was suppressed and
regarded as a subgenus of Sciurus. Several of the species, forming the genus
Sciurus of former classifications, were referred, on account of dental characters,
to the genus Xerus. His arrangement was as follows :
Sciurinae
1 . Rlieiihrosciiiius
2. Xenis
Subgenus Proioxenis (=the modern Protoxeius, Epi.xenis and
Myrsiliis)
Subgenus Xenis
Subgenus Atlautoxenis
Subgenus Paiaxenis (=the modern Paraxenis and Funisciunis)
Subgenus " Eoxenis" (=the modern Fuuambulus, Menetes,
Lan'scus, and Rliinosciurus)
3. Sciurus
Subgenus " Eosciurus" (=Ratufa)
Subgenus Sciurus. American, Palaearctic, and African forms
currently referred to Sciurus, Callosciurus, Heliosciurus.
Subgenus Tamias
4. Spermopliilus ( = CiteUus)
5. Arctomys {^Marmota)
6. Cynomvs
Pteromyinae
7. Sciuropterus (=the modern Pteromys and related genera)
8. Pteromys (—Petaurista)
9. Eupeiaurus
Nannosciurinae
10. Xauuosciurus (=the modern Nannosciurus and Myosciurus)
SCIURIDAE 263
This arrangement was followed by Thomas in 1896 in his classification of
the whole Order except that the Pteromyinae were not regarded as forming a
distinct subfamily, being referred to the Sciurinae, and Tamias was again given
generic rank.
In 1897 (Proc. Zool. Soc. London, p. 933) Thomas proposed that all sub-
genera of Major's classification except Atlantoxerus should be given generic
rank, substituting the name Fimambulus for Eoxenis, and Ratufa for Eosciurus.
In 1908 (Journ. Bombay N. H. Soc, XVIII, 2, p. 244) Thomas gave a revised
list of Asiatic genera of non-flying Squirrels, recognizing altogether twelve.
1. Sciurus. (The group subsequently referred to Callosciunis and
" Tomeutes.")
2. Zetis, which had formerly been referred to Funambiilus, but was
separated in the paper now under discussion; the name is ante-
dated bv Dremomys, Heude.
3. Glyphotes (erected by Thomas, 1898).
4. Ratiifa.
5. Tamiops, which had been erected by Allen for Sciurus maclellandi.
6. Rhinoscitiriis, which had been separated from Fimambulus by Miller.
7. Menetes, which was separated from Funambiilus by Thomas in the
paper now under discussion.
8. Fimambulus. (Restricted to the forms now referred to it.)
9. Larisais. ("Lana," Gray, preoccupied.) (Formerly had been referred
to Funambiilus.)
10. Rlieithrosciurits.
1 1 . Sciurotamias, which had been erected by Miller for Sciurus davidianus.
12. Nannosciiiriis.
This classification is retained in the present work except that I am unable
to regard Tamiops as a genus distinct from Callosciunis.
In the same year (Ann. Mag. Xat. Hist., 8, I, p. i) Thomas revised the
Flying-squirrels, recognizing eight genera :
1. Peiaiirista.
2. Eupetaiiriis.
3. Trogopteriis, which had been previously erected by Heude.
4. lomys.
5. Belomys.
6. Pteromysciis.
7. Petaurillus.
8. " Sciuroptcnis" (=Pteromys), with subgenera Glaucomys, Hylopetes,
and Petinomys; all these subgenera have subsequently been given
generic rank and appear to me to be clearly distinct from Pteromys
as now restricted (Scandinavian, Russian, Siberian, and Japanese
small Flying-squirrels), but more doubtfully so from each other.
The following year (Ann. Mag. Nat. Hist., 8, III, p. 467) Thomas revised
the African genera of Sciuridae, recognizing twelve genera :
264 SCIURIDAE
1. Sciuius. (The group subsequently referred by Thomas to Aetho-
sciurus, and shown by Hollister, Bull. U.S. Nat. Mus., 99, p. 9,
1919, to be not distinijuishable as a full genus from Heliosciurus.)
2. Heliosciurus.
3. Myrsihis. (Separated from Protoxerus.)
4. Paiaxerus.
5. Funisciurus . (Separated from Paiaxerus.)
6. Protoxerus.
7. Epixerus. (Separated from Protoxerus.)
8. Atlantoxerus.
9. Xerus.
' ^ ' ■ ' (Both separated from Xerus.)
11. Geosciurus. 1 ^ ^ '
12. Mvosciurus.
This classification is followed in the present work except that Euxerus and
Geosciurus are regarded as subgenera of Xerus only; Aet/iosciurus, following
Hollister, is referred to Heliosciurus; and I think that with representative material
it is likely that both Myrsilus and Epixerus (here retained) would be better referred
to Protoxerus.
In 1912, Miller (Catalogue of Mammals of Western Europe) regarded the
Flying-squirrels as forming a distinct family, the Petauristidae (the sole char-
acter being the presence of the flving-membrane, p. 940), In 191S in Miller
& Gidley (Classification of Rodentia) he ver)' properly reduced the group to the
rank of subfamily.
In 1915, J. A. Allen (Bull. Amer. Mus. Nat. Hist., XXXIV, p. 147) restricted
the genus Sciurus to the Palaearctic, and divided the Squirrels occurring in
America into no less than seventeen genera. These names, based mostly on
mammary formula (4 or 6) and the relative length of the rostrum, have for the
most part been disregarded, and appear to be based for the most part on specific
groups. His "genera," with remarks on subsequent treatment, are listed
below :
1. Tainiasciurus. Retained by Miller, 1923 (List of North American
Recent Mammals) as a subgenus of Sciurus. Given generic rank
by Pocock (Proc. Zool. Soc. London, p. 237, 1923) on account of
the suppression of the baculum. Retained as a full genus by
Howell. 1938 (North. Amer. Fauna, 56, p. i) in his classification
of genera of North American Sciuridae.
2. Neosciurus. Regarded as a synonym of Sciurus, subgenus Sciurus by
Miller, 192';. Revived as a subgenus of Sciurus by Howell, 1938
(including " Buiosciurus" and " Ecliiriosciurus").
3. Otosciurus. Regarded as a synonym ot Sciurus, subgenus Sciurus by
Miller, 1923. Revived as a subgenus of Sciurus by Howell, 1938.
4. Hesperosciurus. Regarded as a synonym of Sciurus, subgenus Sciurus
bv Miller, 1923. Revived as a subgenus of Sciurus by Howell,
1938-
SCIURIDAE 265
5. Echinosciurus . Regarded as a synonym of Sciuriis, subgenus Sciurus
by Miller, 1923. Regarded as a synonym of Sciurus, subgenus
.Xeosciurus by Howell, 1938.
6. Baiosciiirus. Regarded as a valid subgenus of 5a«r«j by Miller, 1923.
Regarded as a synonym of Sciurus, subgenus Xeosciurus bv Howell,
1938-
7. Parasciurus. Regarded as a synonym of Sciurus, subgenus Guer-
linguetus by Miller, 1923. Revived as a valid subgenus by Howell,
1938.
8. Syntheosciurus. Currently retained as a full genus.
9. Microsciurus. Currently retained as a full genus.
10. Sciurillus (Thomas). Currently retained as a full genus. Transferred
to the subfamily Nannosciurinae by Thomas, and by Miller &
Gidley, 1918. (In the present paper it has been thought de-
sirable to include in this genus certain Squirrels from Celebes
(murinus group), which as far as examined agree in cranial
characters with this genus.)
1 1 . Leptosciurus . Regarded as a subgenus of Sciurus by Thomas (Ann.
Mag. Nat. Hist. 10, H, p. 290, 1928) (as all Neotropical "genera"
of .^llen).
12. A'otosciurus. Remarks as Leptosciurus.
13. Mesosciurus. Regarded as a synonym of Sciurus, subgenus Guer-
linguetus by Miller, 1923, and by Howell, 1938.
14. Guerlinguetus. Regarded as a valid subgenus of Sciurus by Miller,
1923, and bv Howell, 1938. (But in a wider sense than accepted
by Allen.)
15. Hadrosciurus. Remarks as Leptosciurus.
16. Lrosciurus. Regarded as indistinguishable from Sciurus, subgenus
Hadrosciurus bv Thomas, 1928 (Ann. Mag. Nat. Hist. 10, H,
p. 290, 1928). Shown by Lonnberg, 1921, to be not retainable
on cranial characters suggested by Allen.
17. Simosciurus. Regarded as not distinguishable on cranial characters
from either Hadrosciurus or Lrosciurus bv Lonnberg, 1921.
(.Author's note : but dentition normal, noticeably different from
Hadrosciurus and Lrosciurus. Here regarded as a synonym of
Sciurus, subgenus Guerlinguetus.)
Miller and (jidley, 1918, in their classification of the Order Rodentia divided
the family into three subfamilies, the Sciurinae, Pteromyinae and Nannosciur-
inae, the latter based solely on cranial characters (but originally proposed by
Forsyth .Major on dental characters).
Miller (List of North American Recent Mammals, U.S. Nat. Mus. BuU.
128), 1923, listed twelve genera occurring north of Panama:
I. Marmota.
1. Otospermopliilus. (Had been separated since earlier classifications of
Thomas and Forysth .Major, from Citellus.)
266 SCIURIDAE
3. Callospermophihis. (Had been separated since earlier classifications
from Taviias.)
4. Cite I bis.
5. Ainmospeniiophilus . (Remarks as CaUospermopliilus.)
6. Cxnomxs, with subgenus Leucocrossuromys.
7. Eutamim. (Had been given generic rank by Merriam, separated from
Tamias.)
8. Tamitis.
9. Schirus, with subgenera Tamiasaiinis, Sciiinis, Baiosciurus, and
Giierlingiietiis.
10. Microsciurus.
1 1 . Sviitheosciuriis.
12. Ghnicomvs.
Howell, 1938, has made some modifications in this arrangement. Callo-
spermophiltis, AmnwspermopJiihis, and Otospermopliilus are referred to Cite/lus
as subgenera. In the present work, Eutamias is shown to be not a valid genus.
In 1915, Thomas introduced the system of dividing genericallv on the struc-
ture of the penis-bone or baculum, and a few genera have since been erected,
based on this character, alone. These genera are not retained in the present
work, for the following reasons. Out of numerous named forms, very few
appear to have been examined as regards this structure; those that have, have
been shown in some cases to vary in this character from subspecies to subspecies
(Osman Hill, 1936, Eiinambidiis). In other families of Rodents, no generic names
have been given to forms which vary in baculum characters; or at most subgeneric
names only (for instance, Dipodidae (Vinogradov), Cricctinae (Argyropulo)). If
these mammals are given subgenera onlv on this structure, which seems to me
to he scarcely necessarv (or at most of subgeneric value except in cases of total
suppression of the baculum), I fail to see why such names as Tomeutes in the
present family must be given full generic rank. It may also be argued that the
baculum refers to the male animal only. Pocock has suggested that there may be
corresponding modifications in the reproductive parts of the female, and suggests
that these might be worked out later. But the work on this whole problem is so
far from being finished that it seems absurd to recognize names based on the
shape of the baculum alone.
On the other hand, it is admitted that the genera, all currently accepted,
Callosciiinis, Fiinambuhis, Heliosciiirus, and Sciurus are not in all cases dis-
tinguishable from each other on cranial and dental characters. In cases like
these there are wide dift'erences between the few forms heretofore examined in
penial characters; Heliosciiirus is said to have the baculum suppressed; while
the other three are referred to as manv distinct subfamilies by Pocock. These
genera are here provisionally retained, partly on this character, partly on average
differences in cranial and dental characters, partly on account of the great
convenience of so doing, though some doubt is telt on the advisability of their
retention.
Pocock (Proc. Zool. Soc. London, 1923, pp. 209-246) classified the whole
family on characters of the baculum alone, with ears and feet used if the baculum
I
SCIURIDAE 267
had not been examined, but leaving cranial and dental characters out altogether,
liut if this character is given such importance, 1 fail to see how fossil forms are
to be considered; and it seems that if cranial and dental characters have been
used primarily for classification since the days of Linnaeus one cannot be blamed
for wishing to continue to give more importance to these characters than to an
external character which has only been definitely verified in a very small per-
centage of named species and races, and found to be subspecifically variable in
at least one case.
Pocock classified the family (not including the Flying-squirrels) as follows:
Subfamily Sciurinae
Sciurus, with subgenus Tenes for persiciis ; all the American genera or sub-
genera, Neosciurus, Parasciiirus, Echinosciunis, etc., except Tamiasciurus,
(?) Rheithrosciurus.
Subfamily Tamiasciurinae
Tamiasciurus. ("Penis . . . flexible throughout owing to the suppression
of the baculum" (compare Heliosciurus).)
Subfamily Funambulinae
("A highly diversified group of genera, with glans penis exceedingly variable
in size and structure, and baculum either relatively very large {Funambuliis,
Tamiodes), relatively small (Protoxerns, Ratufa), minute {Funisciurus, Paraxerus,
Aethosciurus), or absent (Heliosciurus).")
Funambulus, Tamiodes, Ratufa, Protoxerus, Aethosciurus, Funisciurus, Para-
xerus, Heliosciurus, and probably other African genera admitted by Thomas,
including possibly even Myosciurus.
Subfamily Callosciurinae
Callosciurus, Menetes, Tomeutes, Rhinosciurus, Lariscus, Dremomys, Tamiops,
Nannosciurus, probably others.
Subfamily Xerinae
Atlantoxerus, Xerus, Euxerus, Geosciurus.
Subfamily Marmotinae
Marmota, Marmotops, Cynomys, Citellus, with many subgenera. (?) Tamias,
Eutamias.
The subfamily Nannosciurinae was done away with by Thomas and Pocock
because, as might be expected, the baculum of Myosciurus differs from that of
Xannosciurus. The dental characters of the group diagnosed by Forsyth .Major
appear to be not strictly constant in all cases; and the cranial characters
diagnosed by Miller & Gidley appear to be not so distinct in all cases as was at
268 SCIURIDAE
first supposed; for instance, the genus Microsciiinis (Sciurinae) appears to be
rather transitionary towards Sciurillus (Nannosciurinae) which, in turn, connects
with Xaiiiiosciunis.
\\'inge, 1924 (Pattcdyr Slaegtcr, II, p. .S4), recognized nine genera only of
Sciuridae as here understood, but referred the Castoridae to the family.
Group Sciuri
1 . Tamias.
2. Otospeniiophilus.
3. Sciiirus.
4. Pleroinvs (with Petaurista).
5. Eupetaurus.
6. Xmis.
Group "Arctomyes"
7. Arctomys ( = Marmota).
8. Speniiop/iiliis (^Citelhis).
9. Cynomys.
Howell, 1938, has revised the genera and subgenera occurring in North
America north of Panama, and recognizes:
1 . Tiiiniiis.
2. Eiitdiniiis (subgenus Neotnmias).
3. Mannota (subgenus Maniiolops).
4. Cviioiins, with subgenus Leiicocrossiiromvs.
V Citt'lhis, with subgenera Cilellus {townsendii , washingtoni, richardsonii,
and ptinvi groups); Ictidomvs (tildeceinliiieatns and spi/osoma
groups); Otospermophilus; A'o/oaVf//i« (subgenus n. for aiDiulatus);
A mmospermophllus ; Callospermophihis ; Xerospeniiophiliis (tereti-
caudiis group); Poliocitelliis (subgenus n. ior frankliiui).
6. Glaucom\s.
7. Svntheosciurus.
8. Micyosciurus.
9. Sciiiins (typical subgenus restricted to Palaearctic). Subgenera
Xeosciiinis (with synonyms Baiosciurus and Echinosciunis); Hes-
perosciiinis (griseiis) ; Otoscitirus (dberti) ; Pdiasciunis ( witii synonym
Arue(isciiinis); Giierliiigiiettis (with synonym Mesoscinnis).
10. Tamiasciurus.
This arrangement is followed in the present paper except that Eutamias is
not considered a valid genus, and that Marmotops (based on the presence of a
functionless digit) is regarded as a synonym of Mannota.
In forming the key to the genera, I do not include the following three char-
acters which have frequently been used for generic purposes, but in my opinion
certainly should not be so.
(l) Presence or absence of functionless upper premolar (P. 3). This
feature has been pointed out to be a character of little mipurtance already by
SCIURIDAE 269
Hollister and Pocock. The tooth in question is either present or absent in
the genus Tamiasciurits, and may occasionally appear in typical Heliosciurus,
which was originally given generic rank on the sole character "cheekteeth
\" instead of ^. In any case except in certain Marmots the tooth appears
to have ceased to be of much functional importance.
(2) Colour pattern. Nearly all Squirrels with a Tamias-Viikt series of
longitudinal stripes on the back have received generic names. I do not
think that genera can be retained on this ground alone, unless coupled with
definite characters elsewhere. Examples are " Tamiscus" and " Tamiops."
Citellus tridecemlineatiis, for instance, has the most specialized colour pattern
known in the family, but many other species of CzYeZ/iw are uniformly coloured.
Also a striped colour pattern occurs in Funisciuriis (lemniscatus group)
side by side with species without it.
(3) Geographical distribution. I am not persuaded that because a
Squirrel comes from Africa it is of necessity distinct generically from one
that comes from the Malay region, or even from America.
Cl.\ssific.^tion Here Adopted
Thirteen genera of Flying-squirrels are here retained, and thirty-one genera
of non-flying Squirrels. In this family above all others generic names have been
bestowed freely, and for no apparent reason, again and again. It is not an easy
group, and it may be that the key I have endeavoured to get together will not
hold in all cases; also some of the genera are at the moment only separable on
average characters, or on characters of the baculum which may break down at a
later date when more forms have been examined. I can see no necessity of
retaining more than thirty-one genera of non-flying Squirrels, and would be
quite content personally to regard even several of those that have been retained
as of not more than subgeneric value.
I have divided the non-flying Squirrels into seven sections which may in
some cases be of doubtful value. These will be discussed later.
The Pteromys Group
I have made no changes in this group, the thirteen named genera being
all retained.
The Sciiiriis Group
A. Nannosciurus section (Pygmy Squirrels with abnormal cranial characters).
Genus i. Myosciurits, Thomas.
Sole species : M. pumilio.
Genus 2. Nannosciurus, Trouessart.
Principal species: .V. exilis group; A', whiteheadi
group; A', melanotis group.
Genus 3. Sciurilliis, Thomas.
Principal species: 5. piisillus group; S. murinus
group (Celebes).
270 SCIURIDAE
B. Sciurus section. (Typical Tree-squirrels; all genera except Ratiifa and
possibly Microsciitrus are not easily distinguishable from the genus Sciurus.)
Genus 4. Micrnsciurtis, Allen.
Principal species: M. alf art group.
Genus 5. Svntlteosciiiriis, Bangs.
Sole species: S. brochus.
Genus 6. Sciurus, Linnaeus.
Subgenus a. Sciurus, Linnaeus.
Principal species: S. vulgaris group (with lis).
Subgenus b. Tenes, Thomas.
Principal species: 5. anomalus group.
Subgenus c. Neosciurus, Trouessart. (Considered valid bv Howell,
^938.)
Principal species: S. carohnensts group; S. deppei
group; S. aureogaster group, with poliopus,
colUaci, socialis, griseoflavus, xucatanensis, varie-
giitoidcs, etc.
Subgenus d. Oiosciurus, Nelson. (Considered valid by Howell, 1938.)
Principal species: S. aberti group.
Subgenus e. Hcsperosciurus, Nelson. (Considered valid by Howell,
193S.).
Sole species : S. griseus.
Subgenus/. Parasciurus, Trouessart. (Considered valid by Howell,
. '938.)
Principal species: 5. mger group (oculatus, artzon-
aisis, etc.)
Subgenus g. Guerlinguetus, Gray.
Principal species : S. hojjmani group (with gerrardi,
etc.); S. aestuans group; 5. pucherani group
(Ignitus, boliviensis); S. siramiiieus group.
Subgenus It. Notosciurus, Allen.
Sole species: S. rhoadsi.
Subgenus /. Iladrosciurus, Allen.
Principal species: S. jlaitimifer; S. laiigsdorjfi group
(" Uroscitirus" as understood by Allen).
Genus 7. Tatuiasciurus, Trouessart.
Principal species: T. hudsfniicus group.
Genus 8. Callosciurus, Gray.
Subgenus a. Tamiops, Allen.
Principal species: C. niaclellaiidi group.
Subgenus b. Callosciurus, Gray.
Principal species: C. tenuis group, with jentinki;
C. lozvi group; C. erytliraeus group (with
sladeni, ferrugineus, finlaysoni, flaiiinanus, bo-
courti, gerinaini, griseiinanus, atrodorsalis); C.
caniceps group; C. prevosti group; C. notaius
SCIURIDAE 271
group (with vittatus, nigrovittatus); C. pygery-
ihrus group (with lokroides, phavrei); C. quin-
questriatus group; C. hippurus group (with
pryeri, brooki, melanogasler, philippinensis and
other species from PluHppines); C. leucomus
group ; C. rubriventer group.
Genus 9. i'unambtdus, Lesson.
Principal species: F. palmarum group, with pen-
nanti, tristriatus, wroughtoni ; F. layardi group;
F. siihlineatus group.
Genus 10. Dremomys, Heude.
Principal species: D. lokriah group; D. rufigenis
group; D. pernyi group, with {:)ou'stoni,
{})ez-eretti.
Genus II. Ratiifa, Gray.
Principal species: R. macroura; R. indica; R. bicolor;
R. gigantea; R. melanopepla; R. affinis; R.
ephipphim.
C. Lariscus section. (Not a natural group, hut containing genera from the
Indo-Malayan region, all of which are much specialized and clearly distinct
from Sciurus genericaliy.)
Genus 12. Menetes, Thomas.
Principal species : M. berdmorei and races.
Genus 13. Lariscus, Thomas & Wroughton.
Principal species: L. insignis group; L. hosei.
Genus 14. Glyphotes, Thomas.
Sole species: G. simus.
Genus 15. Rhdthrosciurus, Gray.
Sole species: R. macrotis.
Genus 16. Rhinoscitirus, Gray.
Sole species: R. laticaudatus.
Genus 17. Hyosciiirus, Tate & Archhold.
Sole species: H. heinrichi.
D. African arboreal genera. (All but Heliosciurus are clearly distinct genericallv
from Sciurus. Heliosciurus appears to lead into Paraxerus in cranial and
dental characters.)
Genus 18. Heliosciurus, Trouessart.
Subgenus a. Heliosciurus, Trouessart.
Principal species: H. gambianus group.
Subgenus b. Aethosciurus, Thomas.
Principal species: H. poensis group; H. ruzcenzorii;
H. lucifer.
Genus 19. Paraxerus, Forsyth .Major. (Synonym: Tamiscus, Thomas.)
Principal species: P. cepapi group (with ochraceus);
P. palliatus group; P. flaiivittis group; P. boehmi
group, with emini, etc.
272 SCIURIDAE
Genus 20. Fiinisciiinis, Trouessart.
Principal species: F. lemiiiscatus group; F. congicus
group; F. pvrr/iopns group, with aiiriculatus,
nixstax, carriitliersi, etc.
Genus 21. Protoxcrus. Forsvth Major.
Principal species: P. statigeri and races.
Genus 22. Mxrsiliis, Thomas.
Principal species: M. auhinii.
Genus 23. Epixeriis, Thomas.
Sole species: E. icihoni; E. ebii.
E. Xerus section. (African and some Palaearctic Ground-squirrels with
peculiar cranial characters.)
Genus 24. Atlantoxerus, Forsyth Major.
Sole species: A. getulus.
Genus 2v A'cnw, Hemprich & Ehrenberg.
Subgenus a. Xcrus, Hemprich & Ehrenberg.
Principal species: A', rutilus group.
Subgenus h. Eiixeriis, Thomas.
Principal species: A', ervthropiis group.
Subgenus c. Geoscinrus, Thomas.
Sole species : A', capeiisis, X. princeps.
Genus 26. Sperinophilopsis, Blasius.
Principal species: 5. leptodactxlus.
F. Tamias section. (Chipmunks; semi-terrestrial types, in some ways con-
necting CiteUus-Marmota section with Sciiirus section.)
Genus 27. Sciurolaniias, Miller
Subgenus a. Sciiirotamias, Miller.
Principal species: .S'. daiidianiis.
Subgenus h. RiipcsteSy Thomas.
Sole species : 5. forresti.
Genus 28. Tamias, Illiger.
Subgenus a. Ta»iicis, Illiger.
Sole species: T. striatiis and races.
Subgenus b. Eiitaiiiias, Trouessart.
Sole species: T. sibiricus and races.
Subgenus c. Neota)nias, Howell.
Principal species: T.alpiinis group; 'J'.iiiininiusgroup;
T. amoeinis group; T. quadrhittutus group; T.
toicnsendii group. (As revised by Howell, 193 1 .)
G. Marinotn section. (Ground-squirrels without the peculiarities ot the palate
and lachrymal of the Xerus section, without the peculiarities of the infra-
orbital foramen of the Tamias section, and usually, not always, with abnormal
dental characters.)
Genus 29. Citellus, Oken.
Subgenus a. Citellus, Oken.
Principal species: Palaearctic — C. Julius group;
SCIURIDAE 273
C. pygmaeus group (with erythrogenys and
others); C. citellus group (with xanthoprymnus,
alaschanicus, dauricus); C. suslicus group; C.
eversmanni group. Xearctic (arrangement of
Howell, 1938, followed) — C. townsendii group;
C washingtorii group; C. richardsonii group;
C. parryii group (with columbianus).
Subgenus b. Ictidomys, Allen.
Principal species: C. tridecemlineatus group (with
mexicanus) ; C. spilosoma group.
Subgenus c. Poliocitellus, How'ell.
Sole species : C. franklinii.
Subgenus d. Otospermophilus, Brandt.
Principal species: C. variegatus, C. beecheyi.
Subgenus e. NotociteUus, Howell.
Sole species: C. annulatus, C. adocetus.
Subgenus/. Ammospermophilus, Merriam.
Principal species: C. leucurus.
Subgenus^. Xerospermophihis, Merriam.
Principal species: C. niohavensis, C. tereticaudus.
Subgenus h. Callospermophilus, Merriam.
Principal species: C. lateralis.
Genus 30. Marmota, Blumenbach.
Principal species : .17. monax group ; M. flaviventris
group; M. caligata group (with camtschatica);
M. caudata group (with aiirea, dichrous, etc.);
M. bobak group (with sibirica, baibacina, hima-
layana) ; M. marmota group.
Genus 31. Cynomys, Rafinesque.
Subgenus a. Cynomys, Rafinesque.
Sole species: C. ludoiicianus, C. mexicanus.
Subgenus b. Leucocrossuromys, HoUister.
Principal species : C. gunnisoni group.
All specific groups recognized here, except in cases of genera which have been
definitely revised, must be regarded as provisional.
The Pteromys Group
Geographical Distribltiox. — Indo-Malayan region from Himalayas to
Ceylon, and to Sumatra, Java, Borneo and
the Philippines (not Celebes); Palaearctic, from North Scandinavia across the
northern portion of the region to Japan; Afghanistan, Kashmir, Tibet; much of
China north of the Yangtsekiang. Nearctic; from northern Canada south to
Guatemala.
Ch.\ractkrs. — This group differs from the Sciurus group in the presence of
a flying-membrane attached along the sides of the body,
rising from the wrist, and from the ankles.
lij — Living Uodents — I
274 SCIURIDAE
The cheekteeth are usually, not always, with a tendency towards excessive
complexity of pattern, which reaches its extreme deyelopment in the genera
Belomys and Tiugupterus, in which the cheekteeth are more complex in pattern
than in any other genera in the entire Order so far as my observations go.
Further, as a general rule, the zygomatic plate is low, very little tilted upwards,
and weak in general appearance; though this is not the case in the genus
Pteromvs and perhaps some others. Bullae always prominent. Cheekteeth 'i,
except in the genus lomvs.
The characters of the zygomatic plate and cheekteeth tend to show, in mv
opinion, that this group should he regarded as more primitive than the Sciuriis
group.
ExTF.RN.^L Char.\ctkrs. — The genera referred to this group agree in all
essential characters rather closely. In all genera
the flying-membrane is, as indicated above, attached to the wrist and supported
by a cartilaginous outgrowth. Posteriorly it is attached just above the ankle.
In Petaurista and Aeroiuvs, the tail is more or less narrow and round, and there
is a well-developed interfemoral membrane present; in the remainder, so far
as seen, there is no well-developed interfemoral membrane, and the tail is
wider, flatter, having an appearance very much like a large feather.
In the forefoot there are four well-developed digits, the two centre being
the longest, D.4 slightly or considerably longer than D.3, the two outer digits
subequal and a little shorter; the pollex, as usual in the group, is more or less
untraceable. In the hindfoot, the hallux, though well developed, is the shortest
digit; D.5 is usually slightly shorter than the central three, but may sometimes
tend to be as long as them; D.4 is usually slightly the longest. Claws usually
heavy, curved and powerful. The size is extremely variable; Petaurillus must
be one of the smallest of all Squirrels, while certain species of Petaurista are
as large as any other member of the family excepting certain giant forms of
Mannota. So far as known, the habits of these animals are nocturnal, thereby
differing from the non-flving Squirrels.
Thirteen groups have in this branch of the family been given generic rank
in recent years. The animals are not as common in Museums as the non-flying
Squirrels, and many of the forms are very little known. I think it is reasonable
at the moment to retain all these genera; indeed it may be that even more will
be needed as the Indo-Malavan forms become better known.
Key to the Genera oe the Pteromvs Group
Cheekteeth strongly hvpsodont; (fur excessively thick and heavy). Eupetaurus
Cheekteeth not strongly hvpsodont; (in the majority, fur not excessively
thick and heavy).
Cheekteeth always in the lower series and usually in the upper series
characterized by signs of extreme complication due to wrink-
ling; the essential pattern of the cheekteeth usually more or less
masked.
SCIURIDAE 27S
P.4 conspicuously enlarged. (Bullae not specially inflated.)
Cheekteeth semi-hypsodont ; P.4 extremely enlarged. Trogopterus
Cheekteeth brachyodont; P.4 more moderately enlarged. Belomys
P.4 not specially enlarged.
Bullae much inflated; the basi-occipital narrowed. Pteromyscus
Bullae not specially inflated, the basi-occipital noticeably wide.
(Usually the tail is narrowed.) Petaurist.a
Cheekteeth with a more normal pattern, the wrinkling though some-
times traceable never excessive, and never masking the essential
pattern.
Cusps and ridges of cheekteeth poorly marked; P.4 noticeably
smaller than M.i. Petaurillus
Cusps and ridges of cheekteeth well marked ; P.4 not smaller than
M.I.
Inner side of upper cheekteeth formed by two well-marked
approximately equal-sized cusps, the formation of the
teeth square. Lower cheekteeth with the central depres-
sion considerablv narrowed. (General dental pattern
somewhat simplified in appearance.) lOMYS
Inner side of upper cheekteeth never formed by two well-
marked approximately equal-sized cusps, the formation
of the teeth not obviously square. Lower cheekteeth
with the central depression not becoming narrowed,
excepting the genus Pteromys in which the general
dental pattern is extremely complex in appearance.
Tail rounded and narrowed. Aeromys
Tail broad, flat, feather-shaped.
Bullae low and flattened, scarcely rising above general
level of the base of the skull. Petino.MYS
Bullae without special peculiarities.
M.3 with two clear ridges between the anterior and
posterior margins of tooth ; second main ridge
of P.4, M.I and M.2 with re-entrant folds
cutting off central supplementan,- cusp;
central depression of lower molars, particu-
larly iM.3, tending to become narrow and
reduced; M.3 lower with four ridges and
three depressions; inner side of upper cheek-
teeth usually with three cusps present or
276 SCIURIDAE; BELOMYS
traceable; zygomatic plate strongly height-
ened and tilted upwards; incisive foramina
long. Ptkromys
M.3 with only one ridge between anterior and pos-
terior margins of tooth; second main ridge of
P. 4, M.i and M.2 with no re-entrant folds
cutting off central supplementary cusp;
central depression of lower cheekteeth not
tending to become reduced; M.3 lower never
with tour ridges and three depressions; inner
side ot upper cheekteeth as a rule with only
one long cusp present (as in normal Sciu-
ridae); zygomatic plate low, little tilted
upwards (except Eoglaucomxs); incisive
foramina short.
Cheekteeth relatively simpler, with small extra
ridges and depressions not or barely
traceable.
Zygomatic plate low, little tilted upwards;
hindfoot with no metatarsal pad.
Glaucomys
Zygomatic plate high, well tilted upwards;
hindtoot with metatarsal pad present.
EOGLAUCOMYS
Cheekteeth relatively more complex, with small
extra ridges and depressions normally
present. IIvlopetes
The last three genera it must be admitted are not very clearly distinguish-
able from one another.
The character of the tail, which I have used for retaining the genus Aeromvs,
is I think of sufficient importance to be used in a generic sense, in that the tail
seems to be a definite organ used by these animals for their "flying." Very
much the same state occurs in the Dipodidae, the genera Scirtopoda and
Pygeretnuis being based chiefly on the tail formation, which in these cases is
used for jumping. (Certainly if the tail in these externally specialized forms is
not considered a generic character the genus Pygeretnuis will be indistinguish-
able from the genus Ahictagidus.)
(jenus I. BELOMYS, Thomas
lyoS. Uelomys, Thomas, Ann. Mag. Nat. Hist. 8, I, p. z.
Type Species. — Sciinopterus pearsuni. Gray.
Range. — Indo-Malayan; Sikkim, Assam, Manipur, Tongking; Formosa.
BELOMYS 277
Number of Forms. — Five.
Ch.'vracters. — Skull with depressed frontals, and moderately developed
postorbital processes. Bullae large. Zygomatic plate very
primitive, little tilted upwards, only a little more specialized than the t\'pe
found in Aplodontiidae; zygoma, as in most other members of the group, long
and horizontal, being somewhat reminiscent of the zygoma of the Anomaluridae.
The ridge of the superior portion of the zygomatic plate does not extend further
forward than the level of the upper part of the infraorbital foramen.
Cheekteeth J, excessively wrinkled and complicated. P. 3 is small, P.4
much enlarged in the upper series, its anterior portion extending beyond the
small premolar in front of it, which is closely applied to the inner side of P.4.
The inner side of the upper teeth ditfer from most Sciuridae in that instead of
being formed by one large elongate cusp, there are three cusps present which
evidently do not join, the front one being the smallest. The teeth are extremely
complex; what might become a normal Sciurine pattern can be vaguely traced
among the mountain-like elevations and deep depressions covering the whole
surface of the teeth; the elevations are arranged in three primary longitudinal
rows. A well-marked external projecting angle is present on each upper tooth,
the centre of which is divided by a deep re-entrant fold; this appears to corre-
spond to the space between the two main ridges in normal Sciuridae. M.3, even
in these teeth, appears more simplified than the other molars, this being a very
common feature throughout the family.
Lower teeth exceptionally complicated; M.3 the longest tooth. Four main
cusps present, or may be traced, the anterointernal one as usual the highest.
In M.3 there appear to be at least five transverse ridges extending across the
central part of the tooth, but each is much broken up.
Essential external characters as already described; ear rather large, with a
tuft of long bristles or hairs at base.
Forms seen : pearsoni, kakensis, trichotis, blandus.
I am not convinced that there is more than one species of this rather excep-
tional genus, and accordingly treat all named forms provisionally as races of
the type.
List of Named Forms
(The references and type localities to all forms belonging to this group are
the work of i\Ir. R. W. Hayman.)
1. BELOMYS PEARSONI PEARSONI, Gray
1842. Ann. Mag. Nat. Hist., X, p. 263.
Darjiling, Sikkim.
2. BELOMYS PE.ARSONI BLANDUS, Osgood
1932. Field Mus. Nat. Hist. Zool., XVIH, no. 2, p. 269.
Muong Moun, south of Lai Chau, Tongking.
3. BELOMYS PEARSONI VILLOSUS, Blyth
1847. Joum. Asiat. Soc. Bengal, XVI, p. 866.
Upper Assam.
Fig. 82. Belomys pearsom trichotis, Thomas.
B.M. No. I5.5-5-43. tj; X 2.
Fig. S3. Belomys pearsoni trichotis, Thomas.
B.M. No. I5.5-S-43. S< >- 2-
BELOMYS— TROGOPTERUS 279
4. BELOMYS PKARSONI TRICHOTIS, Thomas
1908. Ann. Mag. Nat. Hist. 8, I, p. 7.
Machi, Manipur.
5. BELOMYS PLARSONI KALKliN'SIS, Swinhoe
1862. Proc. Zool. Soc. London, p. 359.
North Formosa.
Genus 2. TROGOPTERUS, Heude
1898. TROGOPTERUS, Heude, Mem. Hist. Nat. Chinois, IV, pt. i, pp. 46-47.
Type Species. — Pteromys xanthipes, Milne-Edwards.
Range. — China; known from Tibet, Szechuan, Ichang, Shensi, Yunnan,
Chihli, etc.
Number of Forms. — Five.
Characters. — Like Belotnys, but P. 4 even more enlarged in the upper
toothrow, and cheekteeth semi-hypsodont (brachyodont in
Belomys). Zygomatic plate like Belomys, but with a prominent knob under the
infraorbital foramen for muscle attachment; this knob, often present in
Sciuridae, I shall refer to as the " masseter-knob." The upper part of the
zygomatic plate is more ridged than in Belomys. Bullae large. Upper cheek-
teeth excessively wrinkled, the elevations arranged in three primary rows. The
external projection in the main upper teeth present, though usually smaller
than in Belomys. P. 3 present, closely applied to the inner side of P. 4, which
projects anteriorly considerably beyond it, and is extremely large. Teeth large
and heavy; the general effect complex in the extreme. Lower teeth with four
main cusps, one at each corner, but the pattern as complexly wrinkled and
folded as in the upper series. M.3 relatively less enlarged than in Belomys.
Mandible with angular portion rather sharply pulled inwards; coronoid high,
recurved.
Essential external characters as in Belomys; sole may be partly haired.
Forms seen : .xanthipes, mordax, minax, himalaicus, edithae.
Thomas has divided the limited British Museum material into five separate
species. I do not think that there is more than a racial difference between any
of the named forms. Until more material comes to hand it seems to be more
correct to regard all named forms as subspecies of xanthipes.
List of X.'UIEd Forms
1. TROGOPTERUS X.\NTHIPES X.\NTHIPES, Milne-Edwards
1867. .Ann. Sci. Nat. Zool., VIH, p. 376.
Chihh, North China.
2. TROGOPTERUS X,\NTHIPES MORD.\X, Thomas
1914. Joum. Bombay Nat. Hist. Soc, XXHI, 2, p. 230.
Ichang, Yangtze-kiang, China.
3. TROGOPTERUS XANTHIPES H1^L■\LAICUS. Thomas
1914. Joum. Bombay Nat. Hist. Soc, XXIII, 2, p. 231.
Gyantse, Chumbi Valley. Tibet.
28o TROGOPTERUS— PTEROMYSCUS
4. I'ROGOPTERLS XANTHIPES EDITHAE, Thomas
1923. Ann. Mag. Nat. Hist. <j, XI, p. 65S.
North-west flank of Likiang Range, Yunnan.
5. TROGOPTERUS XANTHIPES MINAX, Thomas
1923. Ann. Mag. Nat. Hist. 9, XI, p. 660.
Won Cauen, Upper Min River, .Szechuan, China.
Fig. 84. TROGOPTERUS xanthipes mord.\x, Thomas.
B.M. No. 22.9.1.46, V; X 5.
Genus 3. PTEROMYSCUS, Thomas
1908. PTEROMYSCUS, Thomas, Ann. Mag. Nat. Hist. 8, I, p. 3.
Type Species. — Sciiiropterus pulverukntus, Giinther.
R.'iNGE. — Penang, Sumatra, Borneo.
Number of Forms. — Two.
Characters. — Much like Belomys, but with P. 3 vestigial, and P. 4 not
conspicuously larger than M.i; also differing in details of
the pattern of the upper teeth, and with the bullae relatively very much enlarged.
The teeth when worn appear to present a rather more normal, less wrinkled
appearance, but in younger skulls the complexity of the molars is great. The
external projection in the upper molars is less marked than in Belomys, and
apparently there are only two inner cusps present, the small anterior one being
barely traceable. The lower teeth are more or less as m Belomys; M.3 con-
siderably elongated.
PTEROMYSCUS— PETAURISTA 281
Externally like Belomys, but ear smaller and without tufts.
Forms seen : pulverulentus, borneanus.
List of Named Forms
1. PTIiRO.MYSCUS PULVERULENTUS PULVERULENTUS, Gunther
1873. Proc. Zool. Soc. London, p. 413, pi. x.x.\viii.
Penang, Malay Peninsula.
2. PTEROMYSCUS PULVERULENTUS BORNEANUS, Thomas
1908. Ann. Mag. Xat. Hist. 8, I, p. 7.
Barani, Sarawak, Borneo.
Genus 4. PETAURISTA, Link
1795. Pet.\irist.^, Link, Zool. Beytr., i, pt. ii, pp. 52, 78.
Type Species. — Sciurus petaurista, Pallas.
Range. — Palaearctic and Indo-Malayan; Ceylon, Peninsular India (southern
portion, Surat, Orissa); Punjab, Kashmir; Kumaon, Nepal,
Sikkim; Burma (Chindwin, Chin Hills, Arakan, Shan States), Tenasserim;
Yunnan, Fukien, Hainan, Formosa. Tongking, Siam, Annam, Malay Penin-
sula; Sumatra, Java, Borneo, Natunas. Also in Szechuan, Hupeh, and China
north of the Yangtsekiang; South Kansu; Tibet; Chihli, South Manchuria,
Korea, and Japan.
Number of Forms. — ^About sixty-one.
Characters. — Skull characterized by very large postorbital processes
standing nearly at right angles to the braincase; frontals
deeply depressed ; parietal ridges well marked but showing no signs of joining
or even approaching each other in any seen. Jugal with superior process
pointing upwards below postorbital process, a structure often to be seen in the
present group. Bullae usually large but not extremely so; palate broad. Zygo-
matic plate similar in general type to that of Belomvs, but more prominently
ridged. Cheekteeth somewhat intermediate between the wrinkled type of
Belomys and Trogopterus and the more normal type found in Hxlopetes and
others. In the upper toothrow each main upper tooth has three inner main
cusps originally, but in worn teeth these tend to come together; but a well-
marked posterior re-entrant fold (originally between cusps 2 and 3) appears
always to be retained; sometimes three inner folds are present. The fold which
is retained sometimes appears as a pit; it is present in M.3, which is normally
as complex as the other molars, not simplified as is usual in Sciuridae. The
normal Sciurine pattern of four ridges and three depressions is traceable, but
there is often a tendency towards wrinkling, though less marked than in
Trogopterus and Belomys. P.4 is sometimes rather larger than M.i. P. 3 well
developed. The lower cheekteeth agree with those of Trogopterus and Belomvs,
and are excessively complex; M.3 is not elongated. The central depression is
barely traceable as a rule, and the crown surface when worn usually presents
282
PKTAURISTA
four or more broken up isolated depressions, and with many small ridges
running across the surfaces of the teeth. A well-marked depression in front of
the anteroexternal main cusp present, and usually one between the two outer
main cusps present. Teeth semi-hvpsodont.
In P. siilcatiis, not represented in London, the upper incisors are described
as broad, and grooved. The describer states that fuhimis may have faintly
grooved incisors, and does not consider the character generic; but it makes the
retention of such genera as Srnt/wosciiinis more than doubtful. The upper
incisors are normally plain in this genus.
Size large; up to 464 mm. head and body or perhaps more. Interfemoral
Fig. 85. Petaurist.a philippensis philippensis, Elliot.
B.M. No. 13.8.22.3s, <S; ■■: i.
membrane usually more developed than in other genera; tail usually longer than
head and body, as a rule narrowed and rounded though fully haired; but in
some forms, as leiicogenvs, perhaps on account of the cold climate in which they
live, the tail appears in adult to be much broader, and approaching the formation
found in the smaller Flying-squirrels, though a young leucogenys seen has a
narrow tail, as in normal Petaurista. D.4 considerably longer than D.3 in
manus, as a rule.
Forms seen : dlbiventer, alboiiifiis, aniiamcusis, badiatus, barroiii, baliiaiia,
birrelH, candididus, caniceps, castaneus, Cinderella, cineraccus, clarki, elegans,
fulvimis, gorkhali, grandis, hintoiii, iiiornaiiis, latika, lend, leucogenys, lylei,
magnlficus, marchio, marica, inehmotis, mergidus, nigricaudatus, nikkonis, nitidula,
nobilis, ochraspis, oral, areas, petaurista, philippensis, priinrosei. punctata, rajah,
reguli, senex, sibyl/a, tavlori, tosae, lenningi, xanthotis, yunnanensis.
Fig. 86. Petaurista philippensis philippensis, Elliot.
B.M. No. 13.8.22.35, (J; X I.
Fig. 87. Petaurista philippensis philippe.nsis, Elliot.
Cheekteeth: B.M. No. 13.8.22.35, o; x 4j.
284 PETAURISTA
This genus contains very many standing distinct species, which are most
difficult to arrange in any natural order. Many of the species are known only
by very few specimens; some are based on one skin without a skull.
Wroughton (191 1, Journ. Bombay Nat. Hist. Soc. XX, 4, p. 10 12) has keyed
the majority of the Indo-Malayan species, but does not include the Palaearctic
ones; Robinson & Kloss, with the exception of the Malay Island forms, did not
attempt anv revision as regards the reducing of the more doubtful species to
races.
I am inclined provisionally to recognize four groups, one of which contains
the majoritv of the genus and is divisible into several sections.
The .XLBOKl'FUS group contains forms with a striking and highly specialized
red and white colour-pattern (the head white, the back mostly deep red, with
or without a white or brownish dorsal patch). Large thick-furred forms,
with bushy tail. China, south to Yunnan, east to Tibet; and Formosa
{/eiia).
The remaining groups have no well-marked specialized colour-pattern as
indicated above.
The PET.^i'KisT.A group contams forms which are mostly unicolorous; most
often deep red in general coloration, or in one race blackish; rather short-furred
types; back not grizzled. Chiefly Malay Islands, though a race is named from
South China.
All other groups examined have a certain amount of grizzling apparent,
sometimes strongly marked, on dorsal surface.
The .XLBIVENTER group contains the remaining forms in the genus which
are represented in London, and divides apparently into the following
sections :
puiictatiis section : back with conspicuous white spots present. Malacca,
Borneo, and marica from Yunnan. These are very distinct types,
but do not appear to be well known, and it has been suggested
that the peculiar culoration is in these skins due to disease. Rather
small forms.
plulippcnsis section : brownish grizzled white, the white always conspicuous.
Rather thick-furred species. Tail usually longer than head and body.
Containing pliilippensis and the other species from Ceylon and Peninsular
India, also hlei and cineniceus from Siam. This section rather grades
into the
ulbiventer section : upper portion usually without conspicuous white grizzling,
less frosted in general appearance; frequently more or less reddish in
coloration. Tail not specially broadened, or less densely bushy than in
leucogenys section; fur thick to extremely thick. The most important
species referred to this section are iiiornatus. cuniceps, albiventer (all very
thick-furred), and magnificus, from the Himalayas; and apparently
mcrgulus frf>m islands of the Mergui Archipelago. " Pleroinys" gorkliali
is a Petaurista very closely allied to or perhaps a subspecies of caniceps.
The group appears to be represented in Burma by candidulus, which has
PETAURISTA 285
white grizzling present, hut is much redder in general coloration than
anv member of the pluUppensis section.
kucogenys section : this is closely allied to the last, but appears to differ in
the very general broad appearance of the tail (more so than in other
Petaurista); the immensely thick fur, and rather Eupetaurus-Vikc general
appearance; and contains the Japanese and Manchurian kucogenys, and
xantholis from Tibet. P. melanoplerus, not seen, is usually considered as
allied to this branch.
The SULC.\TL"S group contains one species (not seen) differing apparently
from the others in the grooved incisors, though it must be borne in mind that
the incisors can be grooved in individual cases elsewhere in the genus.
This arrangement must be regarded as provisional.
List of Named Forms
petaurista Group
1. PETAURISTA ELEGANS, Temminck
1839-44. Miiller & Schlegel, Verhandl. Nat. Gesch., pp. 107, 112, pi. xvi, figs. 1-3.
Island south of Nusa Kumbang, South Java.
2. PETAURISTA PETAURISTA PETAURISTA, Pallas
1766. Misc. Zool., p. 54.
West Java.
Synonj-m: nitida, Desmarest, 1818, Xouv. Diet. Hist. Nat., XXVII,
p. 403. Java.
taguan. Link, 1795, Zool. Beytr. i (2), p. 78.
3. PETAURISTA PETAURISTA XIGRICAUDATUS, Robinson & Kloss
1918. Joum. Fed. Malay States Mas., VII, p. 223.
Ongop Ongop, Banjoewangi, East Java.
4. PETAURISTA PETAURISTA MELANOTUS, Gray
1837. Charlesworth's Mag. Nat. Hist.. I, p. 584.
"Nepal" (error), Malay Peninsula substituted.
5. PETAURISTA PETAURISTA CICUR. Robinson & Kloss
1914. .■\nn. Mag. Nat. Hist. 8, XIII. p. 223.
Bandon, Siamese Malaya.
6. PETAURLSTA PETAURISTA R.MAH. Thomas
1908. Ann. Mag. Nat. Hist. 8, I, p. 251.
iVIount Dulit, Baram, Borneo.
7. PETAURISTA PETAURISTA MTIDUEUS, Thomas
1900. Nov. Zool., VII, p. 592.
Bunguran, North Natuna Islands.
8. PETAURISTA PETAURISTA B.\TU.\NA, .Miller
1903. Smiths. Misc. Coll., XLV, p. 27.
Tana Bala, Batu Islands, \V. Sumatra.
Synonym: marchio, Thomas, 1908, Ann. Mag. Nat. Hist. 8, i, p. 251.
Si Ramba, Sumatra.
<). PET.\URISTA PET.AURISTA TERUTAUS, Lyon
1907. Proc. Biol. Soc. Washington, XX, p. 17.
Terutau Island, northern Straits of Malacca.
286 PETAURISTA
10. PETAURISTA PETAURISTA MIMICUS, Miller
1913. Smiths. Misc. Coll., LXI, no. 21, p. 27.
Pulau Rupat, East Sumatra.
11. PETAURISTA PETAURISTA LUMHOLTZI, Gyldenstolpe
igig. Stockholm Vet. Akad. Handl. 60, 6, p. 28.
Poeroek Tjahoe, Central Borneo.
12. PETAURISTA PETAURISTA RUFIPES, G.Allen
1925. Amer. Mus. Nov. 163, p. 13.
Yungan, Fukien Province, China.
13. PETAURISTA GRANDIS, Swinhoe
1862. Proc. Zool. Soc. London, p. 35S, pi. .xlv.
Formosa.
iilhonifus Group
14. PETAURISTA ALBORUEUS ALBORUFUS, Milne-Edwards
1870. Compt. Rend., LXX, p. 342.
Moupin, Szechuan.
15. PETAURISTA ALBORUFUS LEUCOCEPHALUS, Hilzheimer
1906. Zool. Anz., XXIX, p. 298.
Tibet.
16. PETAURI.STA ALBORUFUS CASTANEUS, Thomas
1923. Ann. Mag. Nat. Hist. 9. XII, p. 172.
Ichang, Middle Yangtsekiang, China.
17. PETAURISTA ALBORUFUS C^CHRASPIS, Thomas
1923. Ann. Mag. Nat. Hist. 9. XH, p. 172.
Likiang Range, N.-W. Y'unnan.
18. PETAURISTA LENA, Thomas
1907. Ann. Mag. Nat. Hist. 7, XX, p. 522.
Tapposha, Central Formosa.
albiventer Group
{pimctatus Section)
iq. PETAliRISTA PUNCTATUS PUNCTATUS, Gray
1846. Ann. Mag. Nat. Hist., XVHI, p. 211.
Malacca.
;o. PETAURISTA PUNCTATUS BANKSI, Chasen
1934. Bull. Raffles Mus. 8, p. 194.
Mount Kina Balu, Borneo.
21. PETAURISTA PUNCTATUS MARICA, Thomas
1912. Ann. Mag. Nat. Hist. 8, IX, p. 6S7.
Y'unnan ; probably near Mongtze.
22. PETAURISTA PUNCTATUS SYBILLA, Thomas
1916. Journ. Bombay Nat. Hist. Soc, XXIV, 3, p. 423.
Chin Hills, near Kindat, Upper Burma.
(philippensis Section)
23. PETAURISTA PHILIPPENSIS PHILIPPENSIS, Elliot
1839. Madras Journ. Lit. and Sc, X, p. 217.
Near Madras, India.
Synonym: (}) griseiventer. Gray, 1843, List Mamm., p. 133.
PETAURISTA 287
24. PETAURISTA PHILIPPENSIS ORAL, Tickell
1842. Calcutta Journ. Nat. Hist., II, p. 401, pi. XI.
Singhbum district, Bengal.
25. I'KTAURlSrA PHILIPPENSIS CINDERELLA, Wroughton
1911. Journ. Bombay Nat. Hist. Soc, XX, 4, pp. 1014, 1018.
Sural district, Bombay.
26. PETAURISTA PHILIPPENSIS LANKA, Wroughton
191 1. Journ. Bombay Nat. Hist. Soc, XX, 4, pp. 1014, 1017.
Ceylon.
27. PETAURISTA CINERACEUS CINERACEUS, BIyth
1847. Journ. Asiat. Soc. Bengal, XVI, p. 865.
Arakan.
28. PETAURISTA CINER.\CEUS STOCKLEYI, Carter
1933. .'Xmer. Mus. Nov., 674, p. i.
Melamoong, N.-W. Siam.
2<;. PETAURISTA LYLEI LYLEI, Bonhote
igoo. Proc. Zool. Soc. London, p. 192.
Doi Sritepe, Chiengmai, N. Siam.
30. PETAURISTA LYLEI VENNINGI, Thomas
1914. Journ. Bombay Nat. Hist. Soc, XXIII, i, p. 27.
Kalaw, Southern Shan States, Burma.
31. PET.\URISTA LYLEI BADIATUS, Thomas
1925. Proc. Zool. Soc, London, p. 501.
Ngai-Tio, Central Tonkin.
{albiventer Section)
32. PETAURISTA MERGULUS MERGULUS, Thomas
1922. Journ. Bombay Nat. Hist. Soc, XXVIII, p. 1067.
Ross Island, Mergui Archipelago.
33. PETAURISTA MERGULUS REGULI, Thomas
1926. Journ. Bombay Nat. Hist. Soc, XXXI, p. 22.
King Island, Mergui Archipelago.
34. PETAURISTA MERGULUS PRIMROSEI, Thomas
1926. Journ. Bombay Nat. Hist. Soc, XXXI, p. 22.
Sullivan Island, Mergui Archipelago.
35. PET.\URISTA .\NN.\MENSIS ANNAMENSIS, Thomas
1914. Journ. Bombay Nat. Hist. Soc, XXIII, 2, p. 204.
Bali, Nha-Trang, South Annam.
36. PETAURISTA ANN.\MENSIS BARRONI, Kloss
1916. Journ. Nat. Hist. Soc. Siam, II, p. 33.
Hup Bon, Sriracha, S.-E. Siam.
37- PETAURISTA YUNNANENSIS, .\nderson
1875. Ann. Mag. Nat. Hist. 4, XVI, p. 282.
Momein, Yunnan.
3S. PETAURISTA CANDIUULUS, Wroufihton
1911. Journ. Bombay Nat. Hist. Soc, XX, 4, pp. 1014, 1022.
Kindat, Upper Chindwin, Burma.
288 PETAURISTA
39. PETAURISTA TAYLOR], Thomas
1914. Journ. Bombay Nat. Hist. Soc. XXIII, p. 205.
Bankason, South Tenasserim.
40. PETAURISTA FULVINUS, Wroughton
igii. Journ. Nat. Hist. Soc. Bombay, XX, 4, pp. 1014, 1021.
Simla, West Himalayas.
41. PETAURISTA AI.BIVE.NTER, Gray
1834. 111. Ind. Zool., pi. xviii.
No locality. (Occurs Nepal, Kumaon (Wroughton).)
42. PETAURISTA MAGNIFICUS, Hodgson
1836. Journ. Asiat. Soc. Bengal, V, p. 231.
Nepal.
Synonym : Ho/j/fc, Gray, 1842, Ann. Mag. Nat. Hist., X, p. 263. Darjiling.
chrvwthri.x, Hodgson, 1844, Journ. Asiat. Soc. Bengal,
' XIII, p. 67.
43. PETAURISTA INORNATUS, Gcoffroy
1844. In Jacquemont's Voyage, IV, Mamni,, p. 62, Atlas ii, pi. iv.
North India.
44. PETAURISTA BIRRELLI, Wroughton
1911. Journ. Bombay Nat. Hist. Soc, XX, 4, pp. 1014, 1019.
Murree, Hazara, Punjab.
45. PETAURISTA CAXICEPS, Gray
1842. Ann. Mag. Nat. Hist., X, p. 262.
Sikkim.
Synonym: senex. Hodgson, 1844, Journ. Asiat. Soc. Bengal, XIII, p. 68.
Nepal.
46. PETALRISTA GORKHALI, Lindsay
1929. Journ. Bombay Nat. Hist. Soc, XXXIII, 3, p. 566.
Gorkha, Nepal (12,000 ft.).
47. PETAURISTA CLARKEI, Thomas
1922. Ann. Mag. Nat. Hist. 9, X, p. 396.
Mekong Valley, Yunnan.
(leucogen\s Section)
48. PETAURISTA XANTHOTIS, Milne-Eduards
1872. Ann. Sci. Nat. Zool., p. 301.
"Tibet" (probably Moupin, Szechuan).
49. PETAURISTA LEUCOGENYS LEUCOGENYS, Temminck
1827. Mon. Mamm. i, Tab. Method, p. xxvii.
Japan.
50. PETAURISTA LEUCOGENYS NIKKONIS, Thomas
1905. Ann. Mag. Nat. Hist., 7, XV, p. 488.
Nikko, Central Hondo, Japan.
51. PETAURISTA LEUCOGE.NYS OREAS, Thomas
1905. Ann. Mag. Nat. Hist., 7, XV, p. 488.
Wakayama, South Hondo, Japan.
52. PETAURISTA LEUCOGENYS TOSAE, Thomas
1905. Ann. Mag. Nat. Hist., 7, XV, p. 488.
Tosa, Sikoku Island, Japan.
PETAURISTA 289
53. PETAURISTA LEUCOGENYS HINTONI, Mori
1923. Joum. Mamm. Baltimore, 4, p. 191.
Seoul, Korea.
Synonym: thomasi, Kuroda & Mori, 1923, Joum. Mamm. Baltimore, 4,
p. 27. Seoul, Korea.
54. PETAURISTA W.-^TASEI, Mori
1927. Annot. Zool. Jap., 11, ii, p. 107.
Mukden, S. Manchuria.
55. PETAURISTA MELANOPTERUS, Milne-Edwards
1867. .\nn. Sci. Nat. Zool., VIII, p. 375.
Tcheli, China.
sulcatus Group
56. PETAURISTA SULCATUS, Howell
1927. Joum. Washington Acad. Sci. XVII, p. 82.
Hsinlungshan, 65 miles north-east of Peking, Chih-li, E. China.
Not seen and not allocated to group
57. PETAURISTA RUBICUNDUS, Howell
1927. Joum. Washington Acad. Sci. XVII, p. 82.
Mapientung, Szechuan, China.
58. PETAURISTA HAINANA, G. M. .\llen
1925. Amer. Mus. Nov. 163, p. 14.
Nam Fong, Hainan.
59- PETAURISTA PECTORALIS, Swinhoe
1870. Proc. Zool. Soc. London, p. 634.
Takow, S.W. Formosa.
60. PETAURISTA FILCHNERINAE, Matschie
1908. E.xp. Filchner China & Tibet, Zool. Bot. Ergebn., p. 208.
Si-ning-Fu, China (Upper Hwang-Ho, Kansu).
Probably = xanthotis, according to Howell.
Addenda:
PETAURISTA PETAURISTA PENANGENSIS, Kloss
1918. Joum. Fed. Malay States Mus. VII, p. 224.
Telok Bahang, Penang Island.
PETAURISTA, PUNCTATA SUM-VPRANA, Kloss
1 92 1. Joum. Fed. Malay States Mus. X, p. 230.
Padang Highlands, W. Sumatra.
For references purposes 1 include Wroughton's key to the species of Petau-
risla occurring in India (1919, Journ. Bombay Nat. Hist. Soc. XXVI, No. 2,
p. 354). All these forms are regarded as belonging to the albiventer group.
" General colour blackish or greyish, never rufous or fulvous.
Smaller, hindfoot 70-77.
Smaller, hindfoot 72. oral
Larger, hindfoot 77. Cinderella
19 — Li\iny KtKlent-s — I
290 PETAURISTA— AEROMYS
Larger, hindfoot S0-S5.
Back of ears and toreanii bay; tail drab-grev. cineraceus
No bay marking; tail black.
Limbs and jiarachute dark maruon, under surface salmon buff.
lyki (lylei venningi)
Limbs and parachute like the back, at most with a rufous
tinge; under-surface white.
Limbs and parachute with a rufous tinge. pliilippeiisis
Limbs and parachute like the back. lanka
General colouring rufous or fulvous.
Size larger, hindfoot over 80 mm.
Colour darker; black tufts behind the ears. ta\Iori
Colour paler, dark bav tufts behind the ears. candiduliis
Size smaller, hindfoot 65-77.
Larger, hindtoot 70-77.
A well-marked dark saddle-patch extending forward to the
crown; hindfoot 73. nobilis
No saddle patch.
Back of ears black.
Colour darker, grizzled bay and buff. birrelli
Colour paler, grizzled brown and white. inornatus
Back of ears coloured like head.
Face grey. caniceps
Face like head and back.
Darker (bav); no pale area on shoulders; hindfeet
black. alhiventer
Paler (ferruginous); shoulders slightly paler than
back ; feet coloured like back. fulvimis
Smaller, hindfoot 60-65. sibylla"
The forms oral, Cinderella and lanka are regarded as subspecies o{ philippensis
by Robinson & Kloss in their list of Oriental Sciuridac; sibvlla is regarded as a
race of pimctatns in this paper (as is also marica from Yunnan); these authors
use the name magnifuus instead of nobilis.
Genus 5. AEROMYS, Robinson & Kloss
1915. .\eromvs, Robinson &; Kloss, Journ. Fed. Malay States Mus. VI, p. 23.
TvFE Species. — Pteromys tephromelas, Giinther.
R.\.\GE. — Penang, Borneo and Sumatra.
Nl'mber of Forms. — Three.
Cli.VRACTERS. — External characters, including the interfemoral and the
narrow round tail (which is much narrowed and very long)
essentially as in Petaiirista. Skull near Pctaurista. But cheekteeth with in the
adult no wrinkling, relatively simple, and of similar pattern to Hylopetes (below),
AEROMYS— PTEROMYS 291
with which group Thomas in 1908 associated the genus. Forsyth Major pointed
out that this group agrees in dental characters with the smaller Flying-squirrels,
rather than with the Petaurista type. But in a skull in which the teeth are just
being cut, the wrinkling is extreme.
For remarks on the desirability of retaining this genus see p. 276.
Forms seen : bartelsi, phaeomelas, tepliromelas, thomasi.
Two groups may be recognized here provisionally, tephromelas and phae-
omelas, very dark blackish forms, and thomasi which has a very attractive deep
red colour.
List of Named Forms
tephromelas Group
1. AEROMYS TEPHROMELAS, Giinther
1873. Proc. Zool. Soc. London, p. 413, pi. xxxvii.
Penang, Malay Peninsula.
2. AEROMYS PH.\EOMELAS, Gunther
1873. Proc. Zool. Soc. London, p. 413.
Borneo. (Should be regarded as a race of tephromelas ?)
3. AEROMYS BARTELSI, Body
1936. Natuurk. Tijschr. Ned. Ind. 96, p. 146.
Pagar Djawa, Pematang .Siantar, Deli, N. Sumatra.
(Described doubtfully as Petaurista; now seen to be Aeromys.)
thomasi Group
4. AEROMYS THO^L■\SI, Hose
1900. .Ann. Mag. Nat. Hist. 7, V, p. 215.
Baram, Sarawak, Borneo.
Genus 6. PTEROMYS, Cuvier
1800. Pteromys, Cuvier, Tabl. Elem. Hist. Nat. Anim. p. 135.
1825. SciiROPTERUS, F. Cuvier, Dents, des Mamm. 161-162, pi. 56, p. 255. Scitirus
volanSy Linnaeus.
Type Species. — Sciurus volans, Linnaeus.
Range. — Palaearctic; Northern Scandinavia, Finland, Lithuania, Latvia,
Estonia; European Russia, south to former Minsk, .Smolensk,
Riazan, Vladimir, Kasan and Orenberg governments (Vinogradov); across
wooded Siberia ; quoted by \'inogradov from Pavlodar district. North Kazakstan ;
Anadyr region; Transbaikalia. .Manchuria, Korea, Japan; Sakhalin; Kansu.
Number of Forms. — Thirteen.
Characters. — Zygomatic plate much more specialized than in other members
of the group, being considerably heightened, powerfully
ridged on its superior border, the ridge extending beyond the general line of
the zygomatic plate, which is hollowed to a certain degree. Masseter knob
Fig. 88. Pteromys volans vol.^ns, Linnaeus.
B.M. No. 1. 6. 9. 1, c?; 2-
Fig. 89. Pteromys vol,\ns vola.ns, Linnaeus.
B.M. No. I. e.g. I, rf; .■; 2.
PTEROMYS
293
Fig. 90.
Pteromys volans.
Cheekteeth; X 5.
of infraorbital foramen prominent. Frontals depressed, braincase smooth
and .strongly depressed posteriorly. Bullae large. Incisive foramina larger
(longer) than in most other Flying-squirrels, ^hmdible with angular portion
strongly pulled inwards. Cheekteeth without the excessive wrinkling character-
istic of Belotnxs and allies, but much more complex in general appearance than
in Glaucamys and allies. P. 3 small. Upper series with the inner side of the
tooth composed of three more or less distinct cusps; general pattern otherwise
not far removed froin that of Sciurus, but the second
main ridge of P. 4, M.i and M.2 is cut by a deep re-
entrant fold which together with another depression
next to the raised inner border of the tooth isolates the
ntermediate portion of the ridge as a high and distinct
cusp, traces of this to be seen apparently at all times;
a further peculiarity is that iVI.3 is not simplified, but
retains two high main transverse ridges between the
anterior and posterior margins of the tooth, a very rare
feature in this family. In the lower teeth, M.3 is greatly
elongated ; the cusps and ridges of the teeth are very
prominent; the cusp between the two outer main cusps
usually takes the form of a ridge; the posterointernal
cusp is high and broad, and the central depression
characteristic of most Sciuridae is rather reduced. A
high ridge rounds off the posterior portion of M.3; in
this tooth, usually there are traceable three depressions between four transverse
ridges, the second and third of which are rather low.
External characters as usual in the group; sole densely haired in all examined;
size not large.
The genus Pteromys, " Sciuropteriis" as arranged by Thomas in 1908,
contained Glaucomys, Hylopetes and Petinomys as subgenera. Whatever may be
the fate of these, there is no doubt in my mind that by the unique dental
characters combined with the strongly specialized zygomatic plate the genus
Pteromys must be restricted to the northern Palaearctic small Flying-squirrels,
and is very distinct from all others. It is regrettable that the name "Pteromys,"
which has in the past been used for the large Flying-squirrels of the genus
Petaurista cannot be dropped in favour of the much more widely known
Sciurv[>teriis.
Forins seen: " russicus" { = volans), aliico, atliene, amygdalei, momotiga.
It is not clear whether there is inore than one valid species belonging to this
genus.
List of N.amed Forms
I. PTKROMYS VOLANS VOLANS, Linnaeus
1758. Syst. Nat., loth Ed., voL i, p. 64.
Sweden.
Synonym: rHSjiViM, Ticdcmann, 1S08. Zoologie, vol. i,p. 154. Finland.
sibiricus. Dcsmarcst, Mammologie, II, p. 342, 1822.
vulgaris, Wagner, Schreber, Siiugt. Suppl. Ill, p. 228, 1843.
204 I'TEROMYS— GLAUCOMYS
;. PTEROMVS \C)LANS OGNEVl, Strngnnov
193(1. Zool, J. Moscow, 15, p. 539, 559.
Lake Peno, Kalininschen Region, in estuary of the Volga, Gouv. Twer,
Russia.
3. PTF.ROMYS \X)I..\NS C;L"13AR1. Ogncv
1935. Bull. Soc. Nat. Moscow, 43, 1934. pp. 304, 311.
West Siberia, district of Troitzk, formerly Bijsk.
4. PTEUOMYS \-()LAXS TUROVI, Ogncv
1029. Bull. Pac. Sci. Fishery Res. Sta.. II, pp. 14, 41.
Peninsula Koty, Baikal, Siberia.
5. PTERUMYS VOEAXS BETL'LINUS, Serehrcnnikov
1930. Zeitschr. fiir Saugctierk. 4, Heft 3, p. 142.
Pavlodar, Semipalatinsk, Siberia.
6. PTEROMYS VOLANS IN'CANUS, MiUer
1918. Proc. Biol. Soc. Washington, XXXI, p. 3.
East Siberia; Verkhne Kolymssk.
7. PTERCJMY.S VOLANS ATHENE, Thomas
1907. Proc. Zool. Soc. London, p. 409.
Korsakoff, Saghalien.
8. PTEROMYS VOLANS ALUCO, Thomas
1907. Proc. Zool. Soc. London, p. 464.
Kaloguai, 55 miles north-east of Seoul, Korea.
.). PTEROMYS VOLANS ARSENJIiVI, Ognev
1935. Bull. Soc. Nat. Moscow, 43 (1934), pp. 309, 314.
Ussuri.
10. PTEROMYS BUECHNERI, Satunin
1903. Ann. Mus. .St. Petersb., VII, p. 549.
Kansu, China.
11. PTEROMYS ORII, Kuroda
1921. Journ. Mamni. Baltimore, 2, p. 208.
Llyenai, Iburi Province, Hokkaido, Japan.
12. PTEROMYS MOMONGA MOMONGA, Temminck
1847. Faun. Japon, p. 47, pi. 14.
Interior of Japan.
13. PTEROMYS MOMONGA AMYGDALEI. Thomas
1906. Proc. Zool. Soc. London, p. 344.
Washikaguchi, Nara Ken, South Central Hondo, Japan.
Genus 7. GLAUCOMYS, Thomas
1908. Gl.-m'comys, Thomas, Ann. Mag. Nat. Hist., ser. 8, vol. I, p. 5.
Type Spfx'ie.s. — i\Ins rolaiis, Linnaeus.
R.ANGE. — North America: Alaska, Keewatin, Labrador, Manitoba, Alberta,
British Colimnbia, Washington, Oregon, Idaho, California, LUah,
Texas, Alabama, Florida, Virginia, New York (good distribution maps pub-
lished by Anthony (alter Howell), Field Book North. Amer. Mamm. 1928, for
all forms occurring north of Mexico); South Mexico, Honduras.
GLAUCOMYS 295
Number of Forms. — Thirty.
Characters. — Cheekteeth relatively simple, not essentially different in
general arrangement from Sciuriis, and with no traces of the
extra complications seen in the Malayan Ilylopetes and Petinumys. In the lower
teeth, iM.3 is less enlarged than in Ptcrumys, and the central depression is not
reduced, so far as seen.
In the upper cheekteeth, M.3 is simple, as usual in the family, lacking the
third (second main) transverse ridge of Pteromys. liullae large. Incisive fora-
mina very short. Frontals not depressed. Zygomatic plate low and primitive,
not comparing with Pteromvs, and much lower than in Eoglaucomys in all seen.
Postorbital process relativelv small.
Size rather small. Soles, at any rate in winter, densely haired, the metatarsal
pad characteristic of most members of the group being absent. Mammae 8.
Not many specimens of this genus have been available for examination, but
the genus has been fully revised by Howell (North Amer. Fauna, No. 44, 1918).
In this paper very many skulls are figured, and the genus is fully compared with
Pteromys. Two groups are recognized, the volans group, evidently rather
smaller forms from eastern U.S.A. and Mexico, with the ventral surface lighter;
and the sabrimis group, from Labrador, across much of Canada, to Alaska, and
in the western U.S.A.; usually larger, and with ventral surface darker.
Forms seen : volaiis, sabrimis, alpimis.
List of N.^med Forms
roliiris Group
1. GL.^UCOMYS VOLANS VOLANS, Linnaeus
1758. Syst. Nat., loth Ed., vol. i, p. 64.
Virginia.
Synonym: volucella. True, 1885, Proc. U.S. Nat. Mus. VII, p. 596.
silus, Bangs, 1S96, Proc. Biol. Soc. Washington, X, p. 163.
Katis Mountain, Greenbrier County, West Virginia.
nebrascensis, Swenk, 191 5, Univ. Nebraska Studies, p. 15,
pi. 151.
2. GLAUCOMYS VOLANS S.A.TUR.-\TUS, Howell
1915. Proc. Biol. Soc. Washington, XXVIII, p. no.
Dothan, Henry County, Alabama.
3. GLAUCOMYS VOLANS QUERCETI, Bangs
i8q6. Proc. Biol. Soc. Washington, X, p. 166.
Citronelle, Citrus County, Florida.
4. GLAICOMYS VOLANS TEXENSIS, Howell
1915. Proc. Biol. Soc. Washington, XXVIII, p. no.
Sour Lake, Hardin County, Texas.
5. GLAUCOMYS VOLANS GOLDMANI, Nelson
1904. Proc. Biol. Soc. Washington, X. p. 148.
Twenty miles south-east of Teopisca, Chiapas, Mexico.
6. GLAUCOMYS VOLANS HERKERANUS, Goldman
1936. Journ. Washington .•\cad. Sci., XXVI, p. 463.
Mountains of \'era Cruz, Mexico.
zqb GLAUCOMYS
7. C;LAI COMYS VOLANS MADRENSIS, Cioldman
IQ36. Journ. Washington Acad. Sci. XXVI, p. 463. ,
Sierra Madre, Chihuahua, Mexico.
S. GLAL'COMYS VOLANS UNDERWOODl, Goodwin
1936. Amer. Mus. Nov., no. 89S, p. i.
Zambrano, Tegucigalpa, Honduras (a village halt-way between Teguci-
galpa and Comayagua).
sahn'niis Group
9. GLALCOMYS SABRINUS SABRINU.S, Shaw
iSoi. Gen. Zool. 2, p. 157.
.Severn River, Keewatin, Canada.
.Synonym: hudsonius. True, Proc. U.S. Nat. Mus. VII, 1885, p. 596.
10. GLAUCOMYS SABRINUS MAKKOVIKENSIS, Sornborger
1900. Ottawa Naturalist, XIV, p. 48.
Makkovik, Labrador.
11. GLAUCOMYS SABRINUS MACROTIS, Mearns
1899. Proc. U.S. Nat. Mus. XXI, p. 353.
Catskill Mountains, Green County, New York (Hunter Mountain),
12. GL.AUCOMYS SABRINUS CANESCENS, Howell
191 5. Proc. Biol. Soc. Washington, XXVIII, p. in.
Portage la Prairie, Manitoba, Canada.
13. GLAUCOMYS SABRINUS BANGSI, Rhoads
1S97. Proc. .■Xcad. Nat. Sci. Philadelphia, p. 321 (footnote).
Idaho County, Idaho.
14. GL.^UCOMYS SABRINUS ALPINUS, Richardson
1S28. Zool. Journ. 3, p. 519.
Jasper House, .Alberta, Canada.
15. GLAUCOMYS SABRINUS YUKONENSIS, Osgood
1900. North .•\nier. Fauna, no. 19, p. 25.
Camp Davidson, Y'ukon Ri\er, near ..\laska-Canada boundary, "\'ukon,
Canada.
lb. GLAUCOMYS SABRINUS ZAPHAEUS, Osgood
1905. Proc. Biol. Soc. Washington, XVIII, p. 133.
Cleveland Peninsula (Helm Bay), S.-E. Alaska.
17. GLAUCOMYS SABRINUS OREGONENSIS, Bachman
1839. Journ. .^cad. Nat. Sci. Philadelphia, VIII, p. loi.
Columbia County, Oregon (probably near St. Helen).
iS. GLALCOMYS SABRINUS COLUMBIENSIS, Howell
1915. Proc. Biol. Soc. Washington, XXVIII, p. in.
Okanagan, British Columbia.
19. GL.'^UCOMYS SABRINUS FULIGINOSUS, Rhoads
1S97. Proc. Acad. Nat. Sci. Philadelphia, p. 321.
Cascade Mountains, near Martin Station, Kittitas County, Washington.
20. GLAUCOMYS SABRINUS LATIPES, Hr.weli
1915. Proc. Biol. Soc. Washington, XXVIII, p. 112.
Glacier. British Columbia.
GLAUCOMYS— EOGLAUCOMYS 297
21. GLAUCOMYS SABRINUS OLYMPICUS, Klliot
1899. Field Columb. Mus. Publ. 30. zool. ser., vol. i, p. 225.
Happy Lake, Challam County, Washington.
22. (iLAUCO.MY.S SABRIM S BILL.^TUS, Howell
1915. Proc. liiol. Soc. Washington, XXVIII, p. 113.
Sawtooth Lake, east base of Sawtooth Mountains, Idaho.
23. GLAUCOMY.S SABRINLS KLAM.-XTHENSIS, Merriam
1897. Proc. Biol. Soc. Washington, XI, p. 225.
Fort Klamath, Klamath County, Oregon.
24. GLAUCOMY.S SABRINUS ILAVIVKNTRIS, Howell
1915. Proc. Biol. Soc. Washington, XXVIII, p. 112.
Bear Creek, Trinity County, California.
25. GLAUCOMYS SABRINUS LASCIVUS, Bangs
1899. Proc. New England Zool. Club, I, p. 69.
Tallac, El Dorado County, California.
26. GLAUCOMYS SABRINUS CALIFORNICUS, Rhoads
1897. Proc. Acad. Xat. Sci. Philadelphia, p. 323.
San Bernardino Mountains, San Bernardino Co., California.
27. GLAUCOMYS SABRINUS STKPHENSI, Merriam
1900. Proc. Biol. Soc. Washington, XIII, p. 151.
Sherwood, Mendocino County, California.
28. GLAUCO.MYS SABRINUS GRISEIFRONS, Howell
1934. Joum. Mamm. Baltimore, 15, p. 64.
Lake Bay, Prince of Wales Island, Alaska.
11). GL.^UCOMYS SABRINUS LUCIFUGUS, Hall
1934. Occ. Papers Mus. Zool. Univ. Michigan, 296, p. i.
Summit County, Utah; 12 miles east of Kamas.
30. GLAUCOMYS SABRINUS FUSCUS, Miller
1936. Proc. Biol. Soc. Washington, XLIX, p. 143.
Cranberry Glades, Pocahontas County, West Virginia.
Genus 8. EOGLAUCOMYS, Howell
1915. EOGLAUCOMYS, Howell, Proc. Biol. Soc. Washington, XXVIII, p. 109.
Type Species. — Sciuropterus fimhriatus, Gray.
R.\NGE. — Palaearctic; Afghanistan and Kashmir, Punjab.
Number of Forms. — Two.
Characters. — This species was originally included in Glaucomxs by
Thomas. It differs from Glaucomys, as well as by the
characters such as the depressed frontals and much larger postorbital processes
due to the greater size of the animal, in that P. 3 is divided into two cusps and
that the metatarsal pad is present on the hindfoot. The zygomatic plate is much
more strongly tilted upwards, and tends to approach Pteromvs in this respect.
The upper and lower cheekteeth are much like G/aucomvs, differing from
Hyhpetes in being relatively simpler. Ears longer and more pointed than
Glaucomxs. Mammae 8.
29S EOGLAUCOMYS— HYLOPETES
It differs from Pteiomys in the fact that the inner sides of the upper molars
have only the one main cusp; the second main ridge of P. 4, M.i, 1M.2 has no
portion of it isolated as a cusp in adult; ]\1.3 has, as is usual, only one main
ridge; there is no tendency for the central depression in the lower molars,
particularly iM.3, to become reduced; 1X1.3 louver is not specially lengthened;
the zygomatic plate is less extreme; and the palatal foramina are short (normal
for the group).
Forms seen : Jiinbrldiiis, haberi.
List of Named Forms
i. eoglaucomys fimbriatu8 fimbriatus, gray
1S37. Ann. Mag. Nat. Hist. I, p. 584.
Himalayas: Simla.
2. EOGLAUCOMYS F1MBRL\TUS BARERI, Blyth
1S47. Journ. Asiat. Soc. Bengal, XVI, p. 866.
Mountain district of Nijrow, Kohistan, Afghanistan.
Genus 9. HYLOPETES, I'homas
iqoS. HvLOPFTts, Thomas, Ann. Mag. Nat. Hist. 8, I, p. 6.
Type Species. — Sciuropteius everetti, Thomas.
R.ANGE. — Indo-JMalayan; Sikkim, Nepal, Yunnan; Burma, Tenasserim,
Laos; Malay Peninsula, Sumatra, Java, Borneo, and adjacent
islands; Philippine Islands.
Number of P'orms. — Seventeen.
Ch.vr.acters. — Not essentially different from Euglaiicoiiiys but cheekteeth
with traces in both upper and lower series of a more complex
pattern, and characterized by the usual presence of several small pits and
depressions in addition to the usual Sciurine ridges. The zygomatic plate is
primitive, of Beloiiiys type, not high. In large forms, as tilhoia'ger, the post-
orbital process stands out well, the frontals are depressed; in small species, as
spddiceiis, the frontals are flatter, the postorbital process short. The infraorbital
foramen may be rather well open ; in mirantiacus, the portion of the zygomatic
plate behind it is much narrowed. Upper cheekteeth with the essential pattern
of Eoglaiicomvs, but with the ridges often with small depressions traceable in
them, and the joins of the three original inner cusps are sometimes traceable in
the inner side of the teeth. The difference between this genus and Eog/aiicoiiivs
in dental characters is comparable to that between Sciuriis and CdUosciunis.
Lower cheekteeth with M.3 elongated, and P. 4 rather the smallest tooth; more
complicated as a rule than in Eaghiucoinxs, and with a short well-marked fold
present in front of the anteroexternal cusp except in much worn teeth, and
with many small faint pits and lines present, these more clearly marked in the
larger species. Mammae 0. Tail broad, feather-shaped. The bullae are normal.
I'his genus was originally proposed as a subgenus ot PteitJiiivs, from which
HYLOPETES 299
it is unquestionably distinct. It might be more correct to refer this genus, with
Eoglaucomys, to the genus Glaucomys; but for the present I retain all named
genera in this group.
Forms seen : alboniger, aurtmtiucus, helune, everetti, harrisoni, leonardi, laotum,
nigripes, pltitviirus, prohiis, phayrei, sai^ittii, spadiceus.
'I'his genus divides into two well-marked groups on size characters, the
much larger albonii^er group (head and body length over 200), containing
alboniger from Nepal, Yunnan, Burma, and nigripes from the Philippines; and
the smaller sagitta group, containing all other forms. The size in the second
group is rather variable, but seems to grade down from the largest to the
smallest; approximate head and body measurements of the main species are:
phayrei, about 170; sagitta, about 150; awantiacus, about 140; helone, about
135; spadiceus, about 126; and platyurus about 100. H. amoenus, not seen, is
165. H. everetti is about the same size as phayrei, and rather more brightly
coloured than the majoritv of the remainder.
List of Named Forms
sagitta Group
1. HYLOPETES PL.\TYURLS, Jentink
1S90. Notes Leyden Mus., XII, p. 147, pi. vii, figs. 7, 8.
Deli, N.-E. Sumatra.
2. HYLOPETES SP.ADICEUS, Blyth
1847. Joum. Asiat. Soc. Bengal, XVI, p. 867.
Arakan.
3. HYLOPETES BELONE, Thomas
1908. Ann. Mag. Nat. Hist. 8, XI, p. 305.
Pulau Terutau, Malacca.
4. HYLOPETES S.AGITTA, Linnaeus
1766. .Syst. Nat. 12th ed. i, p. 88.
Java.
5. HYLOPETES LEPIDUS, Horsfield
1824. Zool. Res. Java, p. 173, pi.
Java.
(A synonym of sagitta according to Thomas & Wroughton, 1909,
Proc. Zool. Soc. London, p. 387.)
6. HYLOPETES H.\RRISONI H.\RRISONI, Stone
1900. Proc. Acad. Nat. Sci. Philadelphia, XLII, p. 462.
Menbuang River, .Sarawak, Borneo.
7. HYLOPETES HARRISON] CAROLI, Gyldenstolpe
1919. Stockholm Vet. .-Vkad. Handl. 60, no. 6, p. 29.
East Borneo.
8. HYLOPETES AURANTIACUS, Wagner
1841. Munch. Gel. .Anz., XII, p. 438.
Banka Island, off Sumatra.
9. HYLOPETES AMOENUS, Milkr
1907. Proc. U.S. Nat. .Mus., XXXI, p. 264.
Kundur Island, Rhio .Archipelago, Malaya.
300 HYLOPETES— PETINOMYS
lo. HYLOPETES PHAYREI PHAYREI, BIyth
1S51). Journ. Asiat. Soc. Bengal, XXVIII, p. 27S.
Rangoon. liurnia.
.1 HYLOl'lCTES PHAYREI PK(JBL'S. Thomas
1914. Journ. Bombay Nat. Hist. Soc, XXIII, i, p. 2S.
Mount Popa, Burma.
12. HYLOPETES PHAYREI LAOTUM, Thomas
11)14. Journ. Bombay Nat. Hist, Soc, XXIII, 1, p. 28.
Laos Mountains.
13. HYLOPETES EVERICTTI, Thomas
1895. Nov. Zool. II, p. 27.
Bunguran Island, Natunas.
(i/hoiil<;er Group
14. HYLOPETES ALBONIGER, Hodgson
1836. Journ. .■\siat. Soc. Bengal, V, p. 231.
Nepal.
Synonym: tiinibiilli. Gray, 1837, I'roc Zool. Soc. London, p. 68.
15. HYLOPI-TES LEONARDI, Thomas
1921. Journ. Bombay Nat. Hist. Soc, XXVII, 3, p. 501.
Kachin Province, North Burma.
i(.. HYLOPETES NIGRIPES MGRIPHS, Thomas
1893. Ann. Mag. Nat. Hist. 6, XII, p. 30.
Fuerta Princcsa, Palawan, Philippine Islands.
17. HYLOPETES NIGRIPES ELASSODONTUS, Osgood
1918. Proc Biol. .Soc. Washington, XXXI, p. i.
Bancalan Island, Philippine Islands.
(k-nus 10. PETINOMYS, Thomas
1908. Petino.mys, Thomas, .^nn. Mag. Nat. Hist. 8, I, p. 6.
Type Species. — Sciuroptenis liigens, I'homas.
Range. — Indo-AIalayan; Ceylon and South India; Hainan; Te-nasserim;
Malacca, Sumatra, Java, Borneo; Basilan Island (Philippines).
Number of Forms. — Thirteen.
Characters. — Like Hvlopetes, hut the bullae, although large, are flattened,
described bv Thomas as "fairly large horizontally, but
peculiarly low and flattened, scarcely rising above the general level ot the base
of the skull, their substance unusually thick and opaque." As in Hvlopetes, this
genus includes some large forms, as fuscocapillus, with depressed frontals and
more powerful postorbital processes, and some very small forms, as setosiis, with
less modified skulls. In the larger species the parietal ridges are quite well
developed.
Cheekteeth like Hvlopetes, sometimes tending to be a little more comple.x.
Zygomatic plate generally low and primitive, a little less so than is usual in the
fiiseociipilliis group. Mammae 4 or 6.
PETINOMYS 301
Though the peculiar flattening of ihc buiku- is less strongly marked in
fiiscocapillus than the others, it seems reasonable to regard this group as a genus.
I'our well marked groups have been examined:
the FUSCOCAPILLUS group of South India, large, head and body length about
296 mm., bullae not quite typical of the genus (hindfoot about 52);
the ii.\gi:ni group, about as large; bullae typical of the genus; including
luii^eni and liigens, from Sumatra and adjacent islands, and (?) crinitiis
(head and body 310, not seen), of the Philippines. The head and body
measurement of higens is 230, of hageni, 313 mm. (Jentink) ;
the GENiB.XRBis group: moderate-sized forms, hindfoot about 30; Borneo,
Java, Malacca; the Hainan species (not seen), head and body 172 mm. is
provisionally included here;
the SETOsiis group: pygmy forms; hindfoot 20-24, head and body 120 in
phipsoni, probably 100 or less in setosus; includes /)/»/)io«2 of Tenasserim,
setosus of Sumatra, and vordermanni of Billiton (head and body 100).
Forms seen : horneoensis, fiiscocapillus, gembarbis, hageni, laxardi, Itigens,
malaccaniis, phipsoni, setosiis.
List of Named Forms
setosus Group
1. PETINOMYS SETOSUS, Temminck & Schlegel
1847, Fauna Japon, Mamm., p. 49.
Padang, Sumatra.
2. PETINOMYS PHIPSONI, Thomas
1916. Joum. Bombay Nat. Hist. Soc, XXIV, 3, p. 421.
Tenasserim Town, Tenasserim.
3. PETINOMYS VORDERM.A.NNI, Jentink
1890. Notes Leyden Mus,, XII, p. 150, pi. vii, figs. 13, 14.
Billiton Island, off Sumatra.
genibarbis Group
4. PETINOMYS GENIB..\RBIS GENIB.A.RBIS, Horsfield
1824. Zool. Res. Java (description and plate).
Eastern Java.
5. PETINOMYS GENIB.^RBIS BORNEOENSIS, Thomas
1908. Ann. Mag. Nat. Hist. 8, II, p. 304.
Bakong River, East Sarawak, Borneo.
6. PETINOMYS GENIH.\RBIS MAL.i\CCANUS, Thomas
1908. .\nn. Mag. Nat. Hist. 8, II, p. 304.
Malacca.
7. PETINOMYS KLECTILIS, G. .M. .Allen
1925. Amer. Mus. Nov. 163, p. 16.
Nam Fong, Hainan.
302 PETINOMYS— PETAURILLUS
luigoii Group
S. PKTINOMYS LL"GEXS, Thomas
1S94. Ann. Mus. Stor. Nat. Genova, XIV, p. 666.
Si Oban, Sipora Island, \V. Sumatra.
>). PETINOMYS MAEREN.S, Miller
IQ03. Smiths. Misc. Coll., XLV, p. 26.
North Pagi Island, west of Sumatra.
10. PETIXO.MYS HAGENI. Jentink
iSSS. Notes Leydcn Mus., XI, p. 26.
Deli, Sumatra.
11. PETINOMYS CRINITUS. Hollister
igii. Proc. Biol. Soc. Washington, XXIV, p. 1S5.
Basilan Island, Philippines.
fuscocapillus Group
12. PETINOMYS FUSCOCAPILLUS, Jerdon (in Blyth)
1847. Journ. Asiat. Soc. Bengal, XVI, p. S67.
South India.
13. PETINOMYS LAY.ARDI, Kelaart
1S50. Joum. Roy. Asiat. Soc. Ceylon, XI, p. 328.
Ceylon.
Genus 11. PETAURILLUS, Thomas
190S. Pet.-wrillis, Thomas, .-Xnn. Mag. Nat. Hist. 8, I, p. 3.
Type Species. — Sciuropterus liosei, Thomas.
Range. — Known from Selangor and Borneo.
Number of Forms. — Three.
Characters. — Pygmy Flying-squirrels, rather sharply differentiated from
the other genera by the simpler cheekteeth and the relative
size of the upper teeth. P. 4 is noticeably smaller than INI.i ; but P. 3 is quite
well developed, so that the three anterior teeth decrease evenly in size from ALi
forwards. The cheekteeth with low ridges, the pattern not distinct, though
evidently much as in norma! Sciuridae. P. 4 lower noticeably reduced; cusps
of lower teeth low. Zygomatic plate a little higher and broader than is usual.
Bullae large. Size very small.
The forms of this genus are not well known.
Forms seen : hosei, emiliae, kinloclii.
List of Named Forms
I. PETAURILLUS HOSEI, Thomas
1900. .Ann. Mag. Nat. Hist. 7. V, p. 275.
Tovul River. liaram. Sarawak, Borneo.
PETAURILLUS— lOMYS 303
2. PETAURILLUS EMILIAE, Thomas
1908. Ann. Mag. Xat. Hist. 8, I, p. 8.
Uaram, Sarawak, Borneo.
3. PETALRILLUS KLNLOCHI, Robinson & Kloss
1911. Journ. Fed. Malay States Mus., IV, p. 171.
Kapar, Sclangor, Malay Peninsula.
Genus 12. lOMYS, Thomas
1908. loMYS, Thomas, Ann. Mag. Xat. Hist. 8, I, p. i.
Type Species. — Pteromys horsfieldi, Waterhouse.
Range. — Malacca, Sumatra, Borneo, Java.
Number of Forms. — Five.
Characters. — Skull, including the low zygomatic plate, essentially as
Belomys. Cheekteeth 4 , in appearance square, and differing
noticeably from all other members of the group. The two main ridges of the
upper cheekteeth rise inwardly as well as outwardly into two cusps, so that each
tooth has a well-marked cusp at each corner. The depressions in front of the
two main ridges are well marked, but the posterior depression is obsolete. No
marked discrepancy in size between any of the upper cheekteeth. P. 3 absent.
M.3 with only one main ridge, but the two inner cusps are present. P. 4 with
its small foremost cusp placed in front of the tooth, nearly centrally. Lower
cheekteeth with four well-marked cusps on each tooth, the anterointernal not
or little higher than the others, which is a ver\' rare feature in the family. The
central depression is much narrowed, and appears as a re-entrant fold; opposite
to it is a narrow external fold which is separated from it by a narrow ridge
joining the two outer cusps. This tooth formation rather suggests the specialized
lower molars of Funisciiirus among the Sciurus group.
Externally with no special features. Ear rather large.
Forms seen : thomsoni, horsfieldi, davisoni.
The named forms are all regarded as races of the tvpe by Robinson & Kloss,
1918.
List of N.\med Forms
1. lOMYS HORSFIELDI HORSFIELDI, Waterhouse
1837. Proc. Zool. Soc. London, p. 87.
Java (or Sumatra).
2. lOMYS HORSFIELDI D.AVISONI, Thomas
1886. Proc. Zool. Soc. London, p. 74, pi. vi.
Malacca.
.I. lOMYS HORSFIELDI THO.MSONI, Thomas
1900. .Ann. Mag. Nat. Hist. 7, V, p. 275.
Bakong River, Baram, Sarawak, Borneo.
4. lOMYS HORSFIELDI LEPIDUS, Lyon
191 1. Proc. U.S. Xat. Mus.. XL, p. 78.
Kendawangan River, S.-W. Borneo.
304 EUPETAURUS
5. UJMYS HOKSFIKLDl SIPDRA, Lhascri & Kloss
192S. Proc. Zool. Soc. London. 1927, p. Sig.
Sipora Island. Sumatra.
Genus 13. EUPETAURUS, Thomas
18S8. EiPETAURUS. Thomas. Journ. Asiat. Soc. Bengal. LVII. p. 25b.
Type Species. — Eiipctnuii/s litwiiiis, Thomas.
R.\NGE. — Kashmir.
Number of Forms. — One.
Ch.\r.\cters. — "Skull distinct from that of ' Pteromys' (^-Petaurista) by its
longer trumpet-shaped muzzle, more marked supraorbital
notches, longer anterior palatine foramina, and shorter bony palate. Teeth
strikingly contrasted with those of any of the other Sciuridae by being hypsodont
instead of brachyodont, while their essential pattern remains unchanged. Thus,
while the crown ot each tooth is enormously lengthened vertically, the grooves
ordinarily present on the grinding surface of the molars of ' Pteromvs' are
reproduced as deep vertical foldings of the enamel, which when seen in the
natural section produced by wear give the teeth very much the general appear-
ance of those of many of the Hystricomorpha . . . the teeth also apart from their
hypsodont structure, are distinguishable by their very large proportional size,
by being set more obliquely than is the case in other Squirrels, and by pre-
senting ... a sharp posterointernal angle, markedly diflerent from the evenly
convex internal border of the teeth of 'Pteromys.' The implantation of the large
upper premolar is also peculiar, in that of the three distinct roots it has in the
allied forms, the anteroexternal and the internal have coalesced into a single
broad flat root. . . ." (Thomas). As figured by Thomas, P. 4 is rather longer
than M.i in the upper series.
This genus is represented at the British Museum by a few- skins, but as yet
by no skull. Tail very thickly haired throughout; it appears to be of the
Pteruiiivs rather than the Petaiiiista type, and there is evidently no well-marked
interfcmoral membrane. Ear low. Whole body covered in excessively thick
soft woolly fur, that of the ventral surface being lighter than that of the dorsal.
Even the hindfocit is, except for the pads, heavily haired. Size large.
Forms seen: ciiieieiis (skins).
List of N.\meij For.ms
I. EUPETALRUS CIN'EREUS. Thomas
1888. Journ. ."Xsiat. Soc. Bengal, LVII, p. 258, pis. xxii, xxiii.
Gilgit Valley, Himalayas. Kashmir.
The Sciiirus Group
Geographical Disikibution. — As in the family Sciuridae.
Characters. — This group differs from the Pteromys group in the invariable
absence of flying-membrane. The cheekteeth are never so
excessively complex as in certain Flying-squirrels as Belo?iiys, Tro;;ii[>terus, etc.;
SCIURUS GROUP 30s
but on the other hand some Flying-squirrels as Glaucomys are quite as simplified
dentally as anv of the present group; in the Sciurits group, usually the zygo-
matic plate is higher and more tilted upwards than in the more primitive
members of the Pteromys group.
Key to Genera of Sciurus Group
The rostrum extremely elongated.
Upper incisors much reduced ; cheekteeth tending to wear down to
the roots in adult; infraorbital foramen barely forming a
canal; claws not enlarged. Rhinosciurus
Upper incisors strong, not reduced; cheekteeth evidently without
peculiarities; infraorbital foramen forming a long canal;
claws much enlarged. Hyoscicrus
The rostrum not extremely elongated. (Upper incisors not abnormally
reduced; the cheekteeth never tending to wear down to the roots
during life.)
The upper incisors greatly strengthened, either much thickened
anteroposteriorly, their surfaces with many parallel grooves, or
much broadened.
Upper and lower incisors much thickened anteroposteriorly, with
many parallel grooves on their surfaces. Premolars con-
siderably reduced; toothrows reduced, and cheekteeth
simplified in pattern. Rostrum lengthened. Rheithrosciurus
Upper and lower incisors much broadened, the lower part of the
upper teeth and the upper part of the lower teeth diverging
from each other, their anterior surfaces without grooves.
Cheekteeth (so far as ascertainable) normal. Rostrum short.
Glyphotes
The upper incisors without extreme abnormalities; (in genera in
which these teeth are becoming thickened, their anterior
surfaces are without grooves).
Skull abnormal, the orbit circular, placed far backwards; postor-
bital process much reduced or vestigial, situated above level
of posterior zygomatic root. Lachrymal over middle or
hinder part of toothrow. Zygomatic plate appearing nearlv
vertical.
Infraorbital foramen forming no canal, the portion of the zvgo-
matic plate behind it exceedingly reduced, situated in
front of tooth-row. Ectopterygoid absent. P.4 (upper
series) much reduced. (Cranial characters as indicated
above carried to extreme degree; size smallest of
family.) Myoscu'RUS
20 — Living Rodents — I
3o6 SCIURUS GROUP
Intraorbital foramen forming a canal, the portion of the zygo-
matic plate behind it normal, situated over hinder part
oftoothrow. Ectopterygoid present. P. 4 (upper series)
not specially reduced.
Cranial characters as indicated above carried to extreme
degree; postorbital process vestigial. M.3 more
reduced than is normal in the family. Palate usually
narrowed. Xan'nosciurus
Cranial characters as indicated above not or less extremely
developed. Postorbital process less vestigial. AI.3
not reduced. Palate not narrowed. Sciurii.lus
Skull less abnormal; orbit not circular, not placed unusually far
backwards; postorbital process usually situated consider-
ably in front of posterior zygomatic root (excepting the
genus Microsciiinis); lachrymal usually over or in front of
part of toothrow; zygomatic plate strongly oblique. ^
Cheekteeth simplified, losing all traces of normal pattern
practically from birth. L.\RISCUS
Cheekteeth not simplified, not losing all traces of normal pattern
till adult or usually late in life.
Externally modified for terrestrial life; D.3 in the manus
always (so far as seen, possibly excepting Atlan-
toxerus) longer than D.4. (Cheekteeth in progres-
sive species becoming stronglv hypsodont; tail
shorter than head and body, often considerably
reduced ; infraorbital foramen forming a canal.)
Lachrymal considerably enlarged. Palate extending
conspicuously behind toothrows. Bullae enlarged,
well inflated.
Tail short, little longer than hindtoot; claws of fore-
and hindfeet excessively thickened and de-
veloped; bullae not evenly rounded.
Spermophilopsis
Tail relatively long, sometimes approaching head
and body length; claws of fore- and hindfeet
not excessively thickened, less developed;
bullae evenly rounded. (Fur always bristly,
compare Atlantoxerus.) Xerus
Lachrymal not specially enlarged.
Palate extending conspicuously behind toothrows.
^ I'ossibic exceptions to some of these characters may be shown in the Celebes Callosciurus
leucorrrus, very fe\% skulls ot which have been examined.
SCIURUS GROUP 307
(Bullae large, evenly rounded; fur not
bristly, compare Xerus.) Atlantoxerus
Palate not extending conspicuously behind tooth-
rows. (Usually, upper cheekteeth with ten-
dency towards constriction on inner side, so
that they become roughly three-sided instead
of more or less rounded or four-sided, as is
normal; this constant in Cynomys (strongly
developed), Marmota (moderately developed),
and a large portion of Citelhis (strongly
developed).
Toothrows markedly convergent posteriorly.
Dentition extremely hea\-y. Skull with
prominent ridges for muscle attachment.
Mandible with angular portion strongly
inflected. Pollex not vestigial. Cynomys
Toothrows not or scarcely convergent pos-
teriorly. Dentition rarely or not ex-
tremely hea\-y. Pollex vestigial.
Skull massive, with heavy prominent postor-
bital processes, a strong sagittal crest
normally present. Ridges for muscle
attachment on skull prominent.
Mandible with angular portion norm-
ally less inflected. M.armot.a
Skull lighter, with moderate or weak postor-
bital processes, a sagittal crest most
often not developed. Ridges on skull
for muscular attachment never exces-
sive. Mandible with angular portion
normally strongly inflected. Citellus
E.xternally semi-terrestrial or arboreal in external charac-
ters; D.3 in manus never constantly longer than D.4
(except in the genera Tamias and Sciurotamias there
is a very general tendency for D.4 to be longer than
D.3). (Palate never produced conspicuously be-
hind toothrows; upper cheekteeth with no tendency
towards constriction on inner side.)
Infraorbital foramen forming no canal, and normallv
relatively large, round and well open, usuallv at
maximum for the subfamily. (The position of the
genus EpLxerus must be regarded as provisional.)
3o8 SCIURUS GROUP
D.3 and D.4 in manus normally approximately
equal in length. Skull more or less flat, and
with reduced postorbital processes. (Ventral
surface of body normally furred; cheek-
pouches present; tail not conspicuously
bushy.) Incisors not specially thickened. T.vmias
D.4 in the manus longer than D.3. Skull not
flattened, the postorbital processes not re-
duced. (Tail conspicuously bushy.) Incisors
considerably thickened.
Ventral surface of body normally haired. Cheek-
teeth (of all specimens examined) \yith
clear and well-marked ridges and depres-
sions (compare Proloxeriis). Infraorbital
foramen well open (compare Epixenis.)
Myrsilus
Ventral surface of body poorly haired, often
almost naked.
I'oothrows considerably reduced. Infra-
orbital foramen narrower, less well
open. Epixerus
Toothrows not specially reduced. Infra-
orbital foramen large, well open.
(Cheekteeth usually without clearly
marked ridges and depressions (com-
pare Myrsilus).) Protox[:ri's^
Infraorbital foramen less open, always forming at least
a short canal.
The lower cheekteeth specialized, becoming trans-
yersely ridged, as in the upper series, the
central depression characteristic of normal
genera much reduced and appearing as a
re-entrant fold.
Zygomatic plax norinal, the ridge on its upper
border extending beyond leyel of the
infraorbital foramen. Upper cheekteeth
simplifying early in life, in the adult
usually with only one clear re-entrant fold
retained; the central depression of the
lower molars often becoming isolated.
Menetes
^ With rfpresentati\'e material it may be that Protoxenis, Epixenis, and Myrsilus would be
better considered as all of the one genus Protoxerus only.
SCIURUS GROUP
Zygomatic plate shortened, the ridge on its upper
border stopping abruptly above the infra-
orbital foramen. Upper cheekteeth not
s.rnphfying early in life, usually in adult
JMth three clear re-entrant folds present
Lower teeth with the central depression
normally not isolated.
Lower cheekteeth with cusps obsolete, and
crowns almost completely flat. Funisciurus
Lower cheekteeth with cusps strongly
marked, the crowns not becoming flat.
The lower cheekteeth much less specialized, or not
so not tending to become transversely
ndged as m the upper series ^
Skull flattened, little depressed posteriorly
narrow in general appearance, and with
strongly reduced postorbital process
Avgomat.c plate not strongly tilted up-'
^vards. D.3 and D.4 in manus usually
approximately equal in length. (Infra-
orbital foramen barely forming a^anal
only a httle less open than in tLw' )
ci., 11 . „ SCIUROTAMI.AS
Skull not specially flattened, usually strongly de-
pressed posteriorly, and with postorbital
process never much reduced except in
vm- small species. Zygomatic plate well
t.ted upwards. (Infraorbital '^foramen
clearly forming a canal.)
Postorbital process extremely thick and pro-
m.nent; cheekteeth " with very low
cusps, the pattern almost always in-
distinct; feet much specialized for
arboreal life, the inner side of forefoot
with conspicuous expansion (evidently
taking the functional place of pollex).
p . ,. , Ratufa
Postorbital process usually not e.xtremely
prominent; cheekteeth with moderate
or high cusps, the pattern almost
always clear and definite at least at
some stage of life; feet less con-
spicuously specialized for arboreal life
3IO SCIURUS GROUP
the expansion on the inner side of
forefoot absent or less conspicuous.
Postorbital process tending to be situated
nearly or exactly over the posterior
zygomatic root. (Upper incisors
pro-odont.) Microsciurus
Postorbital process situated clearly in
front of posterior zygomatic root.
(The remainder of the genera are not at all times distinguishable from each
other on cranial and dental characters alone.)
Baculum, so far as known, suppressed
or vestigial.
Zygomatic plate either slanting up-
wards or forwards, with
strong ridge on superior
border, and with prominent
masseterknob present under
the infraorbital foramen; or
with the ridge not approach-
ing the superior border of
rostrum, and stopping ab-
ruptly above the infraorbital
foramen. Lower cheek-
teeth with a narrow trans-
verse valley extending from
first outer main cusp to the
anterointernal cusp.
Heliosciurus
Zygomatic plate without abnorm-
alities. Lower cheekteeth
without well-marked nar-
row transverse valley ex-
tending from first outer
main cusp to anterointernal
cusp (so far as seen).
Tami.\sciurus
Baculum, so lar as known, retained.
(The characters of the genus
Syntheosciiirus in this respect
are not known.)
Rostrum progressively elongated
throughout e\ery species of
the genus, at its extreme
SCIURUS GROUP 311
development becoming ab-
normal. Oremomys
Rostrum never consistently
elongated throughout everj-
species of a genus, at ex-
treme development never
abnormal.
Coronoid process relatively
low; cusps of cheek-
teeth noticeably high,
and central depression
of lower cheekteeth often
tending to be relatively
smaller; zygomatic
plate usually slanting
upwards far forwards,
and rather prominently
ridged (M.3 lower
series not specially elon-
gated). FUNAMBULUS
Coronoid process in the
majority high, well
developed ; cusps of
cheekteeth usually less
noticeably high, and
central depression of
lower cheekteeth not
reduced normally; zy-
gomatic plate most often
not slanting upwards far
forwards, and not con-
spicuously ridged.
Upper cheekteeth with
small outer (third)
cusp usually, not al-
ways, absent or ob-
solete ; pattern of
cheekteeth usually
definite, clear, and
rather more com-
plex; M.3 lower
series normally tend-
ing to be noticeably
elongated. Callosciurus
312 SCIURUS GROUP— MYOSCIURUS
Upper cheekteeth with
small outer (third)
cusp retained or
traceable; pattern
of cheekteeth usually
comparatively in-
distinct; M.3 lower
series rarely or not
elongated.
Upper incisors not pro-
odont, plain. SciURi'S
Upper incisors pro-
odont, one-
grooved.
Syntheosciurus
The genera Callosciurus and Funamhidus are retained, it must be admitted,
more for convenience than because of the conviction that thev are of necessity
distinct generically from Sciurus, though Pocock transferred them on baculum
structure to three different subfamilies. Apart from this structure, they are
separable only on average characters; the same applies to Dremomvs, which
possesses intermediate species grading into Callosciurus to which it evidently
stands nearest. Comparing Funainbulus with Sciuiiis and Callosciurus, Pocock
writes, regarding the baculum of his " Funambulinae" : "It is when present
always a simple bone, without the spatulate expansion at the apex seen in the
Sciurinae and without the accessory blade of the Callosciurinae."
Section A. N.\nnosciurus Section: Pygmy Squirrels with highly abnormal
cranial characters; the Nannosciurinae of Miller & Gidley, and Forsyth
Major.
Genus i. MYOSCIURUS, Thomas
igog. MYOSCIURUS. Thomas, Ann. Mag. Nat. Hist., S, III, pp. 469. 474
Type Species. — Sciurus pumilio, Le Conte.
Range. — West Africa; Cameroons, Gaboon.
Number of Forms. — One.
Ch.aracters. — Skull with extremely broad frontals; postorbital process
vestigial, situated over posterior zygomatic root. Zygomatic
plate almost vertical, slanting upwards over or behind toothrow. Infraorbital
foramen immediately in front of toothrow, the portion of the zygomatic plate
behind it abnormallv narrowed, also placed in front of toothrow. No ectoptery-
goid. Palate straight, considerably narrowed. Jugal broad, as in allied genera.
Incisors pro-odont. Cheekteeth J. According to Forsyth Major, writing of
this genus and Naniiosciuriis, "The pattern of the crown differs trom that found
MYOSCIURUS— NANNOSCIURUS 313
in the Sciuromorpha generally in presenting only three complete transverse
crests in the upper molars instead of four, and three in the lower molars. The
third crest ... is very reduced in these py^my squirrels, sometimes not more
than a minute cusp. A further peculiarity of these molars is the large develop-
ment of the anterior transverse valley both of the superior and inferior molars
. . . sometimes almost equalling that of the posterior valley. This last, owing
to the partial suppression ot the third crest, occupies the area of the median as
well as that of the posterior transverse valley in the teeth of Sciuromorpha."
On this account he referred these genera to a separate subfamily ; but sometimes,
as in skull No. 9.10.2.36 at the British -Museum, the ordinary Sciurine ridges
(four) and depressions (three) may be traced in the main teeth. The toothrow
is reduced. Another peculiarity is that in this genus M.3 is turned over, and
faces outwards. Upper and lower premolars very reduced in size.
Size verv small indeed, head and bodv about 75 mm. Tail much narrowed,
shorter than head and body (about three-quarters this length or slightly more).
Digits as in normal Tree-squirrels.
This genus is undoubtedly the most aberrant of the section, as shown chiefly
by the abnormal infraorbital foramen, and also the lack of ectopterygoid and
the extremely small size.
Forms seen : pumilio.
List of N.wied Forms
(References and type localities of all members of Sciurus group by Mr. R. W.
Hayman.)
I. MYOSCIURUS PUMILIO, Le Conte
1857. Proc. Ac. Nat. Sci. Philadelphia, p. 11.
Gaboon.
Synonym: mimitus, du Chaillu, 1S61, Proc. Boston Soc. Nat. Hist., VII,
p. 366. Gaboon.
miniitidus, Hollister, 1921, Proc. Biol. Soc. Washington,
XXXIV, p. 135.
Genus 2. NANNOSCIURUS, Trouessart
1880. N.\NNoscn.T«us, Trouessart, le Naturaliste, p. 292.
Type Species. — Sciurus tnelanotis, Miiller & Schlegel.
Range. — Indo-Malayan ; Sumatra, Borneo, Java, and the Philippine Islands.
Number of Forms. — Fourteen.
Characters. — Like Mvosciurus but with the ectopter^'goid present, and
infraorbital foramen forming a short canal, the portion of the
zygomatic plate behind it less reduced, normal. Coronoid process, as in
Myosciuriis, much reduced. The cheekteeth are similar to those of Mvosciurus,
though the elements of the usual Sciurine pattern may be sometimes traced, as
in skulls No. 92. 11. 8. 6 and 10.4. 5. 113 at the British Museum. M.3 not facing
outwards, relatively small, more reduced than is usual in Sciurinae; P. 3 present;
P.4asa rule not specially reduced. P. 4 lower smaller than the other lower molars;
Fig. 91. Myosciurus pumilio, Le Conte.
B.M. No. 5-5-23-S, 3; '-< 3j-
Fig. 92. Myosciurus pumilio, Le Conte.
B.M. No. 5.5.23.5, o; 3*-
NANNOSCIURUS
315
cusps in lower teeth nearly obsolete, and the main central depression appears to
give way to a transverse ridge. Palate usually narrow.
E.xternally slightly larger than Myosciurus, or becoming so. Tail tending to be
narrow, shorter tlian head and body length. Arrangement of digits not abnormal.
Fig. 93. (a) Myosciurus pumilio, Le Conte. X2.
(6) SciURiLLUS PUSiLLUs PUSILLUS, Desmarest. X2.
(c) SciLTRiLLUS MURiNus MVRiNus, MuUer & Schlegel. X2.
{d) C.\LLOSCiURUS TENUIS sLT<DUS, Miller. ;■: 2.
Three groups are recognizable in this genus: melanotis group, paler, with
black ears and white face markings; exilis group, darker, without face markings;
and whiteheadi group, like the last but with ear-tufts present and conspicuous,
in some cases extremely long. (These absent in exilis group.)
Forms seen : borneanus, concinmis, exilis, melanotis, pulcher, retectus, samaricus,
whiteheadi.
List of N.^med Forms
exilis Group
I. NANNOSCIURUS EXILIS KXILIS, Miillcr & Schlegel
1838. Tijds. Natur. Ges., p. 148.
Batang Singalur, Sumatra.
3i6 XAXNOSCIURUS
2. NANNOSCIURl'S liXII.lS RICTHCTUS. Thomas
iqio. Ann. Mag. Nat. Hist. S, V, p. 387.
Banguey Island, North Borneo.
3. NANNOSCIURUS EXILIS SORDIDUS. Chascn & Kloss
1928. Joum. Malay Branch Roy. As. Soc, VI, pt. i, p. 44.
Long Tenielan, Middle East Borneo.
4. NANNOSCIURUS CONCINNUS, Thomas
18SS. Ann. Mag. Nat. Hist. 6, XI, p. 407.
Isabella, Basilan Island, Siilu group, Philippines. Considered a sub-
species of e.xilis by Robinson & Kloss, 1918.
5. N.^NNOSCIURUS SAMARICUS, Thomas
1S97. Minutes Zool. Soc. London, 15th June, p. I. 1S9S. Trans. Zool. Soc. London,
XIV, p. 389, pi. 30, fig. 2.
Samar, Philippine Islands.
6. NANNOSCIURUS SURRUTILUS, Hollistcr
1913. Proc. U.S. Nat. Mus. XLVI, p. 313.
Mount Bliss, Mindanao, Philippine Islands.
7. NANNOSCIURUS LUNCEFORDI, Taylor
1034. Philippine Land Mammals, p. 373.
Saub, Cotabato, Mindanao, Philippine Islands.
ivhiteheadi Group
8. NANNOSCIURUS WHITEHEADI, Thomas
1887. Ann. Mag. Nat. Hist. 5, XX, p. 127.
Mount Kina Balu, North Borneo.
mehinotis Group
y. NANNOSCIURUS MELANOTIS MELANOTIS, Miller & Schlege
1844. Temminck's Verhandelinger, Zoologie, pp. 87, 88, pi. xiv, fig. 4.
Java.
Synonym: sonanus, Waterhouse, 1S3S, Cat. Mamm., p. 46.
10. NANNOSCIURUS MELANOTIS SUMATRANUS, Lyon
1906. Proc. Biol. Soc. Washington, XIX, p. 53-
Tarussan Bay, West Sumatra.
.1. NANNOSCIURUS MELANOTIS PULCHI'R, Miller
1902. Proc. Acad. Nat. Sci. Philadelphia, p. 153.
Sinkep Island, near Sumatra.
12. NANNOSCIURUS MELANOTIS BANCANUS, Lyon
1906. Proc. Biol. Soc. Washington, XIX, p. 55.
Klabat Bay, Bangka Island, East Sumatra.
13. NANNOSCIURUS MELANOTIS BORNEANUS, Lyon
1906. Proc. Biol. Soc. Washington, XIX, p. 54.
Sanggan, West Borneo.
14. NANNOSCIURUS MELANOTIS PALLIDUS, Chasen & Kloss
1928. Journ. Malay Branch. Roy. As. Soc, VI, pt. i, p. 43.
Long Poehoes, Middle East Borneo.
SCIURILLUS 317
Genus 3. SCIURILLUS, Thomas
1914. SCIURILLUS, Thomas, Abstr. Proc. Zool. Soc. London, May 12th, p. 36; id,,
Proc. Zool. Soc. London, 1914, p. 416.
Type Species. — Sciurus piisillus, Desmarest.
Range. — Neotropical; Guianas, extending south to the Amazon. Indo-
jNIalayan; Celebes (murimis group provisionally included here).
Number of Forms. — Five.
Characters. — In cranial characters clearly a member of the Nannosciurus
section. Ectopterygoid present. Skull much like that of
Nannosciurus except that the palate is broad and normal, and the whole crania!
effect is a little less extreme owing presumably to the fact that the animals are
rather larger. Jugal in both specific groups included here very broad. Post-
orbital process less vestigial than in Nannosciurus. In the type species the
opening of the intraorbital foramen is carried upwards on front part of zygomatic
plate as a curved groove. P. 3 present. M.3 not reduced. Only much worn
teeth examined in the type species; the ridges not clear, obsolete, the cusps
low. P. 4 lower, somewhat reduced.
Externally (type species) rather larger than Nannosciurus, head and body
reaching 107 mm. Tail more normal, longer, about as long as head and body,
bushy. Digits not abnormal, arboreal type.
There are also at the British Museum three specimens from Celebes labelled
"Sciurus murinus." Whether these represent true murinus or not I have been
unable to find out, as I have not seen any description or reference to this species
which mentions cranial characters. But all cranial characters of the Nanno-
sciurinae as diagnosed by Miller & Gidley, except the fact that the middle of
orbit (like typical Sciurillus) is not behind the middle of the skull, are clearlv
present in these skulls from Celebes. The species is evidently a transitionarv
type between Nannosciurus section and Sciurus section, and is evidently the
Celebes representative of the former; "giant" representatives, if one can call
a Squirrel a giant, which must measure less than six inches in head and body
length. The dentition is about as in normal Squirrels apparently, but much
worn in the three examined; the proportions of the teeth agree with those of
Sciurillus rather than Nannosciurus, as do the main cranial characters. It is not
my intention to burden this Order with more generic names than can be avoided,
so I transfer this group provisionally to Sciurillus, though it mav be that later
the group will need a generic name. Should true murinus prove to belong to
Callosciurus in cranial characters, these skulls must represent a new and un-
described species, but one which I should not feel justified in leaving in Callo-
sciurus on cranial characters. It is interesting to note that Nannosciurus is not
known from Celebes. It is to be hoped that further material will come to hand.
The form evidens, which is described as near murinus, I provisionallv list here,
though 1 have not seen it.
Forms seen : pusillus, "laucinus, murimis.
3i8 SCIURILLUS
List of Named Forms
piisi/liis Group
I. SCIURILI.US lasiLLL'S PUSILIA'S, Desmarest
1822. Mammalogie, pt. 2, p. 337, pi. 77, fig. 2.
South America ; Cayenne.
2. SCIURILLUS PUSILLUS GL,'\UCINUS. Thomas
1914. .\nn. Mag. Nat. Hist. 8, XIII, p. 575.
Great Falls of Demerara River, British Guiana.
murinus Group
3. SCIURILLUS MURINUS MURINUS, iMuilcr & Schlegel
1S44. Vcrhandl. Zool., p. 87.
Celebes.
4. SCIURILLUS MURINUS NECOPINUS, Millir & Hollister
IQ2I. Proc. Biol. Sec. Washington, XXXIV, p. 98.
Goenoeng Lehio, Middle Celebes.
5. SCIURILLUS C-l EVIDENS, Millei & Hollister
1921. Proc. Biol. Soc. Washington, XXXIV, p. 99.
Puloh Lenibeh, N.-E. Celebes.
The infraorbital foramen of the murinus group is normal, without the above-
noted peculiarity of the pusiUus group.
Since the above was written, we have been fortunate enough to obtain three
more of these Celebes Pygmy Squirrels, through Mr. W. Frost. Their cranial
characters are precisely as in the skulls at present in the British Museum and
mentioned above. This indicates that at anv rate these skulls did come from
Celebes, and also apparently that a small Squirrel of this type is common there,
as Mr. Frost writes that Scjuirrels have not been easy to obtain, and these were
the first that came to hand ; and it strengthens my supposition that thev probably
are true murinus, and that the species should certainly not remain in the genus
Callosciurus. The dentition of the new skulls indicates that the pattern is
probably as in normal Squirrels; P. 3 is relatively well developed.
The head and bodv length is 130 mm.; the tail is shorter than this measure-
ment (average 70).
Note. — Since the above was written I have seen an important paper on the
genus Sciurillus (South American section) by Tate & Anthony, Amer. Mus.
Nov. 780, Feb. 14, 1935, notes on South American Mammalia, no. i, Sciurillus.
These authors state that the form kulilii, Gray, 1867, Ann. Mag. Nat. Hist. 3,
XX, p. 433, Pebas, Peru (see the above-mentioned paper, p. 10), is a race of
Sciurillus pusillus, and not a synonym of Sciurus aestuans, as listed here. I have
not seen kuhlii.
Section B. Sciurus Section : typical Tree-squirrels. In this section are placed
very many forms belonging to about eight genera, from the Hularctic,
South America, and the Indo-Malayan.
MICROSCIURUS 319
Except for Microsciurus and Ratufa, the genera are not clearly dis-
tinguishable from one another on cranial and dental characters. On
characters of the baculiim, some of them have been arranged in three
different subfamilies (Pocock); but other than the two genera noted
above all might quite easily be referred to a single genus Sciurus. The
African genus lleliosciurus, which I have placed in section D, is another
genus which is separable only on baculum characters from Sciurus or
its allies.
Genus 4. MICROSCIURUS, Allen
1895. Microsciurus, Allen, Bull. Amer. Mus. Nat. Hist., VII, p. 332.
Type Species. — Sciurus alfari, Allen.
Range. — Neotropical: Nicaragua, Costa Rica, Panama, Colombia, Ecuador,
Peru, and Rio Negro (Amazon).
Number of Forms. — About twenty-one.
Characters. — Skull strongly depressed posteriorly, and postorbital process
situated nearly or exactly over posterior zygomatic root, as in
Nannosciurus and Sciiirillus; but zvgomatic plate slanting gradually upwards as
in normal Squirrels, and orbit less circular than in these genera. Frontals very
broad, nasals short. Jugal broad. Upper incisors pro-odont, usually extending
beyond plane of tip of nasals. Palate normal. Bullae rather small. Cheekteeth
as in Sciurus, though the small outer (third) cusp of the upper molars is often
barely traceable. P. 3 present, and rather well developed, except in the type of
manarius, in which there seem to be no traces of this tooth.
Externally: size small; tail rather narrow or occasionally much narrowed,
rather shorter than head and body as a rule; digits as in normal Tree-squirrels.
This genus suggests the Sciurillus type of skull, but is in all respects a little
nearer to Sciurus in cranial characters. The lachrymal is usually situated rather
further back in relation to the toothrow than in members of the present section.
Forms seen : alfari, avunculus, boquetensis, brozvni, flaviventer, isthmius,
manarius, mimulus, napi, otinus, palmeri, rubrirostris, similis, simonsi.
The species were revised by Allen (Bull. Amer. Mus. Nat. Hist., XXXIII,
p. 145, 1914). All seem very closely related to each other, except perhaps
manarius, as noted above.
List of Named Forms
1. MICROSCIURUS ALF.^RI ALFARI, Allen
1895. Bull. Amer. Mus. Nat. Hist., VII, p. 333.
Volcan de Turrialba, near Jimenez, Costa Rica.
2. MICROSCILRU.S ALFARI VENUSTULUS, Goldman
1912. Smiths. Misc. Coll., LVI, 36, p. 4.
Gatun, Panama.
3. MICROSCIURUS ALFARI BROWM, Bangs
1902. BuU. Mus. Comp. Zool. Harvard Univ., XXXIX, 2, p. 24.
Bogava, Chiriqui, Panama.
330 MICROSCIURUS
4. MICROSCIURUS BOQUETKNSIS, Nelson
1903. Proc. Hiol. Soc. Washington, XVI, p. 121.
Boqucte, Chinqui, Panama.
5. MICROSCIURUS SIMILIS SIMILIS, Nelson
i8qg. Bull. Amer. Mus. Nat. Hist., XII, p. 78-
Cali. Cauca Valley, Colombia.
b. MICROSCIURUS SIMILIS FUSCULUS. Thomas
igio. Ann. Mag. Nat. Hist. 8, VI, p. 503.
Juntas, Rio San Juan, Choco District, Colombia.
7. MICROSCIURUS OTINUS, Thomas
1901. Ann. Mag. Nat. Hist. 7. VII, p. 193-
Medellin, Colombia.
8. MICROSCIURUS ISTHMIUS ISTHMIUS, Nelson
1899. Bull. Amer. Mus. Nat. Hist., XII, p. 77-
Truando River, Isthmus of Darien, Colombia.
Q. MICROSCIURUS ISTHMIUS VIV.\TUS. Goldman
1912. Smiths. Misc. Coll., LX, no. 2, p. 4.
Cana, Pirri Range, Eastern Panama.
10. MICROSCIURUS MIMULUS, Thomas
1S98. Ann. Mag. Nat. Hist. 7, II, p. 266.
Cachavi, Esmeraldes, Ecuador.
,1. MICROSCIURUS PALMKRI, Thomas
1909. .Ann. Mag. Nat. Hist. 8, IV, p. 234.
Sipi, Rio Sipi, Rio San Juan, Choco district, Colombia.
12.
MICROSCIURUS SIMONSI, Th
lomas
1900. Ann. Mag. Nat. Hist. 7, VI, p. 294.
Porvenir, near Zaparal, Bolivar Province, Ecuador.
13. MICROSCIURUS PERUANUS, Allen
1897. Bull. Amer. Mus. Nat. Hist., IX, p. 115.
Guayabamba, N.-W. Peru.
14. MICROSCIIRUS NAPI, Thomas
1900. Ann. Mag. Nat. Hist., 7, VI, p. 295- _, ^ , , , j
Rio Coca, Upper Rio Napo, Ecuador-Colombia boundary.
15. MICROSCIURUS RUBRIROSTRIS, Allen
1914. Bull. Amer. Mus. Nat. Hist., XXXIII, p. 163.
Chanchamayo, Central Peru.
,6. MICROSCIURUS FLORENCIAI-. Allen
ini4. Bull. Amer. Mus. Nat. Hist., XXXIII, p. 164.
Florencia, Caqueta District, S.-W. Colombia.
17. MICROSCIURUS AVUNCULUS, Thomas
1914. Ann. Mag. Nat. Hist. 8, XIII, p. 574-
Gualaquiza, Eastern Ecuador.
iS. MICROSCIURUS SF.PTENTRIONALIS, Anthony
1920. Journ. Mamm. Baltimore, i, p. 81.
Sabalos, on Rio San Juan, at junction of Rio Sabalos, Nicaragua.
SYNTHEOSCIURUS— SCIURUS 321
19. MICROSCIURUS SABANILLAE, Anthony
1922. Amer. Mus. Nov. 32, p. 2.
South Ecuador, Sabanilla, Prov. dc Loja; 5,700 ft.
20. MICROSCIURUS MANARIUS, Thomas
1920. Ann. Mag. Nat. Hist. 9, VI, p. 275.
Acajutuba, Rio Negro, Amazonas.
21. MICROSCIURUS FLAVIVENTER, Gray
1867. Ann. Mag. Nat. Hist. 3, XX, p. 432.
Brazil.
Genus 5. SYNTHEOSCIURUS, Bangs
1902. SYNTHEOSCIURUS, Bangs, Bull. Mus. Comp. Zool. Harvard Univ., XXXIX,
no. 2, p. 25.
Type Species. — Syntheosciurus brochiis, Bangs.
Range. — Panama.
Number of Forms. — One.
Characters. — Skull with no special peculiarities; rostrum rather long,
bullae relatively small, postorbital process moderate. Cheek-
teeth j", evidently like Sciunis (only two skulls with much worn teeth seen).
Upper incisors pro-odont, extending beyond plane of tip of nasals, and one-
grooved.
Fur very thick and soft; tail rather shorter than head and body; digits
normal arboreal type; ear strongly reduced.
Rem.arks. — This genus is retained by North American authors, including
Howell in his recent key to genera occurring north of South America. But the
sole character, pro-odont grooved incisors, appears to me to be questionable.
Elsewhere the incisors may be pro-odont, as in Callosciurus prevosti; and grooves
may appear from time to time, as in, for instance, Heliosciunis, in other genera.
The present species is a little known form. It is probably not more than sub-
generically separable from Sciurus.
Forms seen : brochiis.
List of N.\med Forms
I. SYNTHEOSCIURUS BROCHUS, Bangs
1902. Bull. Mus. Comp. Zool., Harvard Univ., XXXIX, no. 2, p. 25.
Boquete, Chiriqui, Panama. .Altitude, 7,000 ft.
Genus 6. SCIURUS, Linnaeus
1758. Sciurus, Linnaeus, Syst. Nat., loth Ed., vol. i, p. 63.
1899. Baiosciurus, Nelson, Proc. Acad. Sci. Washington, i, p. 31. Sciurus deppei,
Peters.
1 880. EcHiNOSciURUs, Trouessart, le Naturaliste, 2, p. 292. Sciurus hypopyrrhus, Wagner.
1880. Neosciurus, Trouessart, le Naturaliste, 2, p. 292. Sciurus carolinensis, Gmelin.
Valid as a subgenus: see Howell, 1938, N.A. Fauna, 56, p. i.
1899. Hesperosciurus, Nelson, Proc. Acad. Sci. Washington, i, p. 27. Sciurus grisetis,
Ord. Valid as a subgenus: see Howell, 1938.
21 — Living I<o<lents — 1
XXXIV,
p
212.
Sciuriis
XXXIV
p-
213-
Sdurus
XXXIV
p-
280.
Sciunts
XXXIV,
p-
265.
Sciunis
XXXIV,
p-
267.
Scittrus
XXXIV.
p-
199.
Sciiirus
322 SCIURUS
SciURrs ^continued)
i8qg. Otosciurus, Nelson, Proc. .Acad. .Sci. Washington, 1. p. 28. Sciunis aberti,
Woodhouse. Valid as a subgenus: see Howell, 1938.
igog. Texes, Thomas, .\nn. Mag. Xat. Hist. 8, HI, p. 468, footnote. Scinrus persicus,
Erxleben (= anomahis. Guldenstaedt). Valid as a subgenus.
1935. Oreosciurls, Ognev, Abstr. Works. Zool. Inst. Moscow, 2, p. 50. Sciunis
anomalus, Guldenstaedt.
1821. Gi.'ERLiNGLrETL's, Gray, London Med. Repos., XV, p. 304. Sciurus guerlinguetus.
Gray {= Sciurus aesiuans, Linnaeus). Valid as a subgenus.
1823. \L\CROXUS, F. Cuvier, Dents des Manim., p. ibi. Le Guerlinguet (Sciurus
oestuans, Linnaeus).
18S0. Par^cu'RVS, Trouessart, le Naturaliste, II, p. 292. Sciurus niger, Linnaeus.
Valid as a subgenus: see Howell, 1938.
1899. .■Vr.^eosciurus, Nelson, Proc. Washington .-Xcad. Sci., I, p. 29. Sciurus oculatus,
Peters.
1915. Mesosciurus, Allen, Bull. .Amer. Mus. Nat. Hist.,
aestuans hoffmani, Peters.
1915. HiSTRiosciURus, Allen, Bull. .Amer. Mus. Nat. Hist.,
gerrardi. Gray.
1915. SiMOSciURUS, .Allen, Bull. Amer. Mus. Nat. Hist.,
stramineus, Eydoux & Souleyet.
1915. Hadrosciurus, Allen, Bull. Amer. Mus. Nat. Hist.,
flammifer, Thomas. Valid as a subgenus.
1915. Uroscrtjus, Allen, Bull. .Amer. Mus. Nat. Hist.,
tricolor, Poeppig.
1915. Leptosciurus, .Allen, Bull. Amer. Mus. Nat. Hist., XXXIV,
pucherani, Fitzinger.
1914. NoTosciURUS, Allen, Bull. Amer. Mus. Nat. Hist., XXXIII, p. 585. Notosciurus
rhoadsi, .Allen. Valid as a subgenus.
The arrangement of these names subgenerically is as follows :
Subgenus i. Sciurus, Linnaeus (restricted by Howell, 1938, to Palae-
arctic vulgaris group).
Subgenus 2. Tenes, Thomas.
Synonym: Oreosciurus, Ognev.
Subgenus 3. Neosciurus, Trouessart.
Synonym: Echinosciunis, Trouessart ) (see Howell,
Baiusciuriis, Nelson ) 1938)-
Subgenus 4. Hesperosciurus, Nelson.
Subgenus 5. Otosciurus, Nelson.
Subgenus 6. P.\rasciurus, Trouessart.
Synonym: Araeosciurus, Nelson.
Subgenus 7. Guerlinguetus, Gray.
Synonym: Alacroxus, Cuvier.
Mesosciurus, Allen.
llislriosciurus, Allen.
Simosciurus, Allen.
Leptosciurus, Allen.
Subgenus 8. Notosciurus, Allen.
Subgenus 9. Hadrosciurus, Allen.
Synonym: Urosciurus, Allen.
Type Species. — Sciurus vulgaris, Linnaeus.
SCIURUS 323
Range. — Europe, from Ireland eastwards and from North Scandinavia to
the Mediterranean; the greater part of European Russia; south to
Crimea; Caucasus, Asia Minor, parts of Syria and Persia; Siberia, east to
Anadyr region, and south to Altai, Transbaikalia, Amur, Ussuri; North Mon-
golia; Manchuria, Chihli, Korea, Saghalien, Japan. South-eastern Canada
(Ontario); U.S.A. : Oregon, Minnesota, California, Arizona, Colorado, New
Mexico, Te.xas, Louisiana, Florida, eastern States generally; Mexico south
through Central America to Colombia, Ecuador, Peru, Bolivia, Jujuy (North
Argentina), Brazil generally, the (iuianas, Venezuela, Trinidad, Tobago.
Number of Forms. — .\bout a hundred and ninety.
Characters. — The skull, typically, is with moderately developed post-
orbital process; the braincase is strongly depressed post-
eriorly; there are no prominent ridges present for muscle attachment; except
in the South American subgenus Hadrusciurus, the parietal ridges very rarelv
show signs of joining together. The frontals tend to be slightly depressed.
The palate is broad, rather square posteriorly, and terminating just behind
the toothrows. The incisive foramina are small, far in front of toothrows.
The bullae are usually relatively large. The zygomatic plate slants gradually
upwards, its ridge moderately or weakly developed; the zygomatic plate not
tending to become narrowed in the upper border, the front face usually
appearing broad and flat. The infraorbital foramen always forms a canal ; the
masseter knob is weak. The length of the rostrum is in the genus variable, but
never tends to become extreme. Mandible with no special peculiarities.
In the upper cheekteeth there are on the main teeth four transverse ridges,
the second and third of which are higher than the first and fourth which respec-
tively form the anterior and posterior margins of the tooth. The two centre
ridges are nearly parallel and extend across to the inner side of the tooth, which
is rounded and formed as usual in the family by one large inner cusp. The
outer sides of the second and third ridges are formed bv well-marked cusps,
which are usually originally fairly high; between these is placed a small outer
cusp, which may become very low but is usuallv traceable. M.3 has only one
prominent transverse ridge, the second; the third ridge of the other teeth is
obsolete in this tooth, which has a corresponding wide depression behind the
second ridge. P. 3 present or absent; when present, vestigial as a general
rule.
In the lower teeth, the premolar is usually noticeably smaller than the other
teeth, and M.3 is not much elongated as it often is, for instance, in Callosciurus;
each tooth has a more or less well-marked cusp at each corner, and with the
exception of certain forms of the subgenus Hadrosciuriis, the main central de-
pression is always well marked. The anterointernal cusp is the highest, the
posterointernal the lowest. Between these two, and between the two outer main
cusps, is placed a small extra cusp each side.
Externally, size moderately large or moderatelv small; the tail alwavs thickly
bushy, very rarely showing signs of becoming narrow; it may be longer than the
head and body, as in some forms of stramineus, or subequal in length to this
324 SCIURUS
measurement, or considerably shorter, as in itnomalus. In the type species, the
sole of the hindfoot is densely hairy in winter. Tiic digits are arranged as
characteristic of all arboreal Squirrels; D.4 the longest digit in the hindfoot,
D.3 and D.2 successively each a little shorter; D.5 well developed, nearly as long
as D.z; hallux moderate. In the forefoot, D.4 is longer than D.3 (slightly);
the two other main digits are subequal, and a little shorter than the central two
digits. The ear is usually prominent; in z-ii/qaris and aberti, conspicuous ear-
tufts are present, the development of which in vulgaris varies seasonally.
Subgenus SciURUS (Palaearctic range of the genus except the Caucasus,
Syria, Persia, and Asia Minor); this has once again been restricted to the
Palaearctic, by Howell, 1938, though formerly Miller, 1923, regarded
the majority ot the North American species as belonging to it. Plantar
pads 4; ear tufted; sole densely haired in winter. Cheekteeth j.
Mammae 8 (Miller). Includes vulgaris, with numerous races, and the
Japanese Us, which in the absence of knowledge of detail characters such
as the baculum appears to me to be doubtfully specifically distinct from
vulgaris.
Subgenus Tenes (Caucasus, Syria, Persia, Asia Minor); aiioiiialus and races
(synonvm " Oreosciurus," Ognev). Plantar pads 6; mammae 10; baculum
said to differ from vulgaris (Ognev); ears not tufted; cheekteeth normally
:[. (Fur less thickened than is usual in vulgaris.) Tail rather short,
averaging about 70 per cent of head and body length in a comparatively
small series of specimens examined.
Subgenus Neosciurus (Eastern U.S.A., Eastern Canada, Mexico, Honduras,
Guatemala, Nicaragua, Costa Rica). (Synonym " Echiuosciurus," "Baio-
scimus ").
"The skull of S. carolineiisis (type of subgenus Neosciurus) does not
differ widely in general shape from that of S. vulgaris, but is relatively
longer, with braincase shallower and more elongated, . . . and rostrum
longer and relatively narrower; postorbital process shorter and stouter. . .
"In 5. deppei (type of Baiusciurus, Nelson), P. 4 averages slightly
more quadrate than the same tooth in Neosciurus, but examination of a
large series of deppei and carolinensis shows that the character is too slight
and inconstant to serve as a basis for suhgeneric distinction.
" S. a. Iivpopyrrlnis and the large group of Mexican forms associated
with it by Nelson in the subgenus Ecliiiiosciurus differ in general from
carolinensis in having a shorter and relatively broader rostrum, and a more
or less prominent depression in the frontals; these differences, however,
are considered too slight to warrant the recognition of the group"
(Howell, 1938).
The baculum of carolinensis is said to be essentially as in vulgaris;
according to Howell, this character is in deppei and adolphei dorsalis
(one of the aureogaster group ( = " Ecliinosciurus")) essentially as in
carolinensis. There are 6 mammae in deppei, which species appears to
have a rather lower coronoid process than is usual, though this character
SCIURUS 325
may be present also in some of the South American forms. Cheekteeth,
in this subgenus, J.
Subgenus Hespkrosciurus (CaUfurnia, Oregon; "Western Gray Squirrels").
"The skull of S. griseiu resembles that of carolinensis very closely except
that it is larger . . . the jugal relatively lighter (shallower). Baculum
widely different from Neosciurus, resembling more closely that of S.
aberti" (Howell, 1938). P. 3 in our small series relatively well developed.
Subgenus Otosciurus (Arizona, Colorado, New Mexico, and Northern
Mexico; aberli group). "In general shape the skull of S. aberti closely
resembles that of S. vulgaris. P. 3 relatively larger and more strongly
developed, the crown being subject to wear with the rest of the molars.
. . . Compared with Neosciurus, this subgenus differs in having braincase
and interorbital region relatively broader, postorbital breadth less than
interorbital breadth, postorbital process larger and longer, postorbital
notch deeper, P. 3 more strongly developed" (Howell, 1938). The
baculum differs widely from Neosciurus, and is nearer griseus. The
baculum of each of the four subgenera occurring in the United States is
figured by Howell, 1938, N.A. Fauna, no. 56, p. 35.
Subgenus Parasciurus. (Synonym " Araeosciuriis") (Eastern United
States including Florida; Texas, Arizona, Mexico) {iiiger group).
"Compared with Neosciurus, the skull of very similar shape, . . . the
frontals slightly elevated on posterior half; a distinct interorbital notch;
the baculum closely similar to that of 5. carolinensis. Cheekteeth
reduced to :!. The parietal ridges are in the few skulls examined
tending to be prominent, and probably would join, though actually no
skull has been seen with this feature. Relatively large Squirrels.
Subgenus Guerlinguetus. (Synonym " Mesosciurus," " Histriosciurus,"
" Sifnosciurus," " Leptosciurus.") "Skull differs from Parasciurus in
having a shorter rostrum, more swollen braincase, and position of notch
in maxillary plate of zygoma . . . there is one upper premolar, which is
subcircular or quadrate in shape, differing thus from Parasciurus in
which this tooth is subtriangular" (Howell). This group ranges from
Nicaragua southwards over most of South America, south to Jujuy.
In 1915 Allen divided the South American Squirrels into many
"genera." These names were based simply on mammary formula
(6 or 8), the bodily size, the relative length of the rostrum (shown
subsequently in some cases to be incorrect), the geographical distribution.
He keyed these genera as follows :
"Mammae 6.
Premolars ,'
Size small, total length 320-380. Tail shorter than
head and body.
Soles naked, plantar pads normal. Leptosciurus
Soles heavily furred nearlv the whole length;
plantar pads all near base of toes. Notosciurus
326 SCIURUS
Size medium, total length 375-450. Tail subequal
to or shorter than head and body. Mesosciuriis
Mammae 8.
Premolars -] ; tail as long as or longer than head and body.
Size small, tail narrow. Gucrliiiguetus
Size large, total length 41)0-580, tail broad and bushy.
Skull broad and heavy, rostrum short. Hadrusciurus
Skull long and narrow, rostrum slender. Urosciurus
Size large, tail very long and narrow, rostrum very
broad and short. Simosciurus"
It will be seen that " Mesosciui iis" differs from Guerlinguetus in
having mammae 6 instead of 8, and " Simosciiinis" dilTers in being larger.
The majority of these names, based on characters such as these, have
naturallv been currently disregarded. Hadrosciurus, Urosciurus and
Simosciurus have been shown by Lonnberg to be quite indistinguishable
from each other on the cranial characters proposed by Allen.
Thomas stated that Urosciurus was not distinguishable from Hadro-
sciurus. Those forms occurring in South America have not been revised
like the forms occurring north of Panama (Howell, Miller, etc.). I think
there is no doubt that, pending a full revision of all the forms all will
have to stand as synonyms of the oldest name Guerlinguetus except
perhaps Notosciurus (not seen), and certainly Hadrosciurus, which as I shall
endeavour to show is so distinct dentally that it might almost form a genus.
In addition therefore to the acsUians group (tvpical Guerlinguetus),
the subgenus will include the liofjinani group, " Mesosciurus," regarded
as a synonym by both Miller and Howell; the st rami neus group, "Simo-
sciurus," large attractive long-tailed Squirrels from Peru and Ecuador
with, however, absolutely no distinctive features cranially and dentally;
and the pucherani group, " Lepiosciurus" ; this subgenus was described
as " Skidl similar in general form and proportions to iS'. aestuans; differing
from Guerlinguetus in the structure of the upper molars, the outer border
of the crowns having only two prominent cusps instead of three, the
intervening cusplets usually prominent in Guerlinguetus and most other
American Tree-squirrels are practically obsolete or entirely absent." But I
have seen some specimens belonging to this group with this character not
clearly developed; and I have seen specimens of aestuans which appear to
metobe indistinguishable (romLeptosciurus; Ithink it is highlyimprobable
that this is a constant feature dividing the two groups into subgenera.
Subgenus Notosciurus (rhoadsi, Ecuador. Not seen).
"Size small, tail of medium length — naked portion of plantar surface
of hindfeet restricted to distal half, the rest heavily furred; the posterior
pad large, nearly square, occupying the whole breath of the sole, close
to the toe pads ' ' (Allen). This is evidently based on one young individual,
the milk premolar being retained, according to Allen, in the one skull,
so that it is not known whether the permanent dentition would include
SCIURUS 327
P. 3. Originally proposed as a genus; but hairiness of sole is certainly
not a valid character; compare winter specimens of Sciurus vulgaris.
Subgenus Hadrosciurus. (Synonym; " Urosciurus.") (Venezuela, Peru,
Brazil, Colombia, Ecuador, Bolivia).
This is a very distinct group of Squirrels, which may ultimately have
to be regarded as a genus, which seem to be paralleling to a certain
degree such forms as Epixerus and Rheithrosciurus, in that the incisors
are becoming thickened, and the toothrows rather strongly reduced.
The muzzle is pointed, the palate relatively narrow. In the upper teeth,
the cusps are evidently low, at all times, but the normal cusps and ridges
are traceable. In the lower teeth, a rather different appearance from
normal Sciurus is frequently present. There is often a well-developed
transverse ridge extending across the middle of the tooth and connecting
the small intermediate cusps; this ridge may isolate as an island. The
small anterior transverse ridge in the lower teeth regarded by Hollister
as characteristic of the African Heliosciunis is in this subgenus usually
well developed. It would be interesting to examine a large series of
lower molars of this group, and ascertain how far this general pattern is
constant, or if it is present in some races, and not in others, as the material
examined appears to vary somewhat in development. The central
depression in those forms with the pattern fully developed becomes
reduced, which as far as I have seen does not happen elsewhere in the
genus. A short sagittal crest is often present.
Some figures have been obtained in order to see if the group would
stand as a genus on the reduction of the toothrow, but the results are
disappointing as some of the other groups overlap. The percentages of
toothrow compared with total length of skull are given below, throughout
the genus, in a small number of skulls measured.
PERCENTAGE OF TOOTHROW
.\G.\INST TOTAL LENGTH OF SKVLL
A
Subgenus Hadrosciurus
Guerlinguetus aestuam group.
" Leptosciurus" puclierani group.
Guerlinguetus hoffmam group.
Guerlinguetus gerrardi and allies.
Guerlinguetus stramineus group (" Simosciurus").
Subgenus Parasciurus.
Sciurus anomalus.
Sciurus carolinensis.
Sciurus vulgaris.
Sciurus deppei.
Sciurus aberti.
Sciurus aureogaster group.
Sciurus grisetts.
The species referred to Hadrosciurus are relatively large forms.
"ERAGE
EXTREMES
15-3
i4-2-i6-i
15-8
I4-9-I7-8
i6-i
i4-6-i6-7
i6-8
15-17-2
i6-8
i6-4-i7-2
17-6
17-2-18
177
I7-I-I8-6
17-8
I7-1-I8-6
17-8
17-5-18-7
i8-i
17-9-18-4
i8-3
17-3-18-9
i8-4
18-19-1
i8-6
17-4-20-6
i8-7
18-3-19-6
328
SCIURUS
The species of Sciiirus occurring in Western Europe are revised by Miller,
Cat. Mamm. Western Europe, 191 2, p. 897.
Species occurring in Mexico and Central America by Nelson, 1899, Proc.
Washington Acad. Sci., I, pp. 15-106 (under a number of subgcneric names).
Species occurring in South America by Allen, 191s, Bull. Amer. Mus. Nat.
Hist., XXXIV, pp. 147-288 (under a number of generic names).
The North American subgenera are, as indicated above, revised by Howell,
1938, N.A. Fauna, no. 56, which together with a revision of Nearctic Citellus
Fig. 94. SciuRus vulgaris vulgaris, Linnaeus.
B.M. No. 97.4.1 I.I, cJ; -■: li.
clears up much misunderstanding as regards the status of genera and subgenera
of Squirrels occurring in North America, and has made my task very much more
simple, as the North American collection of Seiuridae at the British Museum
is not representative.
Forms seen : aberti, aestuans, alleni, alphonsei, anomaliis, annalium, anthonyi,
argenleiis, argeiitiiiiis, aureogaster, holiviensis, boiidae, boothiae, " bniiuieoniger"
caroUnensis, cast us, cluipmani, cliiiiquensis, "ciiicri'iis," cocalis, cocos, colliaei,
Cdieae, croaticus, '' cusciinis" deppei, doisalis, diirangi, flaminifer, "fraseii,"
Jiiiiiientor, fiilminalus, fuscoater, fiiscoiiigrictiiis, fuscorubeiis, goldmani, griseoflavus,
giiscogena, gn'sciis, guayanus, hyporrlwdus, igiiiventiis, infiiscntiis, ingrami,
irroratiis, itallcus, jura/is, "k/agcsi," laiigsdorffi, kiicotis, /eticoiinis, /i/aetis, lis,
iiiacconiie/li, inaiitchiin'ciis, martcnsi, viedeUinensIs, inelaiua, iiie/aiiotis, iiien'deiisis,
" millcri" ( = leouis), luidvniciisis. navniitensis, niboiixi, nelsoni, nemoraUs, nesaeiis,
niger, " nigrescens," nigiipes, niiiiuiiitlus, vcidntns, " oc/iiesceiis," orientis, pallescens,
SCIURUS
329
paraensis, pucherani, pyrrhinus, pyrrlionotus, quelchi, "quercinus" {=:hernandezi),
quiiidianuSy rigidtis, rufiienter, nipestiis, russtis, segurae, sinaloensis, steinbachi,
stramijieus, taedifer, tepariiis, tobagensis, tricolor, "variabilis" (=gerrardi),
variegatoides, varius, "versicolor" {=inconstans), vulgaris, yiicatanemis.
Fig. 95. SciURUS \t.'lgaris vulgaris, Linnaeus.
B.M. No. 97.4.11.1, S; X ij.
Fig. q6. Sciurus vulgaris vulgaris, Linnaeus.
Cheekteeth; X S-
List of N.\med Forms
"Natios," "morphs" and other sub-sub-species of Russian authors are not
accepted.
330 SCIURUS
Subgenus Schiriis, Linnaeus
1. SCIURUS VULGARIS VULGARIS, L.nnaeus
1758. Syst. Nat., Ed. 10, I, p. 63.
Upsala. Sweden.
S>'nonym: europaeus. Gray, 1843, List, Mamm., p. 139.
riifiis, Kerr, 1792, Anim. Kingd., p. 255.
albonotatiis, BillberK, Synopsis Faunae, Scandinaviae, p. 2
Southern Sweden. 1827.
albus, Billber^, same reference. Skane, Sweden.
niger, Billberg, same reference. Skane, Sweden.
typlnis, Barrett-Hamilton, Proc. Zool. .Soc. London, p. 6
1899.
2. SCIURUS VULGARIS VARIUS, Brisson
1762. Regn. Anim, p. 106.
Northern Europe.
SynonjTii; cinereijs, Fischer, 1829, .Synops. Mamm., p. 353.
3. SCIURUS VULGARIS LEUCOURUS, Kerr
1792. Anim. Kingd., p. 256.
England.
4. SCIURUS VULG.^RIS RUSSUS, Miller
1907. .Ann. Mag. Nat. Hist. 7, XX, p. 427.
Dinan, France.
5. SCIURUS VULGARIS FUSCOATER, Altum
1876. Forstzoologie, 2nd ed., I, p. 75.
Harz Mountains, Germany.
Synonym: nigresceiis, Altum, same reference.
briintiea, Altum, same reference.
graeca, Altum, same reference.
gothardi. Fatio, Arch. Sci. Phys. Nat. Geneve, 4th ser., xix,
p. 512, 1 905 . South slope of Mt. St. Gothard, Switzer-
land.
rtililans. Miller, Ann. Mag. Nat. Hist., 7, XX, p. 426,
1907. Rudolstadt, Thiiringen, Germany.
6. SCIURUS VULGARIS ITALICUS, Bonaparte
183S. Iconog. Faun. Ital., i, fasc. 23.
Italy.
Synonym: meridionalis, Lucifero, Re\ ista Ital. Sci. Nat. Siena, XXVII,
p. 45, 1907. Sila, Calabria, Italy.
7. SCIURUS VULGARIS LILAEUS, Miller
1907. Ann. Mag. Nat. Hist. 7, XX, p. 429.
Agoriani, Greece. (North side of Lyakura (Parnassus) mountams.)
S. SCIURUS VULGARIS ALPINUS, Desmarest
1822. Mamm., II, p. 543.
Pyrenees.
9. SCIURUS VULGARIS NUMANTIUS, Miller
1907. .Ann. Mag. Nat. Hist. 7, XX, p. 428.
Pinares de Quintanar de la Sierra, Burgos, Spain.
10. SCIURUS VULGARIS INFUSCATUS, Cabrera
1905. Bol. Real. Soc. Espafi. Hist. Nat. Madrid, V, p. 227.
Las Navas, Avila, Spain.
SCIURUS 331
11. SCIURUS VULGARIS SEGURAE, Miller
1909. Ann. Mag. Nat. Hist. 8, III, p. 418.
Molinicos, Sierra de Segura, Albacete, Spain.
12. SCIURUS VULGARIS B.\ETICUS, Cabrera
1905. Bol. Real. Soc. Espaii. Hist. Nat. Madrid, V, p. 228.
Alanis, Seville, Spain.
13. SCIURUS VULGARIS SILANUS, Hecht
1931. Zeitschr. filr Saugetierk. Berlin, 6, p. 238.
Silania, Italy.
14. SCIURUS \TJLGARIS AMELIAE, Cabrera
1924. Bol. Real Soc. Espaii. Hist. Nat. Madrid, XXIV, p. 420.
Greece, Kontinoplo, Mt. Olympus.
15. SCIURUS VULG.\RIS CROATICUS, Wettstein
1927. Anz. Akad. Wien, I, p. i.
Croatia, Apatisanska Duliba Forest, south-east of Krasno.
16. SCIURUS VULGARIS BALCANICUS, Heinrich
1936. Bull. Inst. R. Hist. Nat. Sophia, g, p. 41.
WalderamUnterlauf der Kamtschija, Ostauslaufer des Balkan, Bulgaria.
17. SCIURUS VULGARIS ISTRANDJAE, Heinrich
1936. Bull. Inst. R. Hist. Nat. Sophia, 9, p. 41.
Dorf. Karamlek, Strandjabalkan, Bulgaria.
18. SCIURUS VULGARIS RHODOPENSIS, Heinrich
1936. Bull. Inst. R. Hist. Nat. Sophia, 9, p. 41.
Dorf Tschepelare, Central Rhodopen, Bulgaria.
19. SCIURUS VULGARIS N.ADYMENSIS, Serebrennikov
1928. C.R. Acad. Sci. U.S.S.R., p. 422.
Nadym River, West Siberia.
20. SCIURUS VTJLGARIS MARTENSI, Matschie
1901. Archiv. fiir. Naturgesch. Berlin, p. 313.
Lower Yenesei River, Siberia.
21. SCIURUS VULGARIS ALT.AICUS, Serebrennikov
1928. C.R. Acad. Sci. U.S.S.R., p. 422.
Kok-Su River, mouth of Yamanush River, Altai Mountains. (Listed
as a valid race by Vinogradov, 1933.)
22. SCIURUS VULGARIS FUSCONIGRICANS, Dwigubski
1804. Prodromus Faunae Rossicae, p. 85.
Bargusin, Transbaikalia.
23. SCIURUS VULGARIS FUSCORUBENS, Dwigubski
1804. Prodromus Faunae Rossicae, p. 85.
East Siberia.
24. SCIURUS VXFLGARIS DULKEITI, Ognev
1929. Zool. Anz., 83, p. 76.
.\muka River, Great Shantar Islands, east coast of Siberia.
25. SCIURUS \XLG.\RIS MANTCHURICUS, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. 501.
Khingan, Manchuria.
332 SCIURUS
26. SCIURUS VULGARIS RUPESTRIS, Thomas
i(jo7. Proc. Zool. Soc. London, p. 410.
Darine, 25 miles N.W. of Korsakoff, Satjhalien.
27. SCRTRUS VULGARIS EXALBIDUS, Pallas
177S. Nov. Sp. Quaclr. Glir. Ord., p. 374.
Locality not known. (Distribution: Western Siberia.)
28. SCIURUS VLILGARIS JACUTENSIS, Ognev
1930. Bull. Pacif. Sta. Vladivostock, 2, no. 5, pp. 18, 41.
Yakutsk, Siberia.
2q. SCIURUS VULGARIS FEDJUSHINI, Ocnev
1935. Abstr. Works. Zool. Inst. Moscow, 2, p. 43.
Minsk, West Russia.
30. SCIURLIS VULGARIS FORMOSOVl. Ognev
1935. Abstr. Works. Zool. Inst. Moscow, 2, p. 44.
Nijni-Novgorod, Russia.
31. SCIURUS VULGARIS BASHKIRICUS, Ognev
1935. Abstr. Works. Zool. Inst. Moscow, 2, p. 45.
Samara, Russia.
Synonym: vii/goris Imskiriciis natio iiialcnsis, Ogne\', same reference,
p. 46. Ural.
32. SCIURUS VULGARIS JENISSEJENSIS, Ognev
1935. Abstr. Works. Zool. Inst. Moscow, 2, p. 47.
Lower Tunguska, Turuchansk, East Siberia.
33. SCIURUS VULGARIS COREAE, Sowerby
1921. Ann. Mag. Nat. Hist. 9, VII, p. 252.
Kaldguai, 55 miles N.E. of Seoul, Korea.
34. SCIURUS VULGARIS CHILIENSIS, Sowerby
1921. .-Xnn. Mag. Nat. Hist. 9, VII, p. 253.
Tung-ling, 75 miles north-east of Peking, China.
35. SCIURUS VULGARIS KESSLERI, Migulin
192S. Prot. Plant Ukraine, no. 3-4, p. 83.
Zhitomir and Shepetovka, West Ukraine, European Russia.
36. SCIURUS VULGARIS UKRAINICUS, Migulin
192S. Prot. Plant Ukraine, no. 3-4, p. 82.
Sumsk district, Kharkov, Russia.
37. SCIURUS VULGARIS OGNEVI, Migulm
1928. Prot. Plant Ukraine, no. 3-4, p. 84.
Rynski ^'illage, Bororski district, Kharkov govt., Russia.
38. SCIURUS VULGARIS ARCTICUS, Trouessart
1906. Bull. Mus. Hist. Nat. 6, p. 365.
Lena River, North Siberia.
39. SCIURUS VULGARIS ORIENTIS, Thomas
1906. Proc. Zool. Soc. London, 1905, ii, p. 345.
Aoyama, Hokkaido, Japan.
40. SCIURUS VULGARIS K.ALBINENSIS, Selewin
1935. Bull. Univ. Tachkent, 19, pp. 75-77-
Altai, west of Irtish.
SCIURUS 333
41. SCIURUS VULGARIS ARGENTEUS, Kerr
1792. Anim. KinKii., p. 256.
Altai.
iSciurus vulgaris calotus, Hodgson, 1842, Calcutta Journ. Nat. Hist., ii,
p. 221, "hiKh regions of Central Asia," is currently regarded as
unidentifiable.
42. SCIURUS LIS, Temminck
1842. Fauna Japonica, p. 45, pi. xii, figs. 1-4.
Central Japan.
Subgenus Tenes, Thomas
(Synonym: Oreosciurus, Ognev)
43. SCIURUS ANOMALUS ANOIVL'^LUS, Guldenstaedt
1785. Schreber Saugth., IV, p. 78 1.
Sabeka, 25 kms. south-west of Kutais, Georgia, Caucasus.
Synonym: caucasicus, Pallas, 181 1, Zoogeogr., I, p. 186.
russatiis,Wagner, iS42,Schreber's Saugth. Suppl.,III,p. 155.
historicus, Gray, 1867, Ann. Mag. Nat. Hist., 3, XX, p. 273.
44. SCIURUS ANOMALUS PALLESCENS, Gray
1867. Ann. Mag. Nat. Hist. 3, XX, p. 285.
Asia Minor.
45. SCIURUS ANOMALUS FULVUS, Blanford
1875. Ann. Mag. Nat. Hist. 4, XVI, p. 311.
Shiraz, Persia.
46. SCIURUS ANOMALUS SYRIACUS, Ehrenberg
1829. Symp. Phys., I, pi. 8.
Lebanon.
Ognev has also listed persicus (Erxleben, Syst. Regn. Anim. i, p. 417,
1777) as a valid form, but there is reason to belie\e that this name
was based on a Dormouse, Glis glis.
Subgenus Neosciurtis, Trouessart
(Synonym: Baiosciuriis, Nelson.
Echinosciurus, Trouessart.)
caioliiiensis Group
(Revised by Bangs, Proc. Biol. Soc. Washington, X, pp. 153-159, 1896.)
47- SCIURUS CAROLINENSIS CAROLINENSIS, Gmelin
1788. Syst. Nat., i, p. 148.
"Carolina."
Synonym: migraloriiis, Audubon & Bachman, 1854, Quad. I, p. 265.
cinerciis, Schreber, Saugth., IV, p. 766, 1792.
hiemalis, Ord, 1815, Guth. Geog., II, p. 292.
48. SCIURUS CAROLINENSIS EXTIMUS, Bangs
1896. Proc. Biol. Soc. Washington, X, p. 15S.
Miami, Florida. (Dade County.)
49- SCIURUS CAROLINENSIS FULIGINOSUS, Bachman
1838. Proc. Zool. Soc. London, p. 97.
Near New Orleans, Louisiana.
334 SCIURUS
50. SCIURUS CAROi.INENSIS H^'POPHAEUS, Merriam
18S6. Science, VII. p. 351.
Elk River, Sherburne County, Minnesota.
51. SCIURUS CAROLINENSIS LEUCOTIS, Capper
1830. Zool. Journ., V, p. 206.
Between York and Lake Simcoe, Ontario, Canada.
atireogaster Group
52. SCIURUS AUREOGASTER AUREOGASTER, F. Cuvier
1829. Hist. Nat. Mamm. 6, livr. 50, pi. with text (Binomial published at end of work
only, vol. 7, tabl. gen. et. meth., p. 4, 1842).
"California," really Eastern Mexico.
Synonym: rafiventcr, Lichtenstein, 1830, (1827) Ab. K. .Akad. Wiss.
Berlin, 1 16.
lencogaster, Cuvier, Suppl. H.N. Buffon, i, p. 300, 1S31.
iiiusteliiuis, Audubon & Bachman, 1841, Proc. Acad. Nat. Sci.
Philadelphia, p. 100.
ferrugineiveniris, Audubon &: Bachman, same reference, p. loi.
chrysogaster, Giebel, Saugeth., p. 650, 1855.
hypoxaifthiis, GeofFroy, Voy. de la Venus, Zool., p. 158, 1855.
53. SCIURUS AUREOGASTER HYPOPYRRHUS, Wagler
1831. Isis, p. 510.
Mexico, probably in Vera Cruz.
Synonym: morio, Gray, Ann. Mag. Nat. Hist. 3, XX, p. 424, 1867.
niaunis, Gray, same reference, p. 425.
nigresceiis, Bennett, Proc. Zool. Soc, London, p. 41, 1S33.
54. SCIURUS AUREOGASTER FRUMENTOR, Nelson
1898. Proc. Biol. Soc. Washington, XII, p. 154.
Las Vegas, Vera Cruz, Mexico.
55. SCIURUS POLIOPUS POLIOPUS, Fitzinccr
1867. Sitz-Bcr. .^ad. Wiss. Wien. Math. Nat. CI., LV, Abth. i, p. 478.
Cerro San Felipe, Oaxaca, Mexico.
Synonym; iiagiieri. Alien, 1S9S, Bull. Anier. Mus. Nat. Hist., X, p. 453.
albipes, Wagner, 1837, Abh. math.-phys. CI. k. baytr. .\kad.
Wiss. Miinchen, II, p. 501, not of Kerr.
leucops, Gray, Ann. Mag. Nat. Hist., 3, XX, p. 427, 1867.
56. SCIURUS POLIOPUS HERNANDEZI, Nelson
1898. Science, N.S., VIII, p. 783.
Mountains 15 miles west of Oaxaca, Mexico.
Synonym: querfimis, Nelson, 1898, Proc. Biol. Soc. Washington, XII,
p. 150, not of Erxlebcn.
57. SCIURUS POLIOPUS PEREGRINATOR, Nelson
1904. Proc. Biol. Soc. Washington, XVII, p. 149.
Piaxtla, Puebla, Mexico.
58. SCR'RUS POLIOPUS NEMORALIS, Nelson
1898. Proc. Biol. Soc. Washington, XII, p. 151.
Patzcuaro, Michoacan, Mexico.
59. SCIURUS POLIOPUS SENEX, Nelson
1904. Proc. Biol. Soc. Washington, XVII, p. 148.
La .Salada. S.-E. Michoacan, 40 miles south of LIruapan, Mexico.
SCIURUS 335
60. SCIURUS POLIOPUS CERVICALIS, Allen
1890. Bull. Amer. Mus. Nat. Hist., Ill, p. 183.
Hacienda San Marcos, Jalisco, Mexico.
61. SCIURUS POLIOPUS COLIMF.NSIS, Nelson
1898. Proc. Biol. Soc. Washington, XII, p. 152.
Hacienda Magdalena, Colima, .Mexico.
62. SCIURUS POLIOPUS EFFUGIUS, Nelson
1898. Proc. Biol. Soc. Washington, XII, p. 152.
Mountains west of Chilpancingo, Guerrero, Mexico.
63. SCIURUS POLIOPUS TEPICANUS, Allen
1906. Bull. Amer. Mus. Nat. Hist. XXII, p. 243.
Rancho Palo Amarillo, Nayarit, Mexico.
64. SCIURUS NELSONI NELSONI, Merriam
1893. Proc. Biol. Soc. Washington, VIII, p. 144.
Huitzilac, Morelos, Mexico.
65. SCIURUS NELSONI HIRTUS, Nelson
1898. Proc. Biol. Soc. Washington, XII, p. 153.
Tochimilco, Puebla, Mexico.
66. SCIURUS COLLIAEI COLLIAEI, Richardson
1839. Voyage H.M.S. Blossom, Zool., p. 8.
San Bias, Nayarit, Mexico.
Synonym: griseocaudatus. Gray, 1844, Zool. Sulphur, i, p. 34.
67. SCIURUS COLLIAEI NUCHALIS, Nelson
1899. Proc. Washington Acad. Sci. I, p. 59.
Manzanillo, Colima, Mexico.
68. SCIURUS SIN.-VLOENSIS, Nelson
1899. Proc. Washington Acad. Sci. I, p. 60.
Mazatlan, .Sinaloa, Mexico.
69. SCIURUS TRUEI, Nelson
1899. Proc. Washington Acad. Sci. I, p. 61.
Camoa, Rio Mayo, Sonora, Mexico.
70. SCIURUS SOCIALIS SOCIALIS, Wagner
1837. -Abh. math.-phys. CI. k. bayer. .\kad. Wiss. Munchen, II, p. 504.
Near Tehuantepec City, Oaxaca, Mexico.
71. SCIURUS SOCIALIS COCOS, Nelson
1898. Proc. Biol. Soc. Washington, XII, p. 155.
Acapulco, Guerrero, Me.xico.
72. SCIURUS SOCIALIS LITTOR.'^LIS, Nelson
1907. Proc. Biol. Soc. Washington. XX, p. 87.
Puerta .Angel, Oaxaca, Mexico.
73- SCIURUS GRISEOFLAVUS GRISEOFLAVUS, Gray
1867. Ann. Mag. Nat. Hist. 3, XX, p. 427.
Guatemala.
74. SCIURUS GRISEOFL.WUS CHIAPENSIS, Nelson
1899. Proc. Washington Acad. Sci.. I, p. 69.
San Cristobal, Chiapas, .Mexico.
336 SCIURUS
75. SCIURUS YUCATANENSIS YUCATANENSIS, Allen
1S77. Monogr. N. Amer. Rodents, p. 705.
Merida, Yutatan, Mexico.
7b. SCIURUS YUCATANENSIS BALIOLUS, Nelson
igoi. Proc. Biol. Soc. Washington, XIV, p. 131.
Apazote, Campechc, Mexico.
77. SCIURL'S YUCATANENSIS PHAEOPUS, Goodwin
1932. Amer. Mus. Nov. 574, p. i.
Guatemala, .Secanquim, district of Alta Vcrapaz.
78. SCIURUS VARIEGATOIDES VARIEGATOIDES, Ogilby
1S39. Proc. Zool. Soc. London, p. 117.
San Salvador, Central America.
Synonym : pj'/oJo, Lesson, 1842, Rev. Zool. Paris, V, p. 130.
(For a revision of 5. variegatoides and its subspecies see Harris,
1937, Misc. Publ. 38, Univ. Michigan.)
79. SCIURUS VARIEGATOIDES UNDERWOODI, Goldman
1932. Joum. Washington, Acad. Sci. XXII, p. 275.
Honduras ; Monte Redondo, 30 miles north-west of Tegucigalpa.
80. SCIURUS VARIEGATOIDES GOLDMANI, Nelson
1S98. Proc. Biol. Soc. Washington, XII, p. 149.
Huehuetan, Chiapas, Mexico.
81. SCIURUS VARIEGATOIDES BANGSI, Dickey
1928. Proc. Biol. Soc. Washington, XLI, p. 7.
Barra de Santiago, Dept. Ahuachapan, San Salvador.
82. SCIURUS VARIEGATOIDES BOOTHIAE, Gray
1843. List. Spec. Mamm. Brit. Mus., p. 139.
Honduras.
'nynonym : Juscovariegatus, Schinz, 1845, Syn. Mamm. 11, p. 15.
(?) boothiae amialium , Thomas, 1905, Ann. Mag. Nat. Hist., 7,
XVI, p. 309. Honduras.
83. SCIURUS V..\RIEGATOIDES BELTl, Nelson
1899. Proc. Washington, Acad. Sci. I, p. 78.
Escondido River, 50 miles above Bluefields, Nicaragua.
84. SCIURUS VARIEGATOIDES ADOLPHEI, Lesson
1842. Nouv. Tabl. Regn. Anim. Mamm., p. 112.
Realejo, Nicaragua.
85. SCIURUS VARIEGATOIDES MANAGUENSIS, Nelson
189S. Proc. Biol. Soc. Washington, XII, p. 150.
Managua River, Guatemala.
86. SCIURUS VARIEGATOIDES AUSTINI, Harris
1933. Occ. Pap. Mus. Zool. Univ. Michigan, 266, p. i.
Costa Rica: Las Agujas, Prov. of Puntarenas.
87. SCIURUS VARIEGATOIDES ATRIRUFUS, Harris
1930. Occ. Pap. Mus. Zool. LTniv. Michigan, 219, p. 2.
Costa Rica: Tambor, Nicoya Peninsula.
88. SCIURUS VARIEGATOIDES DORSALIS, Gray
184S. Proc. Zool. Soc. London, p. 138.
"Caracas. Venezuela" (erroneous); Liberia, Costa Rica.
Synonym: intermedins, Ciray, 1867, Ann. Mag. Nat. Hist. 3, XX, p. 421,
and iiicoyoiia, Gray, same reference, p. 423.
SCIURUS ' 337
89. SCIURUS VARIEGATOIDES RIGIDUS, Peters
1863. Monatsber. k. preuss. Akad. Berlin, p. 652.
San JosiS, Costa Rica.
90. SCIUKUS VARIEGATOIDKS THOMASI, Nelson
1899. Proc. WashinRton Acad. Sci. I, p. 71.
Talamanca, Costa Rica.
91. SCIURUS VARIEGATOIDF.S MELANIA, Gray
1867. Ann. Mac. Nat. Hist. 3, XX, p. 425.
Point Berica, Costa Rica.
92. SCIURUS VARIEGATOIDES HELVEOLUS, Goldman
1912. Smiths. Misc. Coll. LVI, no. 36, p. 3.
Corozal, Canal zone, Panama.
deppei Group
93. SCIURUS DEPPEI DEPPEI, Peters
1863. Monatsber. k. preuss Akad. Wiss. Berlin, p. 654.
Bapantla, Vera Cruz, Mexico.
SynonjTn: tephrogaster. Gray, Ann. Mag. Nat. Hist. 3, XX, p. 431, 1867.
taeniurus. Gray, same reference.
94. SCIURUS DEPPEI MATAGALPAE, Allen
1908. Bull. Amer. Mus. Nat. Hist. XXIV, p. 660.
San Rafael del Norte, Nicaragua.
QS. SCIURUS DEPPEI VIV.'VX, Nelson
iQOi. Proc. Biol. Soc. Washington, XIV, p. 131.
Apazote, Campeche, Mexico.
96. SCIURUS NEGLIGENS, Nelson
i8g8. Proc. Biol. Soc. Washington, XII, p. 147.
Alta Mira, Tamaulipas, Mexico.
Subgenus Hesperosciurus, Nelson
97- SCIURUS GRISEUS GRISEUS, Ord
1818. Joum. de Phys., LXXXVII, p. 152.
The Dalles, Wasco County, Oregon.
Synonym : /ossor, Peale, 1848, Mamm. Birds. U.S. Explor. Exp., p. 55.
heerniamii, Leconte, Proc. Acad. Nat. Sci. Philadelphia,
p. 149. 1852.
leporiniis, Henshaw, Ann. Rep. Engin., 1876, p. 310.
98. SCIURUS GRISEUS ANTHONVI, Meams
1897. Proc. U.S. Nat. Mus. XX, p. 501, 1898.
Campbell's Ranch, Laguna Mountains, San Diego County', California.
99- SCIURUS GRISEUS NIGRIPES, Br\ant
1889. Proc. Calif. Acad. Sci. 2, II, p. 25.
Coast region of San Mateo County, California.
Subgenus Otosciurus, Nelson
100. SCIURUS ABERTI ABERTI, Woodhouse
1853. Proc. Acad. Nat. Sci. Philadelphia, VI, 1852, p. 220.
San Francisco Mountain, Coconino County, .■Arizona.
Synonym: castanotus, Baird, 1855, Proc. Acad. Nat. Sci. Philadelphia,
VII, p. 332. Copper mines. New Mexico.
22 — Living Rodents — I
338 SCIURUS
loi. SCIURUS ABERTl BARBERI, Allen
1904. Bull. Amer. Mus. Nat. Hist. XX, p. 207.
Colonia Garcia, Chihuahua, Mexico.
102. SCIURUS ABERTl FERREUS, True
iQoo. Proc. Biol. See. Washington, XIII, p. 183.
Loveland, Larimer County, Colorado.
Synonym: concolor. True, 1894, Diagnoses of new N. Amer. Mamm.
p. I. Reprinted Proc. U.S. Nat. IVIus. XVII, p. 241.
(Preoccupied.)
103. SCIURUS ABERTl MIMUS, Merriam
1Q04. Proc. Biol. Soc. Washington, XVII, p. 130.
Hall Peak, Cimarron Mountains, Mora County, New Me.\ico.
104. SCIURUS ABERTl PHAEURUS, Allen
1904. Bull. Amer. Mus. Nat. Hist., XX, p. 205.
La Cienega, N.-W. Durango, Mexico.
105. SCIURUS ABERTl CHUSCENSIS, Goldman
1931. Proc. Biol. Soc. Washington, XLIV, p. 133.
N.-W. New Mexico: Chusca Mountains.
loh. SCIURUS KAIBABENSIS, Merriam
1904. Proc. Biol. Soc. Washington, XVII, p. 129.
Bright Angel Creek, Kaibab Plateau, north side of Grand Canyon of
Colorado, Coconino Co., Arizona.
107. SCIURUS DURANGI, Thomas
1893. Ann. Mag. Nat. Hist. 6, XI, p. 50.
Ciudad Ranch, 100 miles west of Durango City, Durango, Mexico.
Subgenus Parasciunis, 'I'rouessart
(Synonym : Araeoscitirus, Nelson.)
108. SCIURUS NIGER NIGER, Linnaeus
1758. Syst. Nat., Ed. 10, I, p. 64.
South Carolina.
Synonym: vutpiims, Gmelin, 1788, Gni. Syst. Nat. i, p. 147
109. SCIURUS NIGER AVICENNIA, Howell
1919. Journ. Mamm. Baltimore, i, p. 37.
Everglade, Lee County, Plorida.
no. SCIURUS NIGER TEXIANUS, Bachman
1838. Proc. Zool. Soc. London, p. 86.
Coast of Louisiana or Mississippi.
111. SCIURUS NIGER NEGLECTUS, Gray
1867 Ann. Mag. Nat. Hist. 3, XX, p. 425.
Wilmington, Delaware (Newcastle County).
Synonym: cinereus. True, Proc. U.S. Nat. Mus. VII, p. 595. (18S4).
vicinus, Bangs, 1896, Proc. Biol. Soc. Washington, X, p. 150.
(West Virginia.)
112. SCIURUS NIGER BRYANTI, Bailey
1920. Bailey Mus. & Libr. Nat. Hist. Newport News, Va. Bull, i, p. i.
Dorchester County, Maryland.
SCIURUS 339
113. SCIURUS NIGER RUFIVENTER, Geoffrey
1803. Cat. Mamm. Mus. Nat, Hist. Paris, p. 176.
Mississippi Valley.
Synonym: hidoviciamis , Custis, 1806, Burtons Med. & Phys. Joum. 2,
pt. 2. p. 47.
ruber, Rafincsquc, 1820, Ann. of Nat. p. 4.
macroura, Say, Longs Exp. Rocky Mtns. i, p. 115, 1823.
magnicaudiitus, Harlan, Faun. Amer. p. 178, 1825.
(?) subauratiis, Bachman, Proc. Zool. Soc. London, 1838,
p. 87.
(?) audubonii, Bachman, Proc. Zool. Soc, London, 1838,
p. 97.
rubicaudalus, Aud. & Bach. Quadr. N. America II, p. 30,
1851.
sayii, Aud. & Bach. Quadr. N. America II, p. 274, 1851.
atroventris, Engelmann, Trans. Ac. Sci. St. Louis, I, p. 329,
1859.
114. SCIURUS NIGER LIMITIS, Baird
1855. Proc. Acad. Nat. Sci. Philadelphia, VII, p. 331.
Devil's River, Valverde County, Texas.
115. SCIURUS OCULATUS OCUL-^TUS, Peters
1863. Monatsber. k. -Akad. Wiss. Berlin, p. 653.
Mexico, probably near Las Vegas, Vera Cruz.
Synonym: niger melanotus, Thomas, 1890, Proc. Zool. Soc. London,
p. 73. Las Vegas, Vera Cruz, Mexico.
capistratus. Lichtenstein, .\h. Akad. k. Wiss. Berlin, 116,
1830 (1827). (Preoccupied.)
1 16. SCIURUS OCULATUS TOLUCAE. Nelson
1898. Proc. Biol. Soc. Washington, XIT, p. 148.
North slope of Volcano of Toluca, State of Mexico, Mexico.
117. SCIURUS ALLENI, Nelson
1898. Proc. Biol. Soc. Washington, XII, p. 147.
Monterey, Nuevo Leon, Mexico.
118. SCIURUS NAYARITENSIS, Allen
1890. Bull. .Amer. Mus. Nat. Hist., II, p. vii, footnote.
Sierra Valparaiso, Zacatecas, Mexico.
Synonym: a/stoni, Allen, 1889, Bull. .Amer. Mus. Nat. Hist. II, p. 167.
(Not of .Anderson, 1878.)
119. SCIURUS APACHE, Allen
1893. Bull. Amer. Mus. Nat. Hist. V, p. 29.
N.-W. Chihuahua, Mexico.
120. SCIURUS CHIRICAHUAE, Goldman
1933. Proc. Biol. Soc. Washington, XLVI, p. 71.
Chiricahua Mountains, Cochise County, Arizona.
121. SCIURUS ARIZONENSIS ARIZONENSIS, Coues
1867. .Amer. Nat. I, p. 357.
Fort Whipple, Yavapai County, Arizona.
122. SCIURUS ARIZONENSIS HUACHUCA, Allen
1894. Bull. Amer. Mus. Nat. Hist. VI, p. 349.
Huachuca Mountains, Southern Arizona.
340 SCIURUS
123. SCIURUS ARIZONENSIS CATALINAE, Doutt
1931. Ann. Carn. Mus. 20. p. 271.
Santa Catalina Mountains, Arizona.
Subgenus Giiciliiiqiietiis, Gray
(Synonym: Mcsosciiirtts, Allen.
Histriosciurus , .Allen.
Macroxus, Cuvier.
Simosciurus, Allen.
Leptosciurus, .Allen.)
hoffmani Group
124. SCIURUS HOFFM.ANI HOFFM.ANI, Peters
1863. Monatsber. k. Akad. Wiss. Berlin, p. 654.
Costa Rica.
Svnonvm: xanthotis. Gray, 1867, Ann. Mag. Nat. Hist. 3, XX, p. 429.
rufoniger, True, Proc. U.S. Nat. Mus, 1884, VII, p. 595-
125. SCIURUS HOFFM.ANI CHIRIQUENSIS, Bangs
1902. Bull. Comp. Zool. XXXIX, no. 2, p. 22.
Divala, Chiriqui, Panama.
I2b. -SCIURUS HOFFM.A.NI M.-\XAVI, .-Mien
1914. Bull. .Amer. Mus. Nat. Hist. XXXIII, p. 589.
Manavi, Rio de Oro, Ecuador.
127. SCIURUS HOFFMANI QUINDIANUS, Allen
1914. Bull. .Amer. Mus. Nat. Hist. XXXIII, p. 5^7.
Rio Frio, Central Andes. Colombia.
12S. SCIURUS HOFFMANI HYPORRHODUS, Gray
1867. .Ann. Mag. Nat. Hist. 3, XX, p. 419-
Bogota, Colombia.
I2C5. SCIURUS HOFFMANI SODERSTROMl, Stone
1914. Proc. .Acad. Sci. Philadelphia, LXVI, p. 14.
Ecuador.
130. SCIURUS MIRAVALLENSIS, Harris
193 1. Occ. Pap. Zool. Mus. Univ. Michigan, 227, p. i.
Volcan de Miravalles, Costa Rica.
131. SCIURUS RICHMONDI, Nelson
1898. Proc. Biol. Soc. Washington, XII, p. 146.
Escondido River, 50 miles above Bluetields, Nicaragua.
132. SCIURUS GRISEOGENA GRISEOGENA, Gray
1867. -Ann. Mag. Nat. Hist. 3, XX, p. 4^9-
Venezuela. c ^.-iir
Synonym: klagesi, Thomas, 1914. Ann. Mag. Nat. Hist. 8, XI\ , p. 240.
Near Caracas, Venezuela.
133. SCIURUS GRISEOGENA MERIDENSIS, Thomas
1901. .Ann. Mag. Nat. Hist. 7, VII, p. 192-
Escorial, Sierra de Merida, Venezuela.
Svnonym: tamac, Osgood, 1912. Field Mus. Nat. Hist. Zool. Ser. X,
no. 5, p. 48. Paramo de Tama, Colombia-Venezuela
boundary.
SCIURUS 341
134. SCIURUS CHAPMANI CHAPMANI, Allen
1899. Bull. Amer. Mus. Xat. Hist. XII, p. 16.
Caparo, Trinidad.
Synonym : aestuans quebradensis, Allen, 1899, Bull. Amer. Mus. Nat. Hist.
XII, p. 217. Quebrada Secca, Venezuela.
135. SCIURUS CHAPMANI TOBAGENSIS, Osgood
1910. Field Mus. Nat. Hist. Zool. Ser. X, no. 4, p. 27.
Tobago, West Indies.
136. SCIURUS NESAEUS, G. Allen
1902. Proc. Biol. Soc. Washington, XV, p. 93.
Margarita Island, Venezuela.
137. SCIURUS GRISEIMEMBRA, Allen
1914. Bull. Amer. Mus. Nat. Hist. XXXIII, p. 589.
Buenavista, Eastern Andes, Colombia.
138. SCIURUS CANDELENSIS C.A.NDELEXSIS, Allen
1914. Bull. Amer. Mus. Nat. Hist., XXXIII, p. 590.
Huila, Colombia.
139. SCIURUS CANDELENSIS SUNUCO, Cabrera
1917. Trab. Mus. Nac. Ci. Nat. 31, p. 51.
San Jose, East Ecuador.
140. SCIURUS ARGENTINUS, Thomas
1921. Ann. Mag. Nat. Hist. 9, VIII, p. 609.
Jujuy, North Argentina.
141. SCIURUS FERMINAE, Cabrera
1917. Trab. Mus. Nac. Ci. Nat. Madrid, 31, p. 49.
Bueza, East Ecuador.
142. SCIURUS GERRARDI GERR.ARDI, Gray
1 86 1. Proc. Zool. Soc. London, p. 92, pi. XVI.
"New Grenada," probably Medellin, Colombia.
S\Tionym: variabilis, .Alston, 1878, Proc. Zool. Soc. London, p. 665,
not of Geoffrey.
143. SCIURUS GERRARDI LEONIS, Lawrence
I933- Joum. Manim. Baltimore, 14, p. 369.
Colombia.
SynonNTii: milleri, Allen, 19 12, Bull. .\mer. Mus. Nat. Hist. XXXI, p. 91.
(Preoccupied.) Cocal, W. Colombia.
144- SCIURUS GERRARDI INCONSTANS, Osgood
1921. Journ. Mamm. Baltimore, 2, p. 40.
Ecuador.
Synon\TTi: versicolor, Thomas. 1900, .Ann. Mag. Nat. Hist. 7, VI, p. 385.
(Preoccupied.) Cachabi, Prov. Esmeraldas, N. Ecuador.
145- SCIURUS GERR.ARDI MORULUS, Bangs
1900. Proc. New England Zool. Club, II, p. 43.
Loma del Leon, Panama.
146. SCIURUS GERRARDI CHOCO, Goldman
1913. Smiths. Misc. Coll. LX, no. 22, p. 4. '
Cana, Pirri Mountains, Eastern Panama.
34^ SCIURUS
147- SCIURUS GERKARDI SALAQUENSIS, Allen
1Q14. Bull. Amer. Mus. Nat. Hist. XXXIII, p. 592.
Rio Salaqui, N.-W. Colombia.
148. SCIURUS GERRARDI ZULIAE, Osgood
1910. Field Mus. Nat. Hist. Zool. Ser. X, 4, p. 26.
Zulia, Venezuela.
149. SCIURUS GERRARDI CUCUTAE, Allen
1914. 13ull. Amer. Mus. Nat. Hist. XXXIII, p. 592.
El Ciuayabal, Colombia.
150. SCIURUS GERRARDI BAUDENSIS, Allen
1915. Bull. Amer. Mus. Nat. Hist. XXXIV, p. 30S.
Baudo, West Colombia.
151. SCIURUS GERRARDI VALDIVIAE, Allen
1915. Bull. Amer. Mus. Nat. Hist., XXXIV, p. 309.
Puerto Valdivia, Colombia.
152. SCIURUS SPLENDIDUS SPLENDIDUS, Gray
1S42. Ann. Mag. Nat. Hist, i, X, p. 263.
Santa Marta, Colombia.
Synonym: saltuensis magdaleiiae, Allen, 1914, Bull. Amer. Mus, Nat.
Hist. XXXIII, p. 593. Rio ^Iagdalena, Colombia.
153. SCIURUS SPLENDIDUS SALTUENSIS, Bangs
1898. Proc. Biol. Soc. Washington. XII, p. 1S5.
Santa Marta, Colciimbia.
154. SCIURUS SPLENDIDUS BONDAE, Allen
1899. Bull. Amer. Mus. Nat. Hist. XII, p. 213.
Bonda, Santa Marta, Colombia.
155. SCIURUS PYRRHINUS, Thomas
1S98. Ann. Mag. Nat. Hist. 7. II. p. 265.
Vitoc, Peru.
aestuaiis Group
156. SCIURUS AESTUANS AESTUANS, Linnaeus
1766. Syst. Nat. Ed. 12, i, p. 88.
Surinam.
Synonym: kiihlii, Gray,' 1867, Ann. Mag. Nat. Hist. 3, XX, p. 433.
giiianemis, Peters, 1863, Monatsber. Akad. Wiss. Berlin,
p. 655.
157. SCIURUS AESTUANS GILVIGULARIS. Wagner
1843. Archiv. f. Naturg. ii, p. 43.
Borba, Brazil, near mouth of Rio Madeira.
15S. SCIURUS AESTUANS MACCONNELLl, Thomas
190!. Ann. Mag. Nat. Hist. 7, VIII, p. 14S. footnote.
Mt. Roraima, British Guiana.
159. SCIURUS AESTUANS QUELCHI, Thomas
1901. Ann. Mag. Nat. Hist. 7, VIII, p. 147.
Kanuku Mountains, British Guiana.
160. SCIURUS AESTUANS VENUSTUS, Allen
1915. Bull. Amer. Mus. Nat. Hist., XXXIV, p. 260.
Rio Cunacunuma, near Mt. Duida, Venezuela.
' See note on p. 318.
SCIURUS 343
i6i. SCIURUS AESTUANS GARBEI, Pinto
1931. Rev. Mus. Paulista, XVII, p. 294.
Esperito Santo, Bahia, Brazil.
162. SCIURUS ALPHONSEI ALPHONSEI, Thomas
1906. Ann. Mag. Nat. Hist. 7, XVIII, p. 442.
Pemambuco, Brazil.
Synonym: roberti, Thomas, 1903, Ann. Mag. Nat. Hist. 7, XII, p. 463.
(Not of Bonhote). S. Lourenfo, near Pemambuco.
163. SCIURUS ALPHONSEI PARAENSIS, Goeldi
1904. Bol. Mus. Goeldi, IV, p. 70.
Para, Brazil.
164. SCIURUS INGRAMI, Thomas
1901. Ann. Mag. Nat. Hist. 7, VII, p. 368.
Tunnel, Southern Minas Geraes, Brazil.
stramineus Group
165. SCIURUS STR.\MIN'EUS STRAMINEUS, Eydoux & Souleyet
1841. Voy. Bonite, Zool. I, p. 38, pi. IX.
Omatope, Peru.
Synonym: fraseri, Gray, 1867, .\nn. Mag. Nat. Hist. 3, XX, p. 430.
Ecuador.
166. SCIURUS STR.A.MINEUS NEBOUXI, Geoffroy
1855. Voy. de la Venus, Zool. p. 163, pi. xii.
Near Pa>ta, Peru.
167. SCIURUS STRAMINEUS GUAYANUS, Thomas
1900. .Ann. Mag. Nat. Hist. 7, V, p. 150.
Balzar Mountains, West Ecuador.
168. SCIURUS STRAMINEUS ZARUM.AE, .Allen
1914. Bull. Amer. Mus. Nat. Hist., XXXIII, p. 597.
Zaruma, S.-W. Ecuador.
pucherani Group
169. SCIURUS PUCHERANI PUCHERANI, Fitzinger
1867. Sitz.-Ber. .Akad. Wiss. Wien Math. Nat. CI. LV, Abth. i, p. 487.
Vicinity of Bogota, Colombia.
Synonym: rufoniger, Pucheran, 1845, Rev. Zool. VIII, p. 336.
chrysuros, Pucheran, same reference, p. 337.
170. SCIURUS PUCHERANI MEDELLINENSIS, Gray
1872. Ann. Mag. Nat. Hist. 4, X, p. 408.
MedcUin, Colombia.
i-i. SCIURUS PUCHERANI CAUCE.NSIS, Nelson
1899. Bull. .Amer. Mus. Nat. Hist. XII, p. 79.
San Antonio, Western Andes, Colombia.
172. SCIURUS PUCHERANI SALENTENSIS, AUen
1914- Bull. .Amer. .Mus. Nat. Hist. XXXIII, p. 587.
Near Salento. Central .Andes, Colombia.
344 ■ SCIURUS
17.V SCIURUS IGNITUS IGNITUS, Gray
1867. Ann. Mag. Nat. Hist. 3, XX, p. 429.
Yunyas, upper Rio Beni, Bolivia.
-Svnonym: oihresceiis, Thomas, 1914, Ann. Mag. Nat. Hist. 8, XIII,
p. 362. Upper Beni River, Bolivia.
cuscmiis, Thomas, 1899, Ann. Mag. Nat. Hist. 7, III, p. 40.
Ocabamba, Cuzco, Peru.
174. SCIURUS IGNITUS IRRORATUS, Gray
1867. Ann. Mag. Nat. Hist. 3, XX, p. 431.
Upper Rio Ucayali, Peru.
175. SCIURUS BOLIVIENSIS, Osgood
1921. Journ. Mamm. Baltimore, 2, p. 39.
Santa Cruz de la Sierra, Bolivia.
Synonym: leucogaster. Gray, 1867, Ann. Mag. Nat. Hist. 3, XX, p. 430.
(Preoccupied.)
Subgenus Notosciurus, .'Mien
17(1. .SCIURUS RHO.\DSI, Allen
1914. Bull. .-Vmer. Mus. Nat. Hist. XXXIII, p. 585.
Pagnia Forest, Chunchi, Ecuador.
Subgenus Hadrosciurus, Allen
(Synonym: Urosciurus, Allen)
177. SCIURUS FLAMMIFER, Thomas
1904. .Ann. Mag. Nat. Hist. 7, XIV, p. 33.
Caura district. Middle Orinoco, Venezuela.
17S. SCIURUS TRICOLOR, Poeppig
1844. Tschudi Fauna Peruana, I, Therologie, p. 156, pi. xi.
North-east Peru.
Synonym: 0) futiiigatus. Gray, 1867, Ann. Mag. Nat. Hist., 3, XX,
p. 428.
brimnconigcr. Gray, same reference, p. 429.
179. SCIURUS NICiRATUS, Pinto
1931. Rev. Mus. Paulisto, XVII, p. 309.
Rio Jurua, .Amazon.
iSo. SCIURUS DUIDA, .Allen
1914. Bull. An-cr. Mus. Nat. Hist., XXXIII, p. 594-
Rio Cunucunuma, south of Mt. Duida, Venezuela.
181. SCIURUS IGNIVENTRIS IGNIVENTRIS, Wagner
1S42. Wiegmanns Arch. f. Naturgesch. I, p. 360.
Upper Rio Negro, Brazil.
Synonym: morio, Wagner, Abh. Math. Phys. CI. K. B. Akad. Wiss.
Miinchen, V, 1850, p. 275.
1.S2. SCIURUS IGNIVENTRIS TAEDIFER, Thomas
1903. -Ann. Mag. Nat. Hist. 7, XI, p. 487.
50 miles south-east of Bogota, Colombia.
SCIURUS— TAMIASCIURUS 345
183. SCIURUS IGNIVENTRIS COCALIS. Thomas
1900. Ann. Mag. Nat. Hist. 7, VI, p. 138.
Upper Rio Xapo, Ecuador.
184. SCIURUS IGNIVENTRIS ZAMORAE, Allen
1914. Bull. .\mer. Mus. Nat. Hist., XXXIII, p. 594.
Zamora, Ecuador.
185. SCIURUS IGNIVENTRIS FUI.MIN..\TUS, Thomas
1926. Ann. Mag. Nat. Hist. 9, XVII, p. 637.
Manacapuru, Lower Rio Negro, Brazil.
186. SCIURUS PYRRHONOTUS PYRRHONOTUS, Wagner
1842. Wiegmanns .Archiv. f. Naturgesch. i, p. 360.
Borba, near mouth of Rio Madeira, Brazil.
187. SCIURUS PYRRHONOTUS TAP.A.RIUS, Thomas
1926. Ann. Mag. Nat. Hist, g, XVII, p. 635.
Santarcm, Lower .Amazons.
i88. SCIURUS PYRRHONOTUS CASTUS, Thomas
1903. .Ann. Mag. Nat. Hist. 7, XI, p. 488.
Chimate, Upper Rio Beni, Bolivia.
189. SCIURUS PYRRHONOTUS JUR,\LIS, Thomas
1926. .Ann. Mag. Nat. Hist. 9, XVII, p. 636.
Jurua River, Upper Amazons.
190. SCIURUS LANGSDORFFI LANGSDORFFI, Brandt
1835. Mem. .Acad. Sci. St. Petersb. 6, Math. Phys. Nat. Ill, 2, p. 425, pi. xi.
Cuyaba, Matto Grosso, Brazil.
191. SCIURUS LANGSDORFFI URUCUMUS, Allen
1914. Bull. .Amer. Mus. Nat. Hist.. XXXIII, p. 595.
Urucum, Rio Paraguay, Matto Grosso.
192. SCIURUS LANGSDORFFI STEINBACHI, Allen
1914. Bull. Amer. Mus. Nat. Hist., XXXIII, p. 596.
Santa Cruz de la Sierra, Bolivia.
Genus 7. TAMIASCIURUS, Trouessart
1880. Ta-MIASCiurvs, Trouessart, le Naturaliste, 2, no. 37, p. 292.
Type Species. — Sciurus hudsonicus, Erxleben.
Range. — North America: Canada and U.S.A.; forms named from Alaska,
Hudson Bay, Britisli Columbia, Mackenzie, Washington, Oregon,
Idaho, Wyoming, South Dakota, Minnesota, California, Lower California,
Arizona, New Mexico, North Carolina, Alaine, Connecticut.
Number of For.ms. — Twenty-seven.
Characters. — The generic difference between this and Sciurus is the com-
plete suppression of the baculum (Pocock, TuUberg, Howell),
though whether all the forms have been examined in this respect 1 do not know.
It appears, from Tullberg's notes on genera throughout the whole Order,
to be a sufficiently rare character in Rodents on which to warrant the retention
of generic names.
346 TAMIASCIURUS
Skull characters much like normal Sciiirus; the parietal ridges may join;
bullae relatively more enlarged than is usual in Sciunis; braincase not strongly
deflected posteriorly; palate extending slightly behind toothrows. Cheekteeth
near Sciuriis nilgai is in type; P. 3 may be present or absent; in all skulls but
two examined in the British Museum (a very small scries) it is absent; Allen
in his revision of tlie genus, 1S98, remarks that it is absent in about 30 per cent
of those examined.
External characters rather reminiscent of Sciuriis viilg<iris; tail relatively
short (40 per cent total length, Allen); ears tufted seasonally; sole hairy in
most skins seen, also evidently a seasonal character. Digits normal (arboreal
type).
The complete or almost complete suppression of the baculum is also found
in the African genus Heliosciiiriis.
Forms seen : albolimbatiis, douglasi, fremonti, hudsoniciis, loqiuix, ricliardsoni,
vancoiiverensis.
The species and races were revised by Allen, Bull. .Amer. Mus. Nat. Hist.
XX, pp. 249-29S, 1898.
List of Named Forms
1. TAMIASCIURUS HUDSONICUS HUDSONICUS, Erxleben
1777. Syst. Regn. Anim. i, p. 416.
Hudson Strait.
Synonym: ruhroUneatiis. Desmarest, Mamm. II, p. 333, 1S22.
2. TAMIASCIURUS HUDSONICUS GYMNICUS, Bangs
1899. Proc. New England Zool. Club, i, p. 28.
Cireenville, near Moosehead Lake, Maine. (Piscataquis County.)
3. TAMIASCIURUS HUDSONICUS LOQUAX, Bangs
1896. Proc. Biol. Soc. Washington, X, p. 161.
Liberty Hill, New London County, Connecticut.
4. TAMIASCIURUS HUDSONICUS MINNESOTA, Allen
1899. Amer. Nat. XXXIII, p. 640.
Fort Snelling, Hennepin County, Minnesota.
5. TAMIASCIURUS HUDSONICUS D.'\KOTENSIS, Allen
1894. Bull. Amer. Mus. Nat. Hist. VI, p. 325.
Squaw Creek, Black Hills, Custer County, South Dakota.
6. T.AMIASCIURUS HUDSONICUS BAILEYI, Allen
1895. Bull. .Amer. Mus. Nat. Hist. X, p. 261.
Bighorn Mountains, Washakie County, Wyoming.
7. TAMIASCIURUS HUDSONICUS VENTORUM, Allen
1S98. Bull. Amer. Mus. Nat. Hist. X, p. 263.
South Pass City, Wind River Mountains, Fremont County, Wyoming.
8. TAMIASCIURUS HUDSONICUS RICHARDSONI, Bachman
1838. Proc. Zool. Soc. London, p. 100.
Head of Big Lost River, Fremont County, Idaho.
<). TAMIASCIURUS HUDSONICUS STREATORI, Allen
189S. Bull. Amer. Mus. Nat. Hist. X, p. 267.
Ducks, British Columbia, Canada.
TAMIASCIURUS 347
10. TAMIASCIURUS HUDSONICUS VANCOUVERENSIS, Allen
1890. Hull. Amer. Mus. Nat. Hist. Ill, p. 165.
Duncan Station, Vancouver Island, British Columbia.
11. TAMIASCIURUS HUDSONICUS PICATUS, Swarth
1921. Joum. Mamm. Baltimore, 2, p. 92.
Kupreanof Island, S.-E. Alaska, 25 miles south of Kake Village,
southern end of Keku Straits.
12. TAMIASCIURUS HUDSONICUS PETULANS, Osgood
1900. North Amer. Fauna, No. 19, p. 27.
Glacier, White Pass, Southern Alaska.
13. TAMIASCIURUS HUDSONICUS ABIETICOLA, Howell
1929. Joum. Mamm. Baltimore, 10, p. 75.
Highlands, N. Carolina.
14. TAMIASCIURUS HUDSONICUS COLUMBIENSIS, Howell
1936. Proc. Biol. Soc. Washington, XLIX, p. 135.
Raspberry Creek, about 30 miles south-east of Telegraph Creek,
Northern British Columbia.
15. TAMIASCIURUS HUDSONICUS KENAIENSIS, Howell
1936. Proc. Biol. Soc. Washington, XLIX, p. 136.
Hope, Cook Inlet, Alaska.
16. TAMIASCIURUS HUDSONICUS PREBLIEI, Howell
1936. Proc. Biol. Soc. Washington, XLIX, p. 133.
Fort Simpson, Mackenzie, Canada.
17. TAMIASCIURUS REGALIS, Howell
1936. Occ. Pap. Mus. Univ. Mich. no. 338, p. i.
Belle Isle, Isle Royale, Michigan.
18. T.i^MIASCIURUS DOUGLASII DOUGLASII, Bachman
1838. Proc. Zool. Soc. London, p. 99.
Near mouth of Columbia River.
Synonym: belcheri. Gray, Ann. Mag. Nat. Hist. 1842. p. 263.
mckleyi, Baird. Pr. A. Phil. 1855, p. 333.
19. TAMIASCIURUS DOUGLASII MOLLIPILOSUS, Audubon & Bachman
1841. Proc. .^cad. Nat. Sci. Philadelphia, i, p. 102.
Coast of Northern California.
Synonym : orarius. Bangs, 1897, Proc. Biol. Soc. Washington, XI, p. 281.
Philo, Mendocino Co., California.
20. TAMIASCIURUS DOUGLASII CASCADENSIS, Allen
i8g8. Bull. Amer. Mus. Nat. Hist. X, p. 277.
Mount Hood, Oregon.
(According to Osgood, 1907, this will probably stand as lanuginosus,
Bachman, 1S38, Proc. Zool. Soc. London, p. loi. Hunter
Island, British Columbia.)
21. TAMIASCIURUS DOUGLASII ALBOLIMBATUS, Allen
1898. Bull. Amer. Mus. Nat. Hist. X, p. 453.
Blue Canyon, Placer County, California.
SynonjTn: califomicus, .\llen, 1890, Bull. .Amer. Mus. Nat. Hist. Ill,
p. 165. (Not of Lesson.)
34S TAMIASCIURUS— CALLOSCIURUS
22. TAMIASCIURUS DOUGLASII MEARNSI, Townsend
iSgy. Proc. Biol. Sue. Washington, XI, p. 146.
San Pedro iVlartir Mountains, Lower California.
23. TAMIASCIURUS FREMONTI FREMONTI, Audubon & Ijachman
1S54. Qiiadr. N. .-Xmer. 3, p. 237.
"Rocky Mountains," probably in Park region of Central Colorado.
24. TAMIASCIURUS FREMONTI NEOMEXICANUS, Allen
1S98. Bull. .\mer. Mus. Nat. Hist. X, p. 291.
Rayado Canyon, Colfax Co., New Mexico.
25. TAMIASCIURUS FREMONTI LYCHNUCHUS, Stone & Rehn
1903. Proc. Acad. Nat. Sci. Philadelphia, p. 18.
Forks of Ruidoso, Lincoln County, New Mexico.
26. TAMIASCIURUS FREMONTI MOGOLLONENSIS, Mearns
i8go. -Auk, vol. 7, p. 49. Bull. Amer. Mus. Nat. Hist. II, p. 277.
Quaking Asp Settlement, summit of Mogollon Mtns., Yavapai County,
Arizona.
27. T.AMIASCIURUS FREMONTI GRAHAMENSIS, Allen
1894. Bull. .^nier. Mus. Nat. Hist. VI, p. 350.
Graham Mountains, Graham Co., Arizona.
Genus 8. CALLOSCIURUS, Gray
1867. CALLOSCIURUS, Gray, Ann. Mag. Nat. Hist. 3, XX, p. 277.
18S0. Heterosciurus, Trouessart, le Naturaliste, i, p. 290. (Sciiiriis trvthraeiis,
Pallas.)
1915. ToMEUTEs, Thomas, Ann. Mag. Nat. Hist. 8, XV, p. 3S5. (Sch(nis lokroides,
Hodgson.)
1906. T.\,Miops, Allen, Bull. Amer. Mus, Nat. Hist., XXII, p. 475. (Scitints machllandi,
Horsfield.) Valid as a subgenus.
Type Species. — Sciurus rafflesii, Vigors & Horsfield.
Range. — From Tibet (subgenus Tamiops), C'hihli {subgenus Tamiops),
Szechuan, China south of the Yangtsekiang; Hainan, Formosa;
Bengal, Nepal, Sikkim, Assam, Burma, Siam, south through Malay region to
Sumatra, Java, Borneo, Celebes and the Philippine Islands.
Number of Forms. — Appro.ximately three hundred and twenty. This
genus is second in number of named forms to
Ratlus only in the Order.
Cn.\RACTERS. — This genus was originally divided from Sciurus by Thomas
on the structure of the baculum. I have already remarked
that this seems usually a very questionable character on which to base generic
names, particularly on account of the relatively few forms in which this character
has been definitely verified. Thomas wrote: "All Indian and Malay species
hitherto referred to Sciurus have bacula totally ditferent from true Sciurus, and
themselves arc di\isible into two types, with an essential community between
the two." Exactly how many forms have been examined on this character I do
not know, though some months ago I endeavoured to make a list of those I had
CALLOSCIURUS 349
come across which had been; there seems to be a constant wide distinction, as
far as one reads, between those Squirrels currently included in this group and
those currently referred to Sciurus which have been examined, in this character.
Also there is a certain difference in the teeth, not altogether constant, yet
noticeable in most of the leading species referred to this genus. M.3 in the
lower jaw is very often noticeably elongated, particularly on the inner side; the
anterointernal cusp in the lower teeth is as a rule very high; the subsidiary
small cusps on the outer side of the upper molars are verj' frequently almost
obsolete; the ridges of the upper molars are often higher and more definite in
this group; there is often a tendency towards complexity due to the presence of
extra small ridges and depressions on the crown; and the anterior extra ridge
cutting off a depression in the lower teeth and running from the anteroexternal
to the anterointernal cusp, characteristic of the genus Heliosciurus, is usually
present and well marked; further, P. 3 is strong as a rule, whereas in most mem-
bers of Sciurus it is vestigial. Also often, but not always, in the present genus
the upper incisors tend to become proodont.
The group is probably a natural one, though it is largely for convenience
that it is here regarded as a full genus distinct from Sciurus. According to
Thomas and Pocock, Tamiops which I refer to this genus as a subgenus, and the
closely allied Dremomys both agree in bacular characters with the " Tomeutes"
section of this genus. Tomeutes was erected as a genus by Thomas based solely
on the formation of the baculum. I have already commented on the inadvisa-
bility on p. 266 of retaining it. In the present state of our knowledge I do
not think it advisable to retain it even as a subgenus. Pocock, 1923, writing of
Callosciurus and Tomeutes, states that "distinct as the bacula of these two
kinds are, there are indications of intergradation between them," and "in
Tomeutes, the variation is so great that it is impossible to affirm any character
by which the baculum of Tomeutes can be distinguished from the bacula of
Dremomys, Rhinosciurus, Tamiops and Lariscus." This author states that the
species vittatus is a " Tomeutes." This species appears to me to be indistinguish-
able from notatus, or very questionably so, on cranial and external characters.
To refer it to a distinct subgenus seems absurd {notatus is a "Callosciurus" on
penial characters, so far as known). Furthermore there are fortv-sL\ named
subspecies of vittatus. It would be interesting to examine the baculum in all
of them. Probably it might be found that there is a complete intergradation in
the species alone, though this is sheer surmise. Therefore until verj- much more
work is done on this character in these Malay Squirrels, and bearing in mind
that the baculum e\-idently varies from local race to local race in Funambulus,
and in different species of Ratufa, all these Indo-Malayan Squirrels should be
referred to a single genus only.
The skull is in the group not essentially ditferent from Sciurus; in some
species there is a tendency for the rostrum to be slightly elongated ; the parietal
ridges often do not join, but in others quite a sharp crest is formed by them
(further notes on this character below). The cheekteeth are as described above;
the large internal cusp of the upper molars may be seen in some specimens to
be composed of three separate elements when cut. In C. prevosti and others
350 CALLOSCIURUS
the trontals appear unusually broad, and the postorbital processes are strongly
developed. The zygomatic plate is usually much as in Schinis, but in some
species there is a tendency for it to he more prominently ridged, a little inclined
to be narrowed above, and slanting upwards far forwards.
C. rubrivetiter, from Celebes, is the most distinct species seen; it appears
to be larger than all others; the cheekteeth are (in the two skulls seen) rather
worn, but simpler than is usual; the parietal ridges are very prominent, and
unite to form a sagittal ridge more or less close behind the postorbital process,
this ridge being much longer than in any other species seen. This form might
perhaps form a distinct subgenus.
C. leucomus, from Celebes, appears in two skulls seen to be almost transi-
tionary to the Nannosciurus-Sciwillus type of skull, with postorbital process
situated unusually far backwards, and lachrymal situated farther back than is
usual. Whether these characters are constant I do not know. A large collection
of Celebes Squirrels would be most welcome, as it is curious that out of only
about ten skulls examined from this island, every form seems aberrant and
different from the more normal Indo-AIalayan Squirrels; the species murinus
I have had to refer to the genus Sciurilhis, a member of the Naniiosciiirm section.
External characters as in normal Tree-squirrels; D.4 normally longer than
D.3 in manus. The colour, as might be expected, varies enormously throughout
the genus. In S. tenuis, and S. jentinki (small species), the tail is usually much
narrowed. Prominent ear tufts are present in leucomus, rubriventer and rosenbergi.
Tamiops was proposed as a genus by Allen and has usually been accepted.
The upper cheekteeth are, as often in Callosciurus, with the third outer cusp
(which is present in Sciunis) absent or vestigial. This, though the genus was
established mainly on this character, can hardly be regarded as of generic rank.
P. -5 is relatively large, as in CaUosciurus. The lower cheekteeth resemble Callo-
sciurus; M.3 is elongated to a degree, and the cusps are well marked. The skull
is not abnormal. The size is small. Back with conspicuous black stripes
(usually five in number, though these vary in development). Digits as in
normal Tree-squirrels, with D.4 in the manus normally slightly longer than
D.3. Tail usually much narrowed.
The group was compared with Sciurus only by Allen ; but unfortunately in
differentiating between this group and Sciurus on dental characters he has
merely repeated the condition usually found in Callosciurus. The narrow tail
turns up again in Callosciurus tenuis. The colour pattern alone remains of
Allen's characters, which in my opinion is not a valid generic character (see
p. 269).
A skeleton has been compared with Callosciurus skeleton, but does not
show any essential difference in general formation. The only difference appears
to be that in "Tamiops" there are 26 tail vertebrae, whereas in Callosciurus
tenuis there are only 23. But this is hardly a generic character; for instance, in
Ratufa indica, as quoted by Flower (Osteology, p. 85, 1885), there may be
either 24 or 25. Under these circumstances I have no alternative to reducing
Tamiops to a subgenus of the present genus.
Forms seen : alacris, albescens, alhivexilli, andrewsi, annellatus, aoris, aquilo,
CALLOSCIURUS 351
atratus, atrodorsalis, balstoni, buliiensis, bangueyae, barbei, bartoni, bellona, besuki,
bhutanensis, bilimitattis, " bilineatus," blanfordi, blythi, bocki, bocourti, bonliotei,
borneanus, brookei, caniceps, carevi, carimonensis, caroli, castaneoveniris, centralis,
chinensis, ciiinamomeiis, clarkei, cockerelli, concolor, condurenns, contiimax,
crotalius, criimpi, dactylinus, davisoni, dextralis, domelicus, dulitensis, erebus
( —piceus), erythraeus, epomophonis, erythrogaster, famulus, ferrugineus, finlaysoni,
flavimanus, floweri, "flumiiialis," folletti, formosanus, forresti, frandseni, frater-
culus, fryainis, germaini, gloveri, gordoni, grayi, griseicauda, griseitnanus, grisei-
pectus, gutiong, harmandi, harringtoni, hastilis, liendeei, hippurellus, hippurosus,
hippurus, hiimei, ictericus, imarius, imitator, inconstans, inquitiatus, janetta,
jentinki, juvencus, kinneari, kongensis, kuchingensis, lancavensis, hiotum, leucomus,
kucopNS, leucotis, lokroides, loii-i, lylei (race of bocourti), lylei {Tamiops, here
renamed holti), maclellandi, madurae, manipurensis, mapravis, maritimis, mearsi,
mehmogaster, menamicus, rnoitaui, " meticulusus," michianus, maporemis, micro-
rhynchiis, midas, millardi, milleri, miiidanensis, miniatus, moheius, mohillus, moi,
monticolus, nagartim, nakamis, natunensis, navigator, nesiotes, nigrovittatus,
ningpoerisis, notatus, nox, olivacetts, orestes, owensi, panjioli, panjius, pemangi-
lensis, peninsularis, perhentiani, phanrangis, phayrei, philippinensis, pierrei, pipi-
donus, pirata, plasticus, pluto, "portus," prevosti, proteus, pryeri, punctatissimus,
pygerythrus, quantulus, quinquestriatus, rafflesi, roberti, robinsoni, rodolphei,
rosenbergi, rubex, rubrirenter, rufoniger, rupatius, Sylvester, samarensis, sara-
wakensis, scotti, seimundi, shanicus, shortridgei, " siamensis," similis, singapurensis,
sinistralis, sladeni, sordidus, " splendidus," steerei, stevensi, styani, subluteus,
suffusus, sullivanus, siirdus, swinlioei, tabaudius, tacliardi, tacopius, tahan, taman-
sari, tapanulius, telibius, tenuirostris, tenuis, terutavensis, thaiwanensis, tiomanicus,
vanakeni, virgo, vittatus, watsoni, zuellsi, uilliamsoni, wrayi, youngi, simmeensis.
So far as subspecies are concerned, the classification of Robinson & Kloss,
1918 (Rec. Indian Mus. XV, pt. IV, pp. 171-250), is followed.
The forms in which the baculum structure is definitely verified, so far as
at present traced, are: maclellandi, prevosti, notatus, castaneozentris, atrodorsalis,
vittatus, lokroides, hippurus, miniatus, robinsoni, tahan, caniceps, erythraeus, pluto,
sladeni, similis, phayrei, blanfordi, pygerythrus, janetta, pryeri, philippinensis,
melanogaster, tenuis, brooki, lozvi, stevensi, blythi.
This list is probably incomplete.
The division of this genus is very difficult; but provisionally I think it is
reasonable to divide the genus into twelve groups.
It must, however, be borne in mind that this arrangement is very provisional,
and how many of the groups would stand in a detailed revision of the whole
genus I do not know.
1. tenuis group. Small Squirrels, hindfoot usually under 36 mm., and upper
incisors less proodont. Malay Peninsula, Sumatra, Borneo. The tail is
normally much narrowed.
tenuis, jentinki, probably fraterculus.
2. lowi group. Size as in group i, but as far as seen upper incisors strongly
proodont. Tail not much narrowed, but often relatively short. Malay
352 CALLOSCIURUS
Peninsula, Sumatra, Borneo. Both these groups have the bacuhim of the
" Tomeutes" tvpe so far as known.
All other forms of the typical subgenus are larger animals, with
hindfoot measurement usually more than 36 mm.
3. erythraeus group. This contains the majority of the Squirrels belonging to
the genus from the northern part of the range, i.e. South China to Siam,
with the baculum, so far as known, of "Callosciunis" type as diagnosed
by Thomas. Usually, but not always, the upper incisors do not tend to
be proodont. The colour is extremely various, but there appear to be
intermediate forms between all the extremes. As examples may be
quoted some races of ferrugineiis (unicolorous red); cockereUi, mostly
red above, white below; fiiilaysoni, typically unicolorous white; hocourti,
typically black above, white below; germaini subspecies, unicolorous
black. C. erythraeus is typically greenish above, red below, but some
races are whitish below. There are no flank stripes except in a race of
sladeni, which has black ones. The colour is never black above, red below,
as in prevosti. C. atrodorsalis has usually a black mid-dorsal region.
erythraeus, sladeni, ferrugineiis, finlaysoni, hocourti, germaini, flavimanus,
atrodorsalis, cockereUi, griseimanus.
4. caniceps group. Doubtfully distinct from erythraeus group, but upper
incisors very generally tending to be proodont. C. caniceps is typically
orange above, and with black tailtip, but as usual there is much variation
in colour in races (as arranged by Robinson & Kloss). Tenasserim and
Siam to Malacca.
5. prevosti group. Related to the above two groups so far as known in bacular
characters; black above, red below, always as far as seen, with or without
white flank stripe which may in some forms be broadened so that it
takes up most of the back; shades extremely variable; in some cases
these appear to be the most beautifully marked of all Squirrels. Malacca,
Sumatra, Borneo, Celebes. The upper incisors are very generally
proodont.
6. notatus group. Closely allied to the above; bacular characters so far as
known like the "Callosciunis" type of Thomas, except apparently
vittatus (but one? out of forty-six races examined; see remarks on
p. 349). Very generally with a white stripe over a black stripe on the
flanks (one race only in all forms examined with white flank-stripe only).
Body usually green above; red or in iiigrovittatus greyish below. Upper
incisors very generally proodont. notatus, vittatus, nigrovittatus. Malay
Peninsula and islands to Borneo. In the last four groups, the parietal
ridges of the skull, so far as seen, very rarely tend to come together, and
the zygomatic plate is rarely heavily ridged.
7. pvgervthrus group. Squirrels from Burma, Assam, and Nepal, with so far
as known the baculum of the " Tvmeutes" type of Thomas. The parietal
CALLOSCIURUS 353
ridges may come together, though this is not a usual feature of the skull.
There is a tendency for the zygomatic plate to be rather more powerfully
ridged, and it may extend upwards farther forwards than is usual. The
colour is typically duller than in the ervthraeiis group, though this varies;
no flank stripes except in phavrei, in w hich they are black. The incisors
most often do not tend to be projected forwards.
pygerxthrus, lokroides, blvtlii, stevoisi, phayrei.
8. qiiinquestriatus group. Very similar to group 7, but with the belly typically
longitudinally banded black and white, and with a well-developed black
mid-ventral stripe, so far as seen. This feature seems sufficiently rare
to warrant the formation of a group for this species. Yunnan, Burma.
9. hippitrus group. This contains several squirrels agreeing, so far as at present
known, with the pvgervthrus group in the structure of the baculum, from
the southern portion of the range of the genus (Malacca, Sumatra, Borneo,
Philippines). Most often the general colour is somewhat darker than
in r\\e pygerythrus group; frequently more or less unicolor, as in brookei,
philippitiensis, etc. melanogaster is a dark form with a black belly, and
differs from the other species, included here, so far as seen, in the more
proodont upper incisors. The colours are more strongly contrasted in
hippiinis, pryeri, and steerei. The tail in hippurus is extremely bushy;
this species tends to become largest of the genus except rubriventer.
A short sagittal crest is a normal feature of the skull, so far as seen, in
hippurus, melanogaster, and all species examined from the Philippine
Islands, but is not so in brookei.
hippurus, pryeri, brookei, philippinensis, steerei, juvencus, mindanensis,
samarensis ( ? other Philippine species), melanogaster.
10. leucomus group. I have not seen enough material to be able to frame a
definition of this group. About seven species are described as members
of the group (or near leucomus or rosenbergi), which are not represented
in London. The skull oi leucomus seems aberrant, as noted above, though
how far this is a constant feature I do not know. Conspicuous ear tufts
are present in both species seen which are allotted to the group (in this
respect differing from all other groups except the rubriventer group and
the subgenus Tamiops); but this is not a constant character in some other
described forms. C. leucomus has well-marked white spots behind the
ears, not seen in other members of the genus, but these are absent in
rosenbergi and others. Celebes, (leucomus, rosenbergi the sole species
examined.)
1 1 . rubriventer group. Very large ; ears tufted ; upper incisors strongly proodont ;
a long sagittal crest present in both skulls seen. Celebes.
12. maclellandi group (subgenus Tamiops). Usually smaller than the other
species of the genus Tail usually much narrowed. A Tamias-WVji
colour pattern of parallel black stripes on the back. Ear usually tufted.
Burma, South China, Siam, Hainan, Formosa, to Tibet, Chihli.
23 — Living Rodents — I
354 CALLOSCIURUS
This arrangement is, as remarked above, very provisional, and it may be
that many of these groups will break down when the whole genus is carefully
revised; for such a revision there is much need. It seems to me to be a more
natural arrangement at any rate than lumping the species into two "genera"
based on a single external character that cannot vet ha\e been verified in a third
of the named forms.
List of N.^med Forms
Subgenus Tamiops, Allen'
1. CALLOSCIURUS MACLELLANDI M.ACLELLANDI, Horsfield
1X39. Proc. Zool. Soc. London, p. 152.
Assam.
Synonym: pembcrtoni, Blyth, 1842, J. A. S. Bengal, XI, p. SS7. Bhutan.
Icucotis, Temminck, 1853, Esq. Zool. Cote de Guine, p. 252.
Malacca.
2. CALLOSCIL RUS M.^CLELLANDI MANIPURENSIS, Bonhote
igoo. .\nn. Mag. Nat. Hist. 7, V, p. 51.
Aimole, Manipur.
3. CALLOSCIURUS MACLELLANDI MARITIMUS, Bonhote
igoo. .^nn. Mag. Nat. Hist. 7, V, p. 51.
Foochow, China.
4. CALLOSCIURUS MACLELLANDI MONTICOLUS, Bonhote
1900. .Ann. Mag. Nat. Hist. 7, V, p. 52.
Ching Fen Ling, Fokien, China.
5. CALLOSCIURUS MACLELLANDI FORMOSANUS, Bonhote
1900. .\nn. Mag. Nat. Hist. 7, V, p. 52.
North Formosa.
6. CALLOSCIURUS MACLELLANDI BARBEl, Blyth
1S47. Juurn. .\siat. Soc. Bengal, XVI, p. S75.
Ye, Tenasserim.
7. CALLOSCIURUS MACLELLANDI KONGENSIS, Bonhote
1901. Proc. Zool. Soc. London, i, p. 55.
Raheng, Siam.
X. CALLOSCIURUS MACLELLANDI NOVEMLINE.ATUS, Miller
1903. Proc. Biol. Soc. Washington, XVI, p. 147.
Trang, Siamese Malaya.
.,. CALLOSCIURUS MACLELLANDI RODOLPHEI, Milne-Edwardb
1S67. Re\-. Mag. Zool. XIX, p. 227.
Cochin-China.
10. CALLOSCIURUS M.A.CLELLANDI HAINANUS, Allen
1906. Bull. .Amer. Mus. Nat. Hist. XXII, p. 476.
Lei-Mui Mon, Hainan (mountains).
11. CALL(.(SCIL!RUS M.ACLELLANDI SAUTERI, Allen
191 1. Bull. .Amer. Mus. Nat. Hist. XXX, p. 339.
Chip Chip, Northern Formosa.
12. CALLOSCIURUS MACLELLANDI RIUDONI, Allen
1906. Bull. .Amer. Mus. Nat. Hist. XXII, p. 477-
Riudon, Hainan (plains).
' For further notes on these forms, see p. 653.
CALLOSCIURUS 3SS
13. CALLOSCIURUS MACLELLANDI LIANTIS, Kloss
1919. Joiirn. Nat. Hist. Soc. Siam, III, no. 4, p. 370.
Cape Liant, S.-E. Siam.
14. CALLOSCIURUS M.ACLKLLANDI LAOTUM, Robinson & Kloss
1922. .Ann. Mag. Nat. Hist. 9, IX, p. 92.
Pak Hin Bung, Mekong Ri%'er, Laos.
15. CALLOSCIURUS MACLELLANDI MOI, Robinson & Kloss
1922. -Ann. Mag. Nat. Hist. 9, IX, p. 92.
Langbian Plateau, S. Annam.
16. CALLOSCIURUS MACLELLANDI RUSSEOLUS, Jacobi
1923. .\bh. Mus. Dresden, 16, no. i, p. 11.
Tibet.
17. CALLOSCIURUS MACLELLANDI FORREST!, Thomas
1920. .Ann. Mag. Nat. Hist. 9, V, p. 305.
Lichiang Range, Yunnan.
18. CALLOSCIURUS MACLELLANDI OLIVACEUS, Osgood
1932. Field Mus. Nat. Hist. Publ. Zool. Ser. XVIII, p. 292.
Mt. Fan Si Pan, near Chapa, Tongking.
19. CALLOSCIURUS SWINHOEI, Milne-Edwards
1874. Rech. des Mamm. p. 308.
Moupin, Tibet.
(Listed as a distinct species by Robinson & Kloss, 191 8.)
20. CALLOSCIURUS INCONSTANS, Thomas
1920. .Ann. Mag. Nat. Hist. 9, V, p. 306.
Mongtze (?), Yunnan.
21. CALLOSCIURUS CL.ARKEI. Thomas
1920. Ann. Mag. Nat. Hist. 9, V, p. 304.
Yantze Valley, Yunnan (North).
22. CALLOSCIURUS SPENCEI, Thomas
1921. Joum. Bombay Nat. Hist. Soc. XXVII, p. 503.
North Kachin Province, N. Burma.
23. CALLOSCIURUS HOLTI, New Name
(To replace Taniiops (^^Callosciurus) lylei, Thomas)
1920. .Ann. Mag. Nat. Hist. 9, V, p. 307.
Coast 50 miles south of Bangkok, S.-E. Siam. Not Callosciurus lylei,
Bonhote.
24. CALLOSCIURUS VESTITUS, Miller
1913. Proc. Biol. Soc. Washington, XXVIII, p. 115.
Hsin-Lung-Shan, 65 miles north-east of Peking, China.
Subgenus Callosciurus, Gray
tenuis Group
25. CALLOSCIURUS TENUIS TENUIS, Horsfield
1824. Zool. Res. Java, p. 153.
Singapore Island.
SynoniTn: affinis, Horsfield, 1824, Zool. Res. Java, p. 153. Singapore.
35f> CALLOSCIURUS
2f.^ CALLOSCURLS TENUIS SURDUS, Miller
iqoo. Proc. Acad. Sci. Washington, XI, p. So.
Tranc, Siamese Malaya.
27. CALLOSCILRUS TENUIS SORDIDUS, Kloss
iQii. Ann. Mag. Nat. Hist. 8. VII, p. 119.
Great Redang Island, off Trengganu, East Malay Peninsula.
2S. CALLOSCIURUS TENUIS TIOMANICUS, Robinson
1917. Journ. Fed. Malay States Mus. VII, p. 103.
Tioman Island, east coast of Malay Peninsula.
2Q. CALLOSCIURUS TENUIS TAHAN, Bonhote
igo8. Journ. Fed. Malay States Mus. Ill, p. 6.
Mt. Tahan, Pahang, Malay Peninsula.
30. CALLOSCIURUS TENLTS GUNONG. Robinson &: Kloss
1914. Journ. Fed. Malay States Mus. V, p. 119.
Kao Xong, Bandon, Siamese Malaya.
,u. CALLOSCIL^RUS TENUIS MODESTUS, Muller
1839. Temmincks Verhandelinger Zoologie, Inleidung, p. 55.
Mt. Singgalang, Sumatra.
32. CALLOSCIURUS TENUIS ALTITUDINIS, Robinson & Kloss
igi6. Journ. Straits Branch Roy. Asiat. Soc. 73, p. 269.
Korinchi Peak, Sumatra, 7,300 ft.
33. CALLOSCIURUS TENUIS MANSAL.ARIS, Miller
1903. Proc. L*. S. Nat. Mus. XXVI. p. 451.
Mansalar Island, W. Sumatra.
34. CALLOSCIURUS TENUIS B.\TUS, Lyon
1916. Proc. U. S. Xat. Mus. LII, p. 443.
Tana Bala, Batu Islands, W. Sumatra.
35. CALLOSCIURUS TENUIS BANCARUS, Miller
1903. Proc. U. S. Xat. Mus. XXVI, p. 451.
Bangkaru Island, Banjak Islands, W. Sumatra.
36. CALLOSCILRIS TENUIS PUMILUS, Miller
1903. Smiths. Misc. Coll. XLV, p. 15.
South Pagi Island, \V. Sumatra.
37. CALLOSCIURUS TENUIS PARVUS, Miller
1901. Proc. Biol. Soc. Washington, XIV, p. 33.
Sarawak, Borneo.
38. CALLOSCIURUS TENUIS SIANTANTCUS, Chasen & Kloss
1928. Journ. Malay Branch Roy. Asiat. Soc. 6, p. 33.
Anamba Islands, South China Sea.
y). CALLOSCIURUS PROCERUS, Miller
1901. Proc. Washington Acad. Sci., X, p. 122.
Bunguran Island, North Natunas.
40. CALLOSCIURUS JENTINKI, Thomas
18S7. Ann. Mag. Nat. Hist. 5, XX, p. 128.
Kina Balu, North Borneo.
CALLOSCIURUS 35-
41. CALLOSCIURUS FRATERCULUS, Thomas
1895. Ann. Mus. Civ. Stor. Nat. Genova, 2, XIV, p. 10.
Sipora, Mentawti Islands, West .Sumatra.
loui Group
42. CALLOSCIURUS LOWI LOWI, Thomas
1892. .\nn. Mag. Nat. Hist. 6, IX, p. 253.
Sarawak, Borneo.
43. CALLOSCILRL'S LOWI BANGUEYAE, Thomas
1910. Ann. Mag. Nat. Hist. 8, V, p. 387.
Banguey Island, North Borneo.
44. CALLOSCIURUS LOWI N.^TUNENSIS, Thomas
1895. Nov. Zool. II, p. 26.
Sirhassen Island, Natuna group, South China Sea.
45. CALLOSCIURUS LOWI ROBINSONI, Bonhote
1903. Fasciculi Malayenses, Zool. I, p. 24, pi. i.
Bukit Besar, Patani, Malay Peninsula.
46. CALLOSCIURUS LOWI HUMILIS, Miller
1913. Smiths. Misc. Coll. LXI, no. 21, p. 24.
Kateman River, East .Sumatra.
47. CALLOSCIURUS LOWI VANAKENI, Robinson & KJoss
J916. Joum. Straits Branch Roy. .Asiat. Soc. 73, p. 270.
Barisan Range, Korinchi, Sumatra.
48. CALLOSCIURUS LOWI PINIENSIS, MUler
1903. Smiths. Misc. Coll. XLV, p. 14.
Pinie Island, Batu group. West Sumatra.
49. CALLOSCIURUS LOWI BAL.\E, Miller
1903. Smiths. Misc. Coll. XLV, p. 14.
Tana Bala Island, Batu group, W. Sumatra.
50. CALLOSCIURUS LOWI SEIMUNDI, Thomas & Wroughton
1909. Ann. Mag. Nat. Hist. 8, III, p. 440.
Kundur Island, Rhio-Lingga Archipelago, East Sumatra.
51. CALLOSCIURUS LOWI .\LACRIS, Thomas
1908. Ann. Mag. Nat. Hist. 8, II, p. 306.
Semangko Pass, Selangor-Pahang boundar\', Malaya.
52. CALLOSCIURUS LOWI SIBERU. Chasen & Kloss
1928. Proc. Zool. Soc. London {1927), p. 824.
Siberut Island, W. Sumatra.
53. CALLOSCIURUS LINGUNGENSIS, Miller
1901. Proc. Washington Acad. Sci., Ill, p. 123.
Pulo Lingung, North Natuna Islands.
erythraeus Group
54. CALLOSCIURUS ERYTHRAEUS ERYTHRAEUS, Pallas
1778. Nov. Sp. Quadr. Glir. Ord. p. 377.
Locality not known.
358 CALLOSCIURUS
55. CALLOSCIIRLS ERVTHRAKLS BHUTANENSIS, Bonhotc
iQOi. Ann. Map. Nat. Hist. 7, VII, p. 161.
Bhutan.
56. CALLOSCIURUS ERYTHRAKUS NAGARUM, Thomas & Wroughton
191(1. Journ. Bombay Nat. Hist. Soc. XXIV, p. 228.
Sadiya, Assam.
57. CALLOSCIURUS ERVTHRAEUS ERYTHROGASTER, Blyth
1842. Journ. As. Soc. Bengal, XI, p. 970.
Manipur.
58. CALLOSCIURUS ERYTHRAEUS PUNCTATISSIMUS, Gray
1867. Ann. Mag. Nat. Hist. 3, XX, p. 283.
Cachar, Assam.
50. CALLOSCIURUS ERYTHRAEUS INTERMEDIUS, Anderson
1879. Zool. & Anat. Res. Yunnan, p. 241.
Assam.
60. CALLOSCIURUS ERYTHRAEUS GORDON!, Anderson
1S71. Proc. Zool. Soc. London, p. 140.
Bhamo, Upper Burma.
61. CALLCJSCIURUS ERYTHRAEUS KINNEARI, Thomas & WrouRhton
1916. Journ. Bombay Nat. Hist. Soc. XXIV, p. 229.
Tatkon, Kindat, Upper Chindwin, Burma.
b2. CALLOSCIURUS ERYTHRAEUS HYP1:RYTHKUS, Blyth
1855. Journ. As. Soc. Bengal, XXIV, p. 474.
Tenasserim. (? Moulmein.)
63. CALLOSCIURUS ERYTHRAEUS RUBECULUS, Miller
1903. Smiths. Misc. Coll. XLV, p. 22.
Khow Sai Dow, Trong, Siamese Malaya.
64. CALLOSCIURUS ERY'THR.\EUS YOINGI, Robinson & Kloss
1914. .Ann. Mag. Nat. Hist. 8, XIII, p. 224.
Gunong Tahan, 5,000-6,000 ft. N. Pahang, Malaya.
h5. CALLOSCIURUS ERYTHRAEUS CASTANEOVENTRIS, Gray
1842. Ann. Mag. Nat. Hist., X, p. 263.
Hainan.
66. CALLOSCIURUS l-RYTHRAEUS AQITLO, Wroughton
1921. Journ. Bombay Nat. Hist. Soc. XXVII, p. 601.
Dibong River, Sadiya, Assam.
67. CALLOSCIURUS ERYTHRAEUS GLOVERI. Thomas
1921. Journ. Bombay Nat. Hist. Soc. XXVII, p. 502.
Nagohuka, W. Szechuan, China.
68. CALLOSCIURUS ERYTHRAEUS NINGPOENSIS, Bonhote
1901. Ann. Mag. Nat. Hist. 7, VII. p. 163.
Ningpo, China.
Synonym: tsingtaueiisis. Hilzheimer, 1905, Zool. .Anz. XXIX, p.
Tsingtau, China.
6q. CALLOSCIURUS ERYTHRAEUS GRISEOPECTUS, Blyth
1S47. Journ. As. Soc. Bengal, XVI, p. S73.
Locality not known.
CALLOSCIURUS 359
70. CALLOSCIURUS EKYTHRAEUS STYANI, Thomas
1894. Ann. Mag. Nat. Hist. 6, XIII, p. 363.
Betvveen Shanghai and Hangchow, probably Kahing, China.
71. CALLOSCIURUS KRYTHR.XKUS BONHOTEI, Robinson & Wroughton
1911. Journ. Fed. Malay States Mus. IV, p. 234.
Chin Chien San, .Szechuan, China.
72. CALLOSCIURUS KRYTHRAKUS MICHL^^NUS, Robinson & Wroughton
191 1. Journ. Fed. Malay States Mus., IV, p. 234.
Mee-chee, Yunnan.
73. CALLOSCIURUS EKYTHRAEUS HAEMOBAPHES, G. M. Allen
1912. Proc. Biol. Soc. Washington, XXV, p. 177.
Chih-Ping, S.-E. Yunnan.
74. CALLOSCIURUS ERYTHRAEUS TH-AIWANENSIS, Bonhote
1901. Ann. Mag. Nat. Hist. 7, VII, p. 165.
Baksa, S. Formosa.
75. CALLOSCIURUS ERYTHRAEUS CENTR.A.LIS, Bonhote
1901. Ann. Mag. Nat. Hist. 7, VII, p. 166.
Lak-Ku-Li, Central Formosa.
76. CALLOSCIURUS ERYTHRAEUS ROBERTI, Bonhote
igoi. Ann. Mag. Nat. Hist. 7, VII, p. 166.
N.-W. Formosa.
77- CALLOSCIURUS ERYTHRAEUS CRUMPI. Wroughton
1916. Journ. Bombay Nat. Hist. Soc. XXIV, p. 425.
Sedonchen, Sikkim.
78. CALLOSCIURUS ERY'THRAEUS INSULARIS, Allen
1906. Bull. Amer. Mus. Nat. Hist., XXII, p. 473.
Lei-Mui-Mon, Hainan.
79. CALLOSCIURUS ERYTHR.AEUS HENDEEI, Osgood
1932. Field. Mus. Nat. Hist. Publ. Zool. Ser. XVIII, p. 270.
Chapa, Tongking.
80. CALLOSCIURUS ERYTHRAEUS CROT.\LIUS, Thomas & Wroughton
1916. Journ. Bombay Nat. Hist. Soc. XXIV, 2, p. 229.
Hkamti, Chindwin, Burma.
81. CALLOSCIURUS ERYTHR.\EUS WELLSI, Wroughton
1921. Journ. Bombay Nat. Hist. Soc. XXVII, p. 775.
Shangpung, Jaintia Hills, Assam.
82. CALLOSCIURUS ERYTHRAEUS NIGRIDORSALIS, Kuroda
'935- Journ. Manim. Baltimore, p. 281.
Riran, Taito, S.-E. Formosa.
83. CALLOSCIURUS ERYTHRAEUS WOODI. Harris
193 1. Occ. Pap. Mus. Zool. Univ. Mich. 228, p. i.
Lung-Tan, 25 miles east of Nangking, Kiang-Su, China.
84. CALLOSCIURUS FLAVIMANUS FLAVIALANUS. Geoffrey
1832. Mag. Zool. I, Mamm. CI. i, Ann. 2.
Tourane, Annam.
3bo CALLOSCIURUS
55. CALLOSCIL'RUS FLAVIMANUS QUANTULUS, Thomas
1927. Proc. Zool. Soc. London, p. 51.
Xieng Khouang, Laos, Annam.
56. CALLOSCIURUS FLAVIMANUS DACTYLINUS, Thomas
1Q27. Proc. Zool. Soc. London, p. 52.
Dak-to, Annam.
87. CALLOSCIURUS FLAVIMANUS CONTUMAX, Thomas
IQ27. Proc. Zool. Soc. London, p. 52.
Kontoum, south of Dak-to, Annam.
88. CALLOSCIURUS FLAVIMANUS PIRATA, Thomas
1929. Proc. Zool. Soc. London (1928), p. 836.
Napi, Laos, Annam.
8q. CALLOSCIURUS FLAVIMANUS BOLOVENSIS, Osgood
1932. Field. Mus. Nat. Hist. Pub. Zool. Ser. XVIH, p. 276.
Paksong, Boloven Plateau, Laos, Annam.
90. CALLOSCIURUS SLADENI SLADENI, Anderson
1S71. Proc. Zool. Soc. London, p. 139.
Thigyain, L^pper Burma.
Synonym; kc?ii»iisi, Wroughton, 1908, Ann. Mag. Nat. Hist. 8, XI,
p. 491. Katha, LIpper Irrawaddy.
91. CALLOSCIURUS SLADENI MIDAS, Thomas
1914. Journ. Bombay Nat. Hist. Soc. XXIH, p. igS.
Myitkyina, Upper Burma.
.,::. CALLOSCIURUS SLADENI RUIiEX, Thomas
1914. Journ. Bombay Nat. Hist. Soc. XXIH, p. 198.
Lonkin, Myitkyina district. Upper Burma.
93. CALLOSCIURUS SLADENI BARTONI, Thomas
1914. .Ilium. Bombay Nat. Hist. Soc. XXHL p. 199-
Uyu River, 50 miles east of Homalin, Upper Chindwin, Burma.
.,4. CALLOSCIURUS SLADENI SHORTRIDGEI, Thomas S: Wroughton
1916. Journ. Bombay Nat. Hist. Soc. XXIV, p. 232, pi. fig. i.
Hkamti, Upper Chindwin, Burma.
05. CALLOSCIURUS SLADENI FRYANUS, Thomas & Wroughton
1916. Journ. Bombay Nat. Hist. Soc. XXIV, p. 232, pi. fig. 2.
Minsin. LTpper Chindwin, Burma.
96. CALLOSCIURLS SLADENI CAREYI, Thomas & Wroughton
1916. JiHirn. Bombay Nat. Hist. Soc. XXIV, p. 233, pi. fig. 3.
Tamanthe, Upper Chindwin, Burma.
97. CALLOSCIURUS SLADENI HARINGTONI, Thomas
1905. Ann. Mag. Nat. Hist. 7, XVI, p. 314.
Moungkan, Upper Chindwin, Burma.
98. CALLOSCIURUS SLADENI MILLARDI, Th.mias & Wroughton
1916. Journ. Bombay Nat. Hist. Soc. XXIV, p. 234, pi. fig. 5.
Pyaungbyin, 40 miles north of Kindat, Upper Chmdwin, Buniia.
00. CALLOSCIURUS SLADENI SOLUTUS, Thomas
1914. Journ. Bombay Nat. Hist. Soc. XXIII, p. 199.
Homalin, Upper Chindwin, Burma.
CALLOSCIUKUS 361
100. CALLOSCIUKUS FERRUGINEUS FERRUGINEUS, F. Cuvicr'
1829. Hist. Nat. Mamm. iii, pi. 238.
Peyu, Lower Uurma.
loi. CALLOSCIUKUS FERRUGINEUS FRANDSENI, Kloss
1916. Proc. Zool. Soc. London, p. 46.
Koh Chang Island, S.-E. Siam.
102. CALLOSCIUKUS FKKRUGINICUS CINNAMOMEUS, Temminck
1853. Esq. Zool. Cote de Guinc, p. 250.
Cambodia.
103. CALLO.SCIURUS FKRRUGINEUS \VlLLI.\MSONI, Robinson & Kloss
1922. Ann. Mag. Nat. Hist. 9, IX, p. 90.
Xieng Khan, Mekong River, Siam.
104. CALLOSCIURUS FERRL'GI.NKUS HKRliERTI, Robinson & Kloss
1922. Ann. Mag. Nat. Hist. 9, IX, p. 90.
Hup Bon, near Sriracha, S.-E. Siam.
105. CALLOSCIURUS FERRUGINEUS PIERKEI, Robinson & Kloss
1922. Ann. Mag. Nat. Hist. 9, IX, p. 91.
Phu Quoc Island, Cambodia.
106. CALLOSCIURUS FERRUGINEUS PHANRANGIS, Robinson & Kloss
1922. Ann. Mag. Nat. Hist. 9, IX, p. 91.
Tour Cham, near Phanrang, S. Annam.
107. CALLOSCIURUS FERRUGINEUS MEN.'\MICUS, Thomas
1929. Proc. Zool. Soc. London (1928), p. 839.
Nan, N. Siam.
108. CALLOSCIURUS FERRUGINEUS ANNELLATUS, Thomas
1929. Proc. Zool. Soc. London (1928), p. 839.
Angkor, Cambodia.
loy. CALLOSCIURUS COCKERELLI, Thomas
1928. Ann. Mag. Nat. Hist. 10, II, p. 100.
Nan, N. Siam.
110. CALLOSCIURUS FINL.A.YSONI FINL.^YSONI, Horsfield
1824. Zool. Res. Java, p. 151.
Koh Si Chang Islands, Bight of Bangkok, Siam.
Synonym: keraudreni. Lesson, 1830, Cent. Zool. pi. i. Burma.
siamensis. Gray, i860, Ann. Mag. Nat. Hist., 3, \, p. 500.
Siam.
partus. Kloss, 1915, Journ. Nat. Hist. Soc. Siam, I, p. 158.
Koh Si Chang Islands.
111. CALLOSCIURUS FINL.-W.SONI FOLLETTI, Kloss
1915. Journ. Nat. Hist. Soc. Siam, I, p. 159.
Koh Phai, Inner Gulf of Siam.
112. CALLOSCIURUS FINLAV.SONl TACHARUl, Robinson
1916. Journ. Fed. Malay States Mus. VII, p. 36.
Krabin, Central Siam.
113. CALLOSCIURUS FINL.'WSONI TROTTERI, Kloss
1916. Journ. Nat. Hist. Soc. Siam, II, p. 178.
Koh Lan Island, Inner Gulf of Siam.
* looa. Callosciurus fcrru^incus splcndens. Gray; omitted in error; for reference see p. 653.
362 CALLOSCIURUS
114. CALLOSCRRUS BOCOURTII BOCOURTII, Milne-Kduards
1S67. Rev. Zool. p. 193.
Awtha, Siam.
Synonym: leucocephaliis, Honhote, Proc. Zool. Soc. London, 1901, p. 54.
115. CALLOSCR'RL'S BOCOL'RTll IIARMANDl, MiliK-Edwards
1S76. Bull. Soc. Philom. 6, XII, p. 8.
Island Phil Quoc, off Cliantabun, Siam.
116. CALLOSLILRL'S BOCOLRTll SIXISTRALIS, Wroughton
igolS. Ann. Mag. Nat. Hist. 8, II, p. 399.
Pichit, Menam River, Central Siam.
117. CALLOSCIURUS BOCOURTII DEXTRALIS, Wroughton
190S. .\nn. Mag. Nat. Hist. 8, II, p. 400.
Kampeng, Lower Me-Ping Valley, Siam.
118. CALLOSCIURUS BOCOURTI GRUTLI, G\ldenstolpe
1917. Kungl. Svenska Vet. Akad. Handl. LVII, no. 2, p. 37.
Bang Hue Pong, N. Siam.
iiy. CALLOSCIURUS BOCOURTI LYLi:i, Wroughton
190S. Ann. Mag. Nat. Hist. 8, II, p. 401.
Chiengmai, N. Siam.
izo CALLOSCIURL'S BOCOURTI FLOWERI, Bonhote
1901. Ann. Mag. Nat. Hist. 7, VII, p. 455.
Klong Morn, near Bangkok, Siam.
121. CALLOSCIURUS GKRMAINI GERMAINI, Alilne-lidwards
1S67. Rev. Zool. p. 193.
Pulau Condor, off Cambodian coast.
122. CALLOSCIURUS GERMAINI ALBIVEXILLl, Kloss
igi6. Proc. Zool. Soc. London, p. 47.
Koh Kut Island, S.-E. Siam.
123. CALLOSCIURUS GERMAINI NOX, Wroughton
1908. Ann. Mag. Nat. Hist. 8, II, p. 397-
Sea coast south-east of Bangkok, Siam.
124. CALLOSCIURUS .\TRODORSALIS ,\TRODORSALlS, Gray
1842. .Ann. Mag. Nat. Hist. X, p. 263.
Bhutan (error), substitute Moulmein.
125. CALLOSCIURUS ATRODORSALIS THAI, Kloss
1917. Journ. Nat. Hist. Soc. Siam, II, p. 285.
Rahcng, C. Siam.
126. CALLOSCILRUS .ATRODORSALIS SHANICUS, Ryley
1914. Journ. Bombay Nat. Hist. Soc. XXII, p. 663.
Gokteik, N. Shan States, Burma.
127. CALLOSCIURUS ATRODORSALIS ZIMMEENSIS, Robinson S: Wroughton
1916. Journ. Fed. Malay States Mus. VII, p. 91.
Chiengmai, N. Siam.
12S. CALLOSCIURUS ATRODORSALIS T.ACHIN, Rloss
1916. Journ. Nat. Hist. Soc. Siam, II, p. 17S.
Tachm, C. Siam.
CALLOSCIURUS 363
129. CALLOSCILRUS ATRODORSALIS I'RANIS, Kloss
1916. Journ. Nat. Hist. Soc. Siam, II, p. 178.
Koh Lak, Pran, S.-W. Siam.
130. CALLOSCIL'KUS GRISEIMANUS GRISEIMANUS, Milne-Edwards
1867. Rev. Zool. p. 195.
Cambodia.
131. CALLOSCR'RUS GRISEIMANUS LEUCOPUS, Gray
1867. Ann. Mag. Nat. Hist. 3, XX, p. 282.
Cochin China.
132. CALLOSCIURUS GRISEIMANUS VASSALI, Bonhote
1907. Proc. Zool. Soc. London, p. y (footnote).
Ninh Hoa, Annam.
Synonym: Callosaurus griseimanus fumigatus, Bonhote, 1907, Abstr.
Proc. Zool. Soc. London, Jan. 15th, p. 2. Ninh Hoa,
.Annam. Preoccupied hy fumigatus, Gray, 1867.
caniceps Group
133. CALLOSCIURUS IMITATOR, Thomas
1925. Proc. Zool. Soc. London, 2, p. 502.
Thai-Nien, Tongking.
134. CALLOSCIURUS CANICEPS CANICEPS, Gray
1842. Ann. Mag. Nat. Hist. X, p. 263.
Bhutan (error), substitute N. Tenasserim.
Synonym: chrysonotus, Blyth, 1847, Journ. Asiat. Soc. Bengal, XVI,
p. 873. Amherst, Tenasserim.
epomophorus fluminalis, Wroughton & Robinson, igii,
Journ. Fed. Malay States Mus. IV, p. 233. Meping
Rapids, N. Siam.
135. CALLOSCIURUS CANICEPS D.AVISONI, Bonhote
1901. Ann. Mag. Nat. Hist. 7, VII, p. 273.
Bankachon, S. Tenasserim.
136. CALLOSCIURUS CANICEPS INEXPECTATUS, Kloss
1916. Journ. Nat. Hist. Soc. Siam, II, p. 178.
Koh Lak, Pran, S.-\V. .Siam.
Synonym: {}) helgi'i, Gyldenstolpe, 1017, Kungl. Svenska. Vet. Ak.
Handl. LVII, 2, p. 34. South of Koh Lak, S.-W. Siam.
137. CALLOSCIURUS CANICEPS SULLIVANUS, M.ller
1903. Smiths. Misc. Coll. XLV, p. 17.
Sullivan Island, Mergui Archipelago.
138. C.M.LOSCIURUS CANICEPS DO.MELICUS, Miller '
1903. Smiths. Misc. Coll. XLV, p. 18.
Domel Island, Mergui Archipelago.
139. CALLOSCIURUS CANICEPS BENTINC.^NUS, Miller
1903. Smiths. Misc. Coll. XLV, p. ig.
Bentinck Island, ^IerRui .Archipelago.
140. CALLOSCIURUS CANICEPS .M.Vn'HAEUS, Miller
1903. Smiths. Misc. Coll. XLV, p. 19.
St. Matthew Island, Mergui .■\rchipeIago.
364 CALLOSCIURUS
141. CALLOSCIURUS CANICEPS LUCAS, Miller
1903. Smiths. Misc. Coll. XLV, p. 20.
St. Luke Island, Mergui Archipelago.
142. CALLOSCIL'RUS CANICEPS CASENSIS, Miller
1003. Smiths. Misc. Coll. XLV, p. 20.
Chance Island, Mergui Archipelago.
143. CALLOSCIURUS CANICEPS ALTINSULARIS, Miller
1903. Smiths. Misc. Coll. XLV, p. 21.
High Island, Mergui Archipelago.
144. CALLOSCIURUS CANICEPS EPOMOPHORUS, Bonliote
1 00 1. Ann. Mag. Nat. Hist. 7, VII, p. 272.
Salanga or Junk Ceylon Island, Siamese Malaya.
145. CALLOSCIURUS CANICEPS PANJIUS, Thomas & Robinson
1921. Ann. Mag. Nat. Hist, g, VII, p. 119.
Telok Pah, east of Salanga, Pulau Panjang. Peninsular Siam.
146. CALLOSCIURUS CANICEPS PANJIOLI, Thomas & Rob.nson
1921. Ann. Mag. Nat. Hist. 9, VII, p. 120.
Pulau Panjang Anak, north of P. Panjang, Peninsular .Siam.
147. CALLOSCIURUS CANICEPS NAKANUS, Thomas & Robinson
1921. Ann. Mag. Nat. Hist. 9, VII, p. 120.
Koh Naka, near Salanga, Peninsular Siam.
14S. CALH)SCIURUS CANICEPS MAPRAVIS, Thomas S: Robinson
1921. Ann. Mag. Nat. Hist. 9, VII, p. 120.
Koh Maprau, near Salanga, Peninsular Siam.
140. CALLOSCIURUS CANICEPS PIPIDONIS, Thomas & Robinson
1921. Ann. Mag. Nat. Hist. 9, VII, p. 121.
Koh Pipidon, near Salanga, Peninsular Siam.
150. CALLOSCRIRUS CANICEPS TACOPIUS, Thomas & Robinson
1921. Ann. Mag. Nat. Hist, g, VII, p. 121.
Koh Rah, Tacopah, Peninsular Siam.
151. CALLOSCIL'RUS CANICEPS TABAUDIUS, Thomas
1922. Journ. Bombay Nat. Hist. Soc. XXVIII, 4, p. 1067.
, Tavoy Island, Mergui Archipelago.
152. CALLOSCIURUS CANICEPS HASTILIS, Thomas
1923. Journ. Hombay Nat. Hist. Soc. XXIX, 2, p. 377.
Hastings Island, Mergui Archipelago.
153. C.ALLOSCURUS C.\NICEPS MILLERI, Robinson & Wroughton
tgii. Journ. Fed. Malay States Mus. IV, p. 233.
Trong, Siamese Malaya.
154. CALLOSCIURUS CANICEPS SAMUIENSIS, Robinson &: Kloss
1914. .Ann. Mag. Nat. Hist. 8, XIII, p. 226.
Koh Samui, Bandon, Siamese Malaya.
155. CALLOSCIL'RUS CANICEPS F.ALL.4X. Robinson & Kloss
1914. Ann. Mag. Nat. Hist. 8, XIII, P- 225.
Koh Pennan, Bandon. Siamese Malaya.
156. CALLOSCIURUS CANICEPS LANC.AVENSIS, Milki
1003. Smiths. Misc. Coll. XLV, p. 16.
Pulau Langkawi, Straits of Malacca.
CALLOSCIURUS 365
157. CALLOSCIURUS CANICEPS ADANGF.NSIS, Miller
1903. Smiths. Misc. Coll. XLV, p. 17.
Pulau .\dang, Butang Archipelago, Straits of Malacca.
158. CALLOSCIIRUS CANICKPS TERUTAVENSIS, Thomas & Wroughton
1909. .\nn. Mag. Nat. Hist. 8, IV, p. 535.
Pulau Terutau, Straits of Malacca.
159. CALLOSCIURUS CANICEPS HELVUS, Shamel
1930. Journ. Mamtn. Baltimore, p. 72.
Koh Tau, east coast of Malay Peninsula.
160. CALLOSCIURUS CANICEPS CANIGENUS, Howell
1927. Joum. Washington, Acad. Sci., XVII, p. 8i.
Hayenhsien, Hangchow Bay, Chekiang, China.
161. CALLOSCIURUS CANICEPS CONCOLOR, Blyth
1855. Joum. As. Soc. Bengal, XXIV, p. 474.
Malacca.
Synonym: erubescens, Cabrera, 1917, Bol. Real. Soc. Espan. 17, p. 518.
Selangor.
162. CALLOSCIURUS CANICEPS TELIBIUS, Thomas & Robinson
1921. Ann, Mag. Nat. Hist. 9, VII, p. I2i.
Pulau Telibun, coast of Trang, Peninsular Siam.
163. CALLOSCIURUS MOHEIUS MOHEIUS, Thomas & Robinson
1921. Ann. Mag. Nat. Hist. 9, VII, p. 122.
Pulau Mohea (north), near Trang, Malaya.
164. CALLOSCIURUS MOHEIUS MOHILLIUS, Thomas & Robinson
1921. -Ann. Mag. Nat. Hist. 9, VII, p. 122.
Pulau Mohea (south), near Trang, Malaya.
prevosti Group
165. CALLOSCIURUS PREVOSTI PREVOSTI, Desmarcst
1822. Mamm., p. 335.
Malacca.
Synonym: rufogularis. Gray, 1842, -Ann. Mag. Nat. Hist. X, p. 263.
prevosli prevosti "subsubspecies" meticiilosus, Robinson,
1916, Joum. Fed. Malay States Mus. VII, p. 20. Triang,
S.-\V. Pahang, Malaya.
166. C.\LLOSCIURUS PREVOSTI RAFFLESI, Vigors & Horsfield
1828. Zool. Joum. IV, p. 113, pl- 'V.
Sumatra; probably Bencoolen.
167. CALLOSCIURUS PRF:V0STI HUMEI, Bonhote
1901. .\nn. Mag. Nat. Hist. 7, VII, p. 170.
Klang, Selangor, Malay Peninsula.
168. CALLOSCIURUS PREVOSTI \VR.-\YI, Kloss
1910. Journ. Fed. Malay States Mus. IV, p. 148.
Kuala Lipis, Pahang, Malaya.
160. CALLOSCIURUS PREVOSTI MELANOPS, Miller
1902. Proc. Acad. Nat. Sci. Philadelphia, p. 151.
Indragiri River, S.-E. Sumatra.
366 CALLOSCIURUS
170. CALLOSCIURUS PRHVOSTI PENIALIUS, Lyon
1905. Proc. U. S. Nat. Mus. XXXIV, p. 637.
Pulau Penjali, E. Sumatra.
171. CALLOSCIURUS PREVOSTI HARRISON!, Stone & Rchn
1002. Prnc. Acad. Nat. Sci. Philadelphia, LIV, p. 132.
Gunong Sugi, Lampongs, S.-E. Sumatra.
172. CALLOSCIURUS PRHVOSTI C0NDURP:NSIS, Miller
1006. Proc. U. S. Nat. Mus. XXXI, p. 260.
Pulau Kundur, Rhio-Lingga Archipelago.
173. CALLOSCIURUS PREVOSTI CARIMONENSIS, Miller
igo6. Proc. U.S. Nat. Mus. XXXI, p. 261.
Great Karimon Island, Rhio-Lingga Archipelago.
174. CALLOSCIURUS PREVOSTI BANGKANUS, Schlegel
1S63. Ned. Tijds. Dierk, i, p. 26, pi. i, fig. 2.
Bangka Island, E. Sumatra.
175. CALLOSCIURUS PREVOSTI MENDANAUUS, Lyon
1906. Proc. U.S. Nat. Mus. XXXI, p. 5S9.
Pulau Mendanau, west of Billiton.
176. CALLOSCIURUS PREVOSTI CARIM.ATAE, Miller
1906. Proc. U.S. Nat. Mus. XXXI, p. 57.
Karimata Island, ofl" Bomean coast.
177. CALLOSCIURI'S PREVOSTI SANGGAUS, Lyon
1907. Proc. U.S. Nat. Mus. XXXIII, p. 554.
Sanggau, W. Borneo.
17S. CALLOSCIURUS PREVOSTI ARMALIS, Lyon
loii. Proc. U.S. Nat. Mus. XL, p. 82.
Pulau Panebangan, west coast of Borneo.
179. CALLOSCIURL'S PREVOSTI PELAPIS, Lyon
191 1. Proc. U.S. Nat. Mus. XL, p. 82.
Pulau Pelapis, west coast of Borneo.
iSo. CALLOSCIL'RUS PREVOSTI BORNEOENSIS, Miiller & Schlegel
1S39-44. Verhandl. p. 86.
Pontianak, Borneo.
181. CALLOSCIURUS PREVOSTI PALUSTRIS, Lyon
1907. Proc. U.S. Nat. Mus. XXXIII, p. 553.
North bank of Kapuas River, W. Borneo.
1S2. CALLOSCIURUS PREVOSTI PROSERPINAE, Lyon
1907. Smiths. Misc. Coll. XLVIII, p. 275.
Pulau Temaju, W. Borneo.
183. CALLOSCIURUS PREVOSTI SARAWAKENSIS, Gray
1867. Ann. Mag. Nat. Hist. 3, XX, p. 277.
,Sarawak, Borneo.
184. CALLOSCIL"Rl'S PREVOSTI Kl'CHINGENSIS, Bonhote
1001. Ann. Mag. Nat. Hist. 7, VII, p. 170.
Kuching, Sarawak, Borneo.
185. CALLOSCIURUS PREVOSTI ATRICAPILLUS, Schlegel
1S63. Ned. Tijd. Dierk, i, p. 27, pi. ii, fig. i.
Kapuas River, W. Borneo.
CALLOSCIURUS 367
186. CALLOSCIURUS PREVOSTI ATROX, Mille
1913. Smiths. Misc. Coll. LXI, 21, p. 23.
Talisaian Mountain, Dutch S.-E. Uorneo.
187. CALLOSCIURUS PKHVOSTl CAROLl, Bonhote
1901. Ann. Mag. Nat. Hist. 7, VII, p. 173.
Baram, Borneo. (Low country.)
188. CALLOSCR'RUS PREVOSTI GRISEICAUDA, Bonhote
1901. Ann. Mag. Nat. Hist. 7, VII, p. 174.
Mount Kalulong, IJarani, Borneo.
189. CALLOSCIURUS PREVOSTI ERYTHROMELAS, Temminck
1853. Esq. Zool. Cote de Guine, p. 248.
Menado, N.-\V. Celebes.
190. CALLOSCIURUS PREVOSTI SCHLEGELI, Gray
1867. Ann. Mag. Nat. Hist. 3, XX, p. 278.
Koma, Celebes.
iqi. CALLOSCIURUS PREVOSTI BALUENSIS, Bonhote
1901. .■Xnn. Mag. Nat. Hist. 7, VII, p. 174.
Mt. Kina Balu, N. Borneo.
192. CALLOSCIURUS PREVOSTI SUFFUSUS, Bonhote
1901. Ann. Mag. Nat. Hist. 7, VII, p. 175.
Tutong River, N.-W. Borneo.
193. CALLOSCIURUS PREVOSTI RIFONIGER, Gray
1842. Ann. Mag. Nat. Hist. X, p. 263.
Labuan Island, iSl.-W. Borneo.
194. CALLOSCIURUS PREVOSTI PLUTO, Gray
1867. .Ann. Mag. Nat. Hist. 3, XX, p. 283.
Sarawak, Borneo.
195. CALLOSCIURUS PREVOSTI PICEUS, Peters
1866. Proc. Zool. Soc. London, p. 429.
Type locality uncertain.
Synonym: erebus. Miller. 1903, Proc. U.S. Nat. Mus. XXVI, p. 456.
Tapanuli Bay, N.-\V. Sumatra.
iy6. CALLOSCIURUS PREVOSTI NYX, Miller
1908. Proc. U.S. Nat. Mus. XXXIV, p. 638.
Pulau Rapat, E. Sumatra.
197. CALLOSCIURUS PREVOSTI NAVIGATOR, Bonhote
1901. Ann. Mag. Nat. Hist. 7, VII, p. 171.
Sirhassen, Natuna Islands.
198. CALLOSCIURUS PREVOSTI MIMELLUS, Miller
1900. Proc. Washington .Acad. Sci., II, p. 218.
Pulau Wai, Tambelan Islands.
199- CALLOSCIURUS PREVOSTI MIMICULUS, Milkr
1900. Proc. Washington Acad. Sci., H, p. 219.
St. Barbe Island, S. China Sea.
200. CALLOSCIURUS PREVOSTI CAEDIS, Chasen & Kloss
1932- Bull. Raffles Mus. 6, p. 25.
Balambangan Island, N. Borneo.
36S CALLOSCIURUS
201. CALI.OSCILRIS PRKVOSTI BANKS], Chasen
19^3. Bull. Raffles Mus. S, p. igj.
Baram district, Borneo.
202. CALLCISCICRCS PREVOSTI SL'.MATR.ANA, Schlc-gtl
I1S63. Xed. Tijds. Dierk. i, p. 25.
Sumatra.
203. CALLOSCIURUS PREVOSTI REDIMITUS, Boon Mesch
iSig. Verh. Ned. Ind. Inst. Amsterdam, II, p. 243.
(?) Sumatra. ("Kast Indies"); quoted by Robinson in Sumatran
Mammals list.
iiutatus Group
204. CALLOSCIURUS NIGRO\ITTATUS .MGROVITT.^TUS, Horsfield
1S24. Zool. Res. Java. p. 149.
Java; probably east central parts.
205. CALLOSCIURUS MGROVITT.'\TUS BESUKI. Kloss
1921. Journ. Fed. Malay States Mus. X, p. 231.
Tamansari, Idjen Massif, 1600 ft., E. Java.
2of.. CALLOSCIURUS NIGROVITTATUS BILIMITATUS, Miller
1903. Smiths. Misc. Coll. XLV, p. 8.
Tanjong Laboha, Trengganu, E. Malay Peninsula.
207. CALLOSCIURUS NIGROVITT.ATUS JOHOREXSIS, Rubinson & Wroughton
1911. Journ. Fed. Malay States Mus. IV, p. 166.
Pelepak, Johore, Malaya.
208. CALLOSCIURUS XIGROVITT,-\TUS MICRORHYXCHUS, Klo.ss
1 90S. Journ. Fed. Mai. States Mus. II, p. 144.
Juara Bay, Pulau Tioman, coast of Pahang.
20Q. CALLOSCIURUS NIGROVITTATUS BOCKI, Robinson & Wroughton
1911. Journ. Fed. Malay States Mus. IV, p. 167.
Pajo, Padang Highlands, W. Sumatra.
210. CALLOSCIL'RUS NIGROVITTATUS ORESTES, Thomas
1S95. .Ann. Mag. Xat. Hist. 6, XV. p. 530.
Mt. Dulit, Baram, Borneo.
211. CALLOSCIL'RUS NKiROVITTATUS KLOSSI, Miller
igoo. Proc. Washington, .Acad. Sci. II, p. 225.
Saddle Island, Tanibelan Group.
212. CALLOSCIURUS NIGROVITTATUS MADSOEDI, Sody
1929. Xat. Tijds. Xed. Ind. 89, p. 163.
Moeriah, Java.
213. CALL0SCIL:RUS XOTATUS NOTATUS, Boddacrt
1785. Elench. .Anim. p. 119.
West Java.
Synonym: hadgiitg, Kerr, An. Kingd. 1792, p. 262. Java.
plaiilaiii, Ljung. Vet. Akad. Handl. iSoi, p. 99. Java.
hilincatus, Desmarest, 1817, Xouv. Diet. Hist. Xat. X, p. 106.
Java.
gingiauus, Shaw, Cjen. Zool. 2, p. 147, 1801.
griseivcitlcr, GeofFroy, 1834, Voy. Bel. p. 147.
CALLOSCIURUS 369
214. CALLOSCIURUS NOTATUS TAMANSARI. Kloss
IQ2I. Journ. Fed. Malay States Mus. X, p. 230.
Tamansari, Idjcn Massif, 1600 ft., E. Java.
215. CALLOSCIURUS NOTATUS MADURAE, Thomas
1910. Ann. Map. Xat. Hist. 8, V, p. 386.
.Marengan, near Soemenep, E. Madura Island, N.-E. Java.
216. CALLOSCIURUS NOTATUS BALSTONI, Robinson & Wroughton
191 1. Journ. Fed. Malay .States Mus. I\', p. 234.
Tjilatjap, S. Central Java.
217. CALLOSCIURUS NOT.\TUS -STRESEMANNI, Thomas
1913. Ann. Mag. Nat. Hist. 8, XI, p. 503.
Baleling, Bali.
218. CALLOSCIURUS NOT.ATUS .MICROTIS, Jentink
1879. Notes Leyden Mus. i, p. 41.
Saleyer Island, Java Sea.
219. CALLOSCIURUS NOTATUS GUILLEMARDI. Kloss
1926. Journ. Malay Branch Roy. Asiat. Soc. 4, p. 260.
Tenggol Island, Malay Peninsula.
220. CALLOSCIURUS NOTATUS VANHEURNI, Sody
1929. Nat. Tijds. Ned. Ind. 88, p. 327.
Tjipanas, Garoet, Java.
221. CALLOSCIURUS NOT.-\TUS VERBEEKI, Sody
1929. Nat. Tijds. Xed. Ind. 88, p. 330.
Bandar, Distr. Padangan, Rembang, Java.
222. CALLOSCIURUS NOTATUS MALAWALI, Chasen & Kloss
1932. Bull. Raffles Mus. 6, p. 26.
Mallewalle Island, N. Borneo.
223. CALLOSCIURUS NOTATUS NICOTIANAE, Sody
1936. Nat. Tijds. Xed. Ind. 96, p. 217.
Kampong Silalas, near Medan, Deli, N. Sumatra.
224. CALLOSCIURUS ATRISTRLATUS, Miller
1913. Smiths. Misc. Coll. LXI, no. 21, p. 22.
Lo Bon Bon, Dutch S.-E. Borneo.
225. CALLOSCIURUS ANDREWSI, Bonhote
1901. .'^nn. Mag. Nat. Hist. 7, VH, p. 456.
Tjigombong, Java.
226. CALLOSCIURUS VITT.ATUS VITTATUS, Raffles
1822. Trans. Linn. Soc. XIII, p. 259.
Bencoolen, W. Sumatra.
Synonym: tarussamis , Lyon, 1907, Smiths. Misc. Coll. XI. VIII.
p. 279. Tarussan Bay, W". Sumatra.
227. CALLOSCIURUS VITT.-\TUS SATURATUS, Miller
1903. Proc. U.S. Nat. Mus. XXVI, p. 453.
Pulau Mansalar, oflf Tapanuli Bay, \V. Sumatra.
228. CALLOSCIURUS VITT.^TUS PRETIOSUS, Miller
1903. Proc. U.S. Nat. Mus. XXVI, p. 454.
Pulau Bangkaru, Banjak Islands, W. Sumatra.
24 — Living Rodents — I
370 CALLOSCIURUS
22<i. CALI.USCILRLS \ TITATUS LIBICRICOLOR, Miller
1901,. I'roc. U.S. Nat. Mus. XXVI, p. 455.
Pulau Tuanyku, Hanjak Islands, W. Sumatra.
230. CALLC)SLIIRL:S VITTATLS TAPANULIUS, Lyon
1907. Smiths. Misc. Coll. XLVIII, p. 280.
Tapanuli Bay, W. Sumatra.
231. CALLOSCIURUS VITTATUS PENINSULARIS, Milk-r
1903. Smiths. Misc. Coll. XLV, p. 10.
N. bank of Endau River, S.-E. Pahang, Malaya.
232. CALLOSCIURUS VITTATUS RUP.ATIUS, Lyon
190.S. Proc. U.S. Nat. Mus. XXXIV, p. 640.
Pulau Rupat, E. .Sumatra.
233. CALLOSCIURUS VITTATUS SUBLUTEUS, Thomas & Wroughton
1909. Ann. Mag. Nat. Hist. 8, III, p. 440.
Si Karang, S.-E. Johore, Malaya.
234. CALLOSCIURUS VITTATUS SINGAPLTRENSIS, Robinson
191(1. Journ. Fed. Malay States Mus. VII, p. 73.
Changi, Singapore Island.
235. CALLOSCIURl'S VITTATUS MAPORENSIS, Robinson
1916. Journ. Fed. Malay States Mus. VII, p. 64.
Pulau Mapor, Rhio-Lingga Archipelago.
236. CALLOSCIURUS VITTATUS NESIOTES, Thomas & Wroughton
1909. .Ann. Mag. Nat. Hist. 8, III, p. 440.
Pulau Hatam, Rhio-Lingga Archipelago.
237. CALLOSCIURUS VITTATUS TENUIROSTRIS, Miller
1900. Proc. Washington Acad. Sci., II, p. 221.
Tioman Island, off coast of Pahang.
238. CALLOSCIURUS VITTATUS ANAMBENSIS, Miller
1900. Proc. Washington Acad. Sci., II, p. 223.
Pulau Siantan, Anamba Islands.
239. CALLOSCIURUS VITTATUS ABBOTTI, Miller
1900. Proc. Washington Acad. Sci., II, p. 224.
Big Tambelan Island, S. China Sea.
240. CALLOSCIURUS VITTATUS AORIS. Miller
1903. Smiths. Misc. Coll. XLV, p. ro.
Pulau Aor, near Pulau Tioman.
241. CALLOSCIURUS VITTATUS FA.ML'LUS, Robinson
191 2. Ann. Mag. Nat. Hist. 8, X, p. 592.
Pulau Dayang, near Pulau Aor, S. China Sea.
242- CALLOSCIURUS VITTATUS PAxNNOVIANUS, Miller
1903. Smiths. .Misc. Coll. XLV, p. II.
Pulau Panau, Atas Islands, S. China Sea.
243. CALLOSCIURUS VITTATUS PEMANGILENSIS, Miller
1903. Smiths. Misc. Coll. XLV, p. 9.
Pemangil Island, near Pulau Tioman.
CALLOSCIURUS 37i
244. CALLOSCIURUS VITTATUS ICTERICUS, Miller
1903. Smiths. Misc. Coll. XLV, p. 12.
Tana Bala, Hatu Island, W. Sumatra.
245. CALLOSCIURUS VITTATUS SERUTUS, Miller
1906. Proc. U.S. i\"at. Mus. XXXI, p. 58.
Pulau Scrutu, Karimata Islands.
246. CALLOSCIURUS VITTATUS DIRECTOR, Lyon
1909. Proc. U.S. Nat. Mus. XXXVI, p. 509.
Direction Island, S. China Sea.
247. CALLOSCIURUS VITT.ATUS LUTESCENS, Miller
1901. Proc. Washington Acad. Sci., Ill, p. 124.
Sirhassen Island, Natuna Islands.
248. CALLOSCIURUS VITT.VPUS LAMUCOTANUS, Lyon
191 1. Proc. U.S. Xat. Mus. XL, p. 85.
Pulau Lamukotan, W. Borneo.
249. CALLOSCIURUS VITTATUS D.\TUS, Lyon
1911. Proc. U.S. Nat. Mus. XL, p. 86.
Pulau Dato, W. Borneo.
250. CALLOSCIURUS VITTATUS SIRIENSIS, Lyon
191 1. Proc. U.S. Nat. Mus. XL, p. 87.
Pulau Mata Siri, Java Sea.
251. CALLOSCIURUS VITTATUS ARENDSIS, Lyon
1911. Proc. U.S. Nat. Mus. XL, p. 87.
.Arends Island, Java Sea.
252. CALLOSCIURUS VITT.\TUS MARINSULARIS, Lyon
1911. Proc. U.S. Nat. Mus. XL, p. 89.
Pulau Laut, off S.-E. Borneo.
253. CALLOSCIURUS VITTATUS LAUTENSIS, Miller
1901. Proc. Washington Acad. Sci., Ill, p. 128.
Pulau Laut, N. Natuna Islands.
254. CALLOSCIURUS VITTATUS RUTILIVENTRIS, Miller
1901. Proc. Washington Acad. Sci., Ill, p. 126.
Pulau Midei, S. Natuna Islands.
255. CALLOSCIURUS VITTATUS RUBIDIVENTRIS, Miller
1901. Proc. Washington Acad. Sci., III. p. 127.
Bunguran, Natuna Islands.
256. CALLOSCIURUS VITTATUS Si:iCA.IAE, Miller
1901. Proc. Washington Acad. Sci., Ill, p. 125.
Pulau Seraia, Natuna Islands.
257. CALLOSCIURUS VITT.ATUS ALBESCENS, Bonhote
1901. Ann. Mag. Nat. Hist. 7, VII, p. 446.
.•\cheen, N. Sumatra.
258. CALLOSCIURUS VITTATUS DULITENSIS, Bonhote
1901. Ann. Mag. Nat. Hist. 7, VII, p. 451.
Mount Dulit. Baram, Borneo.
259. CALLOSCIURUS VITT.VrUS DILUTUS, Miller
1913. Smiths. Misc. Coll. LXI, no. 21, p. 23.
Tanjong Batu, S.-E. Borneo.
372 CALLOSCIURUS
;(.o. CAI.H).SCR RL S VITTATUS COMPUS, Lyon
irjii. I'roc. Biol. Soc. Washington, XXIV. p. t)8.
Pamiiiiang Bay, S. Borneo.
Synonym: poliopiis, Lyon, 191 1, Proc. L'.S. Nat. Hist. XL, p. 88, pre-
occupied.
zb\. CALLOSCIURUS VITTATUS TICDONGUS, Lyon
ii;o7. I'roc. L'.S. Nat. Mus. XXXI, p. 591.
Tanjong Tedong, Banka Island.
2(1-. CALLOSCIURUS VITTATUS BILLITONUS, Lyon
1907. Proc. U.S. Nat. Mus. XXXI, p. 592.
Buding Bay, Billiton Island.
263. CALLOSCIURUS VITTATUS MINIATUS, Miller
I93D. Proc. Washington .\cad. Sci., II, p. 79.
Trang, Siamese Malaya.
;fM CALLOSCIURl'S VITTATUS SCOTTI, Kloss
191 1. .Ann. Mag. Nat. Hist. 8, VII, p. 117.
Bedimg Island, off Trengganu, E. Malay Peninsula.
265. CALLOSCIURl'S VITTATUS PLASTICUS, Kloss
1911. .Ann. Mag. Nat. Hist. 8, VII, p. 117.
Crreat Redang Island, off Trengganu, E. Malay Peninsula.
zbl: CALLOSCIURUS VITTATUS PERHENTIANI, Kloss
191 1. .Ann. Mag. Nat. Hist. 8, VII, p. 118.
W. Perhentian Island, off Trengganu, E. Malay Peninsula.
267. CALLOSCIURUS VITT.ATUS PROTEUS, Kloss
1911. Ann. Mag. Nat. Hist. 8, VII, p. 118.
E. Perhentian Island, off Trengganu, E. Malay Peninsula.
26S. CALLOSCIURUS VITTATUS WATSONI, Kloss
191 1. Ann. Mag. Nat. Hist. 8, VII, p. 118.
Lantinga Island, off Trengganu, E. Malay Peninsula.
26Q. CALLOSCIURUS VITTATUS LIGHTI, Chaseii & Kloss
1924. Journ. Malay Branch Roy. Asiat. Soc. 2, p. 58.
Penang Island, Malaya.
270. CALLOSCIURUS VITTATUS STELLARIS, Chascn & Kloss
1924. Journ. Malay Branch Roy. Asiat. Soc. 2, p. 58.
Bintang Island, Malaya.
271. CALLOSCIURUS VITTATUS LUNARIS, Chasen & Kloss
1924. Journ. Malay Branch Roy. Asiat. Soc. 2, p. 58.
Bulan Island, Malaya.
272. CALLOSCIURUS ADAMSI, Kloss
1 92 1. Journ. Straits Branch Roy. Asiat. Soc. 83, p. 151.
N. Sarawak, Borneo (Baram Riyer).
pvgeivthnis Group
273. CALLOSCIURUS LOKROIDES LOKROIUES, Hodgson
1S36. Journ. .As. Soc. Bengal, V, p. 232.
Sikkim.
.Synonym; assamensis. Gray, ex McClelland, 1843. List. Mamm. p. 143;
nom. nud.
CALLOSCIURUS 373
274. CALLOSCIURUS LOKROIDES OWENSI, Thomas & Wroughton
1016. Journ. Bombay Nat. Hist. Soc. XXIV, p. 236.
Minsin (east bank). Upper Chindwin, Burma.
275. CALLOSCIURUS LOKKOIUKS SI.MILIS, Gray
1867. Ann. Mag. Nat. Hist. 3, XX, p. 281.
Sikkim.
276 CALLOSCIURUS BLYTHI BLVTHI. Tytler
1854. .Ann. Mag. Nat. Hist. 2, XIV, p. 172.
Dacca, Bengal, India.
277. CALLOSCIURUS BLVTHI MEARSI, Bonhotc
1906. Ann. Mag. Nat. Hist. 7, XVIII, p. 337.
Chinbyit, Lower Chindwin, Burma.
278. CALLOSCIURUS BLYTHI BELLONA, Thomas & Wroughton
1916. Journ. Bombay Nat. Hist. Soc. XXIV, p. 420.
Kin, Middle Chindwin, Burma.
279. CALLOSCIURUS BLYTHI VIRGO, Thomas & Wroughton
1916. Journ. Bombay Nat. Hist. Soc. XXIV, p. 421.
Tatkon, Upper Chindwin, Burma.
280. CALLOSCIURUS STEVENSI, Thomas
1908. Journ. Bombay Nat. Hist. Soc. XVIII, p. 246.
Beni-Chang, .Abor-Miri Hills, Upper Assam.
281. CALLOSCIURUS PYGERYTHRUS PYGERY'THRUS, Geoffrey
1832. Mag. Zool. CI. I ; Belanger Voy. Zool. p. 145, pi. vii, 1847.
Pegu, Burma.
Synonym: inornatus. Gray, .Ann. Mag. Nat. Hist. 1867, 3, XX, p. 282.
282. CALLOSCIURUS PYGERYTHRUS JANETTA, Thomas
1914. Journ. Bombay Nat. Hist. Soc. XXIII, p. 203.
Mandalay, Upper Burma.
283. CALLOSCIURUS PH.A.YREI PHAY'REI, Blyth
1855. Journ. .Asiat. Soc. Bengal, XXIV, pp. 472, 476.
Aloulmein, Burma.
284. CALLOSCIURUS PH.AYREI BLANFORDI, Blyth
1862. Journ. Asiat. Soc. Bengal, XXXI, p. 333.
Ava, Upper Burma.
quinquestriatus Group
285. CALLOSCIURUS QUINQUESTRI.\TUS QUINQUESTRIATUS, Anderson
1871. Proc. Zool. Soc. London, p. 142, pi. x.
Ponsee, Kakhyen Hills, Yunnan border.
Synonym: beebei, Allen, 1911, Bull. .Amer. Mus. Nat. Hisl. XXX,
p. 338. Kuching, Sarawak (erroneous).
286. CALLOSCIURUS QUINQUESTRIATUS IMARIUS, Thomas
1926. .Ann. Mag. Nat. Hist. 9, XVII, p. 640.
Kachin, N. Burma.
287. CALLOSCIURUS QUINQUESTRI.ATUS SYLVESTER, Thoma.-:
1926. Ann. Mag. Nat. Hist. 9, XVII, p. 641.
Schweli-Salween Divide, W. Y'unnan.
374
CALLOSCIURUS
liippurus Group
zSS. CALLUSCIL'RUS Hll'l'l RLS HIl'PURUS, Gcoffroy
1S32. Mag. Zool. CI. I, pi. VI.
Java (erroneous), substitute Alalacca.
Svnonvm: riifogastcr. Gray, 1842, Ann. Mag. Nat. Hist. X, p. 263.
Malacca.
28q. CALLOSCIURUS HIPPURUS GRAYI, Bonhote
1901. Ann. Mag. Nat. Hist. 7, VH, p. 171.
Sarawak, Borneo.
200. CALLO.SCIURUS HIPPURUS HIPPUROSUS, Lyon
1907. Smiths. Misc. Coll. L, Pt. i, p. 26.
Tarussan Bay, \V. Sumatra.
2Q1. CALLOSCIURUS HIPPURUS HIPPURELLUS, Lyon
1907. Smiths. Misc. Coll. L, Pt. i, p. 27.
Batu Ampar, Landak Range, W. Borneo.
292. CALLOSCIURUS PRYERI PRYERI, Thomas
1892. Ann. Mag. Nat. Hist. 6, X, p. 214.
Near Sandakan, British N. Borneo.
293. CALLOSCIURUS PRYERI INQUINATUS Thomas
190S. Journ. Bombay Nat. Hist. Soc. XVIII, p. 247.
Lawas River, N.-W. Borneo.
294. CALLOSCIURUS MELANOGA.STER MELANOGASTER, Thomas
1895. Ann. Mus. Civ. Stor. Genoa, XIV, p. 668.
Sipora, Mentawei Islands. W. Sumatra.
295. CALLOSCIURUS MELANOGASTER ATRATUS, Miller
1903. Smiths. Misc. Coll. XLV, p. 13.
N. Pagi Island, W. Sumatra.
296. CALLOSCIURUS MELANOGASTER MENTAWI, Chasen & KIoss
1928. Proc. Zool. Soc. London, p. 822.
Siberut Island, W. Sumatra.
297. CALLOSCIURUS BROOKEI. Thomas
1S92. Ann. Mag. Nat. Hist. 6, IX, p. 253.
Sarawak, Borneo.
298. CALLOSCIURUS SAMARENSIS, Steere
1890. List Birds. Mamm. Steere Exp. Philippines, p. 30.
Saniar, Philippine Islands.
299. CALLOSCIURUS MINDANENSIS, Steere
1890. List Birds. Mamm. Steere Exp. Philippines, p. 29.
Mindanao, Philippine Islands.
Synonym: cagsi, Meyer, 1890, Proc. Zool. Soc. London, p. 600.
Davao, S. Mindanao.
300. CALLOSCIURUS PHILIPPINENSIS, Waterhouse
1839. Proc. Zool. Soc. London, p. 117.
Mindanao, Philippine Islands.
301. CALLOSCIURUS STEERII, Gunthcr
1876. Proc. Zool. Soc. London, p. 735, pi. xix, fig. i.
Balabac Island, Philippine Islands.
CALLOSCIURUS 375
302. CALLOSCIURUS JUVENCUS, Thomas
1908. Ann. Mag. Nat. Hist. 8, 11, p. 498.
Palawan, Philippine Islands.
303. CALLOSCIURUS MOLLICNDORFI-I, Matschie
1898. Sitz. Ber. Ges. Nat. Fr. Berlin, 5, p. 41.
Calamianes, Philippine Islands.
304. CALLOSCIURUS ALBICAUDA, Matschie
1898. Sitz. Ber. Ges. Nat. Fr. Berlin, 5, p. 42.
Calamianes, Philippine Islands.
leucomus Group
305. CALLOSCIURUS LEUCOMUS LEUCOMUS, Muller & Schlegel
1839. Verhandl. Nat. Gesch., p. 87.
Celebes.
306. CALLOSCIURUS LEUCOMUS OCCIDENTALIS, Meyer
1898. Abh. Mus. Dresden, no. 4, p. 2.
West Celebes.
307. CALLOSCIURUS TOPAPUENSIS, Roux
1910. Zool. .Anz. 35, p. 518.
Mt. Topapu, Central Celebes.
308. CALLOSCIURUS MOWEWENSIS, Roux
1910. Zool. Anz. 35, p. S19.
Mowewe, S.-E. Celebes.
309. CALLOSCIURUS ELBERTAE, Schwarz
1911. Ann. Mag. Nat. Hist. 8, VII, p. 639.
E. Kabaena, off Celebes.
310. CALLOSCIURUS TONKEANUS, Meyer
1896. Abh. Mus. Dresden, no. 6, p. 25, pi. x, fig. i.
Tonkean, Celebes.
311. CALLOSCIURUS SARASINORUM, Meyer
1898. Abh. Mus. Dresden, no. 4, p. i.
Central Celebes.
312. CALLOSCIURUS WEBERI, Jentink
1890. Weber. Zool. Ergebn. i, p. 115, pis. viii, x.
Central Celebes.
313. CALLOSCIURUS TINGAHI, Meyer
1896. .A.bh. Mus. Dresden, no. 6, p. 27, pi. x, fig. 4.
Sangir Islands, Celebes.
314. C.-\LLOSCIURUS ROSENBERGI, Jentink
1879. Notes Leydcn Mus. p. 37.
Sangir Islands, Celebes.
rubriventer Group
315. CALLOSCIURUS RUBRIVENTER, Forslen
1839. Muller & Schlegel, Verhandl. Nat. Gesch. p. 86.
Minahassa, N. Celebes.
376 CALLOSCIURUS— FUNAMBULUS
incerlac sedis; not allocated to nroiips
316. CALLOSCIURUS ALSTON'L Anderson
iSyg. ZooL \V. Yunnan, p. 252, pL xxi.
(?) Borneo.
317. CALLOSCIURUS DIARDL Jintink
1S79. Notes Leyden Mus. i, p. 39.
Nusa Kambangan, off Tjilatjap, Java.
318. CALLOSCIURUS CHINENSIS, Gray
1867. Ann. Mag. Nat. Hist. 3, XX, p. 282.
"China" (based on a specimen of tenuis^).
Addenda: , ,, ,. „
maclcllandi Group
CALLOSCIURL'S MACLKLLAXDl DOLPHOIDES, Kloss.
1921. Journ. Nat. Hist. Soc. Siam, IV, p. loi.
Kompong Som Bon, Cambodia.
ervt/iraciis (jroup
CALLOSCIURUS FINLAYSONI R,\JASLMA, Kloss.
1920. Journ. Nat. Hist. Soc. Siam, IV, p. 103.
Lat Bua Kao, East Siam.
CALLOSCIURUS FINL.'\YSOXI PRACHIX. Kloss.
1916. Journ. Nat. Hist. Soc. Siam, II, p. 16.
Krabin, Central Siam.
There are also at the British Museum skulls bearing the names '' nigro-
vittatiis rubrigula" and " caniceps tahitiis." The references to these races have
not been traced.
Genus 9. FUNAMBULUS, Lesson
1S35. FuNAMBULUs, Lesson, Illustr. de Zool. (15) pi. 43, 2 pp. text.
1S90. EoxERUS, Forsyth Major, Proc. Zool. Soc. London, p. 1S9, part, subgenus of
Xenis containing species now referred to this genus, ^lenctes, Lariscus and Rhino-
sciiirus.
1923. Tamiodes, Pocock, Proc. Zool. Soc. London, p. 215. (Fimamhulus trislriatns,
Waterhouse.)
Type Speciks. — Fiiiuiiiihidiis indiciis, l^csson — Scimus palmanim, Linnaeus.
Range. — Ceylon, Southern Peninsular India, north to Sural, Palanpur,
Central Provinces, Bihar, Rawalpindi (North Punjab), and
Baluchistan.
Number of For.ms. — Twenty-four.
Characters. — Skull with more or less elongate rostrum; postorbital process
not large; parietal ridges frequently joined together. Zygo-
matic plate as a rule slanting upwards far forwards, and relatively prominently
ridged. Infraorbital foramen normal. Palate normal. Coronoid process
usually low. Cheekteeth with upper series characterized in the young by notice-
ably high cusps; P. 3 well developed; the ridges high and the depressions deep,
FUNAMBULUS 377
in the main teeth ; the small outer third cusp, usually present in Sciurus, is gener-
ally absent or scarcely traceable in this genus. Lower cheekteeth with four well-
marked cusps each tooth, the small subsidiary cusps of Sciurus usually not clear;
the central depression tending to be rather smaller than is usual in Sciurus or
Ctillusciurus; the transverse ridge extending from theanteroexternaltotheantero-
internal cusp present.
Forsyth Major transferred this group to Xertis (with Menetes, Lariscus,
Rbinosciurus, etc.), evidently rather on cranial than dental characters, remarking,
"the less semihypsodont oriental S. tristriatus and S. palmarum tend to connect
the Xerus type with the Sciurus vulgaris type (of tooth) in approaching the form
of molar of most of the middle-sized Oriental Squirrels" (referring to Callo-
sciurus).
But no one, so far as I have traced, has ever defined this genus, and to do so
is no easv matter. Thomas proposed that all subgenera of Forsyth Major (of
Xerus and Sciurus) should be given generic rank ; but the abov'e-mentioned dental
characters are scarcely of generic importance. The characters of the baculum,
so far as I have read, while very variable within the genus, are not those either
of Sciurus or Callosciurus. Other than the fact that the cusps are noticeably
high as a rule in the cheekteeth (when young), and that the coronoid process
appears rather low, and the zygomatic plate which seems different from Sciurus,
though not from Callosciurus in every case, I can find no constant difference
between this genus and Callosciurus on the one hand, Sciurus on the other. As
I feel uncertain to which of these Funambulus stands nearest, and it has long
been regarded as forming a natural group, I retain it. Externally with three
prominent white stripes usually present (five in pennanti); the central one is
mid-dorsal. In sublineatus, a small thick-furred type with the tail normally
more narrow than is usual, these stripes much reduced. In layardi, all but the
mid-dorsal stripe are becoming reduced. These two species are rather darker
than the other forms. Fur normally rather coarse and short. Tail not reduced
in length; digits normal (arboreal type).
Pocock erecxedl'amiodes in 1923, based solely on formation of baculum, for
F. tristriatus. In 1936, Osman Hill (Ceylon Journ. Sci. Section B, Zool. Sc Geol.,
XX, pt. I, p. 100) reviewed the penial characters of the squirrels of Ceylon, and
remarks: "The Striped Squirrels . . . form a very difficult problem. According
to Pocock's definitions, the Ceylon race of palmarum would fall into the genus
Tamiodes, whilst lavardi, with its conical appendage on the tip of the glans would
fall into Funambulus. Probably sublineatus on the characters of its glans would
fall between the two, though on its baculum it would require a new genus. It
seems almost absurd that different geographical races of what would otherwise
be regarded as one and the same species should on their penial characters require
separate genera, though theoretically one is bound to admit that this is the correct
procedure. Until more species have been examined ... I consider that it is
best to retain Funambulus for all these Striped-squirrels, despite their penial
differences. The alternative is to re-define the genera Tamiodes and Funambulus
to fit the new knowledge, and probably in addition to institute a third genus for
sublineatus." In the same paper he writes: " Ratufa would appear to be very
37S FUNAMBULUS
different from the smaller Squirrels in its penial characters, but there is appar-
ently less uniformity through the genus than would have been expected. The
differences of R. tnacroura from the others cannot at this stage, however, be
granted to be of generic importance, though differences of similar order have
been used bv Pocock in separating some of the smaller Squirrels generically."
It is clear that if each species (or subspecies) which differs in this respect is
to be given a new generic name, we shall soon have to deal with over a thousand
genera in this Order! But whereas differences in baculum may be valid as
regards solving the problem of whether two forms belong to the same species or
not, I am strongly of opinion that no generic names based solely on the shape of
this organ can be retained. This view is held by Howell, 1938, who remarks:
"The writer does not believe, however, that in the absence of trenchant
cranial characters, the morphology of the baculum alone should be considered
of generic value."
List of N.\med Forms
ptdmarum Group
1. FUN.^MBULUS P.\LM.^RL'M P.-\LMARUM, Linnaeus
1766. Syst. Xat. 1, p. 86.
Madras.
Svnonym; penidUatus, Leach, 1814, Zool. Misc. i, p. 6, pi. i.
indicus, Lesson, 1835, lUustr. de Zool. (15) pi. 43.
2. FUNAMBULUS PALMARUM COMORINUS, WrouKhton
1905. Jiuirn. B.mihay Nat. Hist. Soc. XVI, p. 411.
Trivandrum, Travancore.
3. FUNAMBULUS PAL^LARUM BELLARICUS, Wroughton
1916. Journ. Bombay Xat. Hist. Soc. XXIV, p. 647.
Vizayanagar, Bellary, South India.
4. FUNAMBULUS PALMARUM FAVONICUS, Thomas & Wroughton
1915. Journ. Bombay Nat. Hist. Soc. XXIV, p. 39.
Udugama, Southern Province, Ceylon.
5. FUNAMBULUS PALMARUM KEL,\ARTL Layard
1849 Blyth, Journ. Asiat. Soc. Bengal, XVIII, p. 603. footnote, id. op. cit. XX, p. 166,
1S52.
Hambalotte, Ceylon.
6. FUN.^MBULUS PALMARUM BRODIIU, Blyth
1849. Journ. Asiat. Soc. Bengal, XVIII, p. 602.
Point Pedro, Ceylon.
7. FUN.-\MBULUS P.\LMARl'M OLYMPIUS, Thomas & Wroughton
1915. Journ. Bombay Nat. Hist. Soc. XXIV, p. 41.
Urugalla, Highlands of Central Ceylon.
8. FUNAMBULUS P.^LMARUM ROBKRTSONT, Wroughton
1916. Journ. Bombay Nat. Hist. Soc. XXIV, p. 647.
Pachmarhi, Hoshangabad, Central Provinces. India.
>,. FUNAMBULUS PALMARUM BENGALENSIS. Wroughton
191b. Journ. Bombay Nat. Hist. Soc. XXIV, p. 648.
Hazanbagh, Bengal (now Bihar).
FUNAMBULUS 379
10. FUNAMBl'LUS PALMARLM MATLCJAMKNSIS, Lindsay
1926. Journ. Bombay Nat. Ilist. Soc. XXXI, p. 239.
Matugama, Western Province, Ceylon.
11. FUNAMBL'l.US THOMA.SI, Wrouehton & Davidson
1919. Journ. Bombay Nat. Hist. Soc. XXVI, 3, p. 729.
Khandalla, Bombay Presidency.
12. FUNAMBULUS GOSSEI, Wroughton & Davidson
1919. Journ. Bombay Nat. Hist. Soc. XXVI, 3, p. 730.
Kotaf;iri, Nilgiris, India.
13. FUNAMBULUS PENNANTl PENNANTI, WiouKhton
1905. Journ. Bombay Nat. Hist. Soc. XVI, 3, p. 411.
Mandvi Taluka, Sural district, Bombay.
14. FUNAMBULUS PENNANTI ARGENTESCENS, Wroughton
1905. Journ. Bombay Nat. Hist. Soc. XVI, p. 413.
Rawalpindi, North Punjab.
15. FUNAMBULUS PENNANTI LUTESCENS, Wroughton
1916. Journ. Bombay Nat. Hist. Soc. XXIV, p. 430.
Deesa, Palanpur.
16. FUNAMBULUS TRISTRIATUS TRISTRIATUS, Waterhouse
1837. Charlesworth's Mag. Nat. Hist, i, pp. 496-9.
Madras (by designation).
Synonym: dussumieri, Milne-Edwards, 1867, Rev. Zool. XIX, p. 226.
Malabar.
17. FUNAMBULUS TRISTRIATUS NUMARIUS, Wroughton
1916. Journ. Bombay Nat. Hist. Soc. XXIV, p. 646.
Helwak, Satara district. Western Ghats, Bombay.
18. FUNAMBULUS TRISTRIATUS ANNANDALEI, Robinson
1917. Rec. Ind. Mus. XIII, p. 41.
Shasthancotta, west of Western Ghats, Travancore.
ig. FUNAMBULUS WROUGHTONI, Ryley
1913. Journ. Bombay Nat. Hist. Soc. XXII, p. 437.
Makut, S. Coorg.
layardi Group
20. FUNAMBULUS LAYARDI LAYARDI, Blyth
1849. Journ. Asiat. Soc. Bengal, XVIII, p. 602.
Ambegamoa Hills, Ceylon.
21. FUNAMBULUS LAYARDI DRAVIDIANUS, Robinson
1917. Rec. Ind. Mus. XIII, p. 42.
West side Western Ghats, Travancore.
22. FUNAMBULUS L.\YARDI SIGN.'^TUS, Thomas
1924. Ann. Mag. Nat. Hist. 9, XIII, p. 241.
Ratnapura, S. Ceylon.
sublineatus Group
23. FUNAMBULUS SUBLINEATUS SUBLINEATUS, Waterhouse
1838. Proc. Zool. Soc. London, p. 19.
Nilgiris, India.
SynonjTn: delesserti, Gcrvais, 1841, L'Institut, p. 171. Nilgiris.
3So FUNAMBULUS— DREMOMYS
24. FUNAMBULUS SUBLINEATUS OBSCIKUS. Pclzuln & K.ilil
1SS6. Verh. Zool. Rot. Ges. Wien, XXXV, p. 525.
Uplands of Ceylon.
Synonym: kat/tleenae, Thomas & VVroii^hton, Jonrn. Boniha\' Xat.
Hist. Soc. XXIV, p. 3S. 1915. Kottawa, .South
Province, Ceylon.
irilinealus, Kclaart, Prodr. Faun. Zeylon, p. 54, 1852; for
status see Robinson S: Kloss, Nominal List Oriental
Sciuridae, Rec. Indian Mus. XV, pt. 4.
Forms seen: aygciitcscon, hellaricus, brodiei, coinorinus, fuvoiiiciis. gossei,
katlih'i'iuie, kchuiiti. /iivtiicli, liitcscens, mntiigtiineiisis, iniinan'iis, ohscunis, o/vmpiis,
pahiuirum, pviuuinti, lohcrtsoni, siginitiis, subliiieatiis, thomasi, tristriaiiis, lurough-
toni.
Genus 10. DREiMOiMY.S, Ileude
iSqS. Dremomys, Heude, Mem. Hist. Nat. Emp. Chinois. IV, pt. 2, p. 54.
igoS. Zetis, Thomas, Journ. Bombay Nat. Hist. Soc. XVIII, p. 245. (Sciurns rufigenis,
Blanford.)
Typi; Species. — Sciurns penivi, Milne-Edwards.
Range. — Indo-Malayan chiefly, but touching extreme south of Palaearctic
China. Ilupeh, Szechuan; Fukien, Kweichow, Anhwei, Kwan-
tune, Yunnan; Formosa, Hainan; Nepal, Sikkim, Assam, Burma, Tenasserim;
Tongking, Annani, south to Selangor; Borneo.
Number of Forms. — About twenty-nine.
Ch.'\r.\cters. — This genus, which is said to agree with the " Toiiieiites"
section of Callosciiiriis in the characters of the baculum, is
only separable on average characters from that genus. The rostrum becomes
progressively elongated until at extreme development it is abnormal, being
second only to the extreme genus Rliinosciiirus. But in D. lokn'a/i, which is
the shortest-nosed species of the genus, the rostrum is probably not longer than
in some forms of CaUosciitrus.
The lachrymal is situated farther back in relation to the toothrows than is
normal, and the postorbital process is not very far in front of the level of the
posterior zygomatic root. The parietal ridges evidently do not join, or very
rarely tend to come together. Bullae often relatively small. Zygomatic plate
moderately ridged, not essentially different from Sciurns. Frontals broad;
postorbital process moderate. Cheekteeth evidently not essentially different
from Sciurus. Lower cheekteeth without abnormalities. P. 3 present. Forsyth
Major referred a species of this genus to Sciurns, in his paper on the dental
characters of the family; later the group was transferred, with Rhiiiosciurus,
Meuetes, and Lariscus, to the genus Fuiuiinhulus; still later Thomas erected Zetis
for the group, which is antedated by Dremomys, I leude. It is evidently a natural
group, though as indicated above very close to Callosciurus.
Tail rather shorter than head and body. Fur thick and soft. Flindfoot
rather narrow; digits arranged in the manner characteristic of normal Tree-
squirrels. The rostrum apparently reaches its extreme development in members
of the rufigenis group.
DREMOMYS 381
Forms seen : adumsom, bclfielJi, hhutia, culidiur, chintulis, everetii, jlavior,
fiisciis, ^aroniim, griselda, ^iihiris, lioivelli, imus, laomaclie, licliiensis, lukriah,
mcicmi/hini, mentosiis, mudestiis, opimus, ornatus, owstoni, pernyi, pyrrhomerus,
rufigenis, senex, " subflaiiientris."
I am inclined provisionally to recognize three groups in this genus:
rufigenis group : rostrum usually extreme, cheeks usually red, underside of
tail bright red throughout its length.
lokriah group: rostrum apparently shortest of genus; belly bright yellow.
D. macmiUoni appears to me to he not more than racially distinct from
lokriah. Underside tail and cheeks not red, so far as seen.
pernyi group: the other species; rostrum moderate to extreme; underside
tail and cheeks not red, so far as seen ; belly usually white (transitionary
towards lokriah in D. ozvstoni). D. everetti, Borneo, appears to have a
narrower shorter tail than is usual in the genus.
List of Named Forms
lokriah Group
1. DREMOMYS LOKRI.^H l.OKRI.'VH, Hodgson
1836. Journ. Asiat. Soc. Bengal, V, p. 232.
Nepal.
2. DREMOMYS LOKRIAH BH(JTIA, Thomas
1916. Journ. Bombay Xat. Hist. Soc. XXIV, p. 426.
Sedonchen, E. Sikkim.
Synonym: (?) subflmiventris. Gray, 1843, Hand List Mamm. Brit. Mus.
p. 144. (?) Assam. Considered a «om«i«H(f«m by Robin-
son & Kloss. 1918.
3. DREMOMYS LOKRIAH GARONUM, Thomas
1922. Journ. Bombay Nat. Hist. Soc. XXVni, 2, p. 430.
Tura, Garo Hills, Assam.
4. DREMOMYS LOKRIAH MACMILLANI, Thomas
1916. Journ. Bombay Nat. Hist. .Soc. XXIV, p. 238.
Kindat, Chindwin River, Upper Burma.
(Listed as a valid species by Robinson & Kloss.)
pernyi Group
5. DREMOMYS PERNYI PERNYI, Milne-Edwards
1867. Rev. et Mag. Zool. p. 230, pi. XIX.
Szechuan, China.
6. DREMOMYS PERNYI FLAVIOR. Allen
1912. Proc. Biol. Soc. Washington, XXV, p. 178.
S.-E. Yunnan, China.
7. DREMOMYS PERNYI C;RISELDA, Thomas
1916. Ann. Mag. Nat. Hist. 8. XVII, p. 392.
Nagchuka, \V. Szechuan, China.
DREMOMYS
'^' S DRVMC.MVS PERNVl MODESTUS, Thomas
„ DREMOMVS PKRNYI SENF.X, Allen
,o:. Mem. Mus. Harvard, XL, r.0. 4. P- "9-
^ Ichanc China.
.0 DRKMOMYS PERNYl CHINTALIS, Thomas
K Ann Mag Nat. Hist. 8, XVII, p. 394-
Kj.b. .Ann. ^la|;^-^^^^.j,_ ^h-we,, China.
„ DREMOMYS PERNYl CALIDIOR. Thomas
/ inn Mac Nat. Hist. 8, XVII, p. 394,
,g,6. Ann. ^I'^B.^^^^^ j^ .w. Fo-K.en, China.
„ DREMOMYS PERNYl LICHIENSIS, Thomas
,,./ Ann. Ma^^N- H.st^9, X, ^3.^ ^^__^^
,, DREMOMYS PERNYl HOWELLl. Thomas
,,,._: Ann. >I^S,,^-^,,';JfKa;,':;;fr-T°engyueh, Upper Irrawaddy, Burma.
„ DREMOMYS PERNYl MENTOSUS, Thomas
inn Mag Nat. Hist, y, X, p. 401-
„ DREMOMYS PERNYl IMUS, Thomas
--■^-^'^li^;^'^-'^^~
, OREMOMYS OWSTONl, Thomas
,- DREMOMYS EVERETTl, Thomas
o ' inn Mag Nat. Hist. 6, VI, p. 71-
1890. Ann. Mag.^.^^^^^^^ ^^^^^^^^^ g^^^„.
nifn^enis Group
,. DREMOMYS -^--i^^r^rS^^S
, DREMOMYS RUFIGENIS BELFIELDI, Bonho.e
k;. DREMO III, p. 9, pl- ' •
,908. Journ. F\^j^,'„t^f ;f selangor, Malay Peninsula.
.0. DREMOMYS RUEIGEN.S FL^;;^';;«'^"f ^';„, ^oc. London, .907. P- xo.
,007 Abstr. Proc. Zool. Soc. London, p. 2, Froc. A
^ Bali. Annam.
„ DREMOMYS RUFIGENIS ADAMSONl Thomas
,01; Journ. Bombay Nat. Hist. hoc. XXIII, p. 2,.
,914. jour M3^,„„o, Upper Burma.
,,. DREMOMYS RUFIGENIS ORNATUS, Thomas
,0,4 Journ. Bombay Nat. Hist. Soc^XXIII, p. 26.
^^ Near Mong-tze, Yunnan.
DREMOMYS— RATUFA 383
23. DRIIMOMYS RUFIGENIS OFIMUS, Thomas
1916. Journ. Bombay Nat. Hist, Soc. XXIV, p. 237.
Hkamti, Upper Chindwin, Uurma.
24. URICMOMYS RLFIGKMS I'VRRIIOMKRLS, Thomas
1895. Ann. Mag. Nat. Hist. 6, XVI, p. 472.
Ichang, China.
25. DRKMOMYS RUFIGENIS RIUDONENSIS, .Allen
iyo6. Bull. Amer. Mus. Nat. Hist. XXII, p. 472.
Riudon, Hainan.
26. DRliMOMV.S RUFIGENI.S GUL.ARIS, asgood
1932. Field Mus. Nat. Hist. Publ. Zool. Scr. XVIII, p. 284.
Mt. Fan Si Pan, near Chapa, Tongking.
27. DREMOMYS RUFIGICNIS I..\OM.JiCHE, Thomas
1921. Ann. Mag. Nat. Hist. 9, VII, p. 182.
Ban Hoi Mak, near Pak Hin Bun. Mekong River, Laos.
28. DRE.MOMYS RUFIGENIS LENTUS, Howell
1927. Journ. Washington Acad. Nat. Sci., XVII, p. 80.
Wenchaunshein, Szechuan, China.
incertae sedis
29. DRE.M(JMYS .MELLI. Matschie
1922. Beitr. Faun. Sinica, 88, 10, p. 23.
Mountains east of Shiuchow, Kwantung Province, China.
Genus II. R.\TUFA, Gray
1867. Ratufa, Gray, Ann. Mag. Nat. Hist. 3, XX, p. 273.
1880. EosciURUS, Trouessart, Le Naturaliste, ii, no. 37, p. 291. {Sciunis bicolor,
Sparrmann.)
1867. Ri;k.ma, Gray, .Ann. Mag. Nat. Hist. 3, XX, pp. 275-276. (Sciurus macrouriis.
Pennant.)
Type Species. — Sciurus indicus, Erxleben.
Range. — Indo-Malayan ; Ceylon, Southern India (Malabar, Coorg, Mysore);
Bombay, Sural, Central Provinces, Orissa; Nepal; Bengal, Assam,
Burma, Tenasserim; West Yunnan; Hainan; Annam, Siam, Malay Peninsula
and small adjacent islands ; Sumatra, Java, Borneo ; Banka, Billiton, Bali, Xatunas.
Number of Forms. — About seventy-two.
Ch.\racters. — Very large arboreal Squirrels with hea\y broad skull, very-
prominent postorbital processes, brachyodont cheekteeth in
which the cusps are low and the pattern as a rule not clear, and feet considerably
specialized for arboreal life.
Skull with somewhat depressed frontals, and large heavy postorbital process
which stands out noticeably from the skull in all species of the genus. Braincase
smooth, the hinder portion depressed downwards posteriorly. Rostrum short
and broad; frontals very broad. The parietal ridges evidently show no sign of
coming together. Zygomatic plate broad, moderately ridged on its upper border
and slanting gradually upwards, as in Sciurus. Infraorbital foramen normal,
384 RATUFA
torming canal, and with niassctcr knoli present. Mandible normal, angidar
process not much inflected. Bullae relatively large; palate broad, normal.
Incisors without special peculiarities.
Cheekteeth ^. Originallv there is evidently a pattern characteristic of the
family in the upper series, but the cusps are always extremely low, and the
pattern is usually obscured by many sTiiall depressions and pits, and appears
always less definite than in Sciiinis. The lower cheekteeth with the central
depression normally moderately well marked, but the cusps much flatter than
in Sciiiriis and allies, even the anterointernal cusp normally being only very
slightly raised above the general level of the tooth, and sometimes wearing down
altogether. A short re-entrant fold between the two outer main cusps normally
traceable.
Size large, usually over 250 mm. or even over 300 mm. up to 470 head and
bodv length, or perhaps more. Tail long, thickly bushy, rarely a little shorter
than head and bodv, often much longer. Forefoot extremelv broad, and rather
reminiscent in some ways of that of the Erethizontidae; D.4 longer than D.3;
D.5 and D.2 shorter, subequal; the inner pad is very much expanded and
probably takes the place of the pollex and is used for gripping. Hindfoot
broad, with well-developed hallux and normal arrangement of the digits, D.4
being the longest. Claws thick, powerful. The plantar and palmar pads, which
are evidently considerably specialized, have been described by Pocock, Proc.
Zool. Soc. London, 1922, p. 11S5.
As noticed under the genus Fiiiuiiiihii/iis there is some variation in the shape
of the baculum in this genus.
Forms seen: affiiiis, aiireizriitcr, hii/ieiisis, btinimeiisis, benaci/ensis, bicolor,
biiiigiiraiit'Hsis, can'iiiuiiensis, celaciiopepla, centralis, ceylonica, coiidiireiisis, con-
spictia, cotlnin:ata, dtindolciia, dealbiita, decolorata, ephippium, jelli, fretensis,
gigaiitea, hainana, indica, insignisjohoreiisis, laoiata, Icucogenvs, httrina, macroura,
macntioides, marana, masae, maxima, iiielaiioclirci, mc/anopepla, naiiogigas,
palliata, penangensis, pliaeopcphi, pinioisis, pvrsonuta, sandakanensis, sinliala,
sinus, sirbassensis, smithi, stigmusa, siiperans, tcnncnti, tiuimineiisis.
The classification of Robinson & Kloss, 1918, Rec. Indian Mus. X\', pt. IV,
pp. 171-250, Nominal List of Oriental Sciuridae, is accepted.
The genus apparently does not divide clearly into groups; it may be men-
tioned that the ear is heavily tufted in indica, all races of which except dealhata are
coloured red so far as seen, and which appears distinct from most of the other
forms; the ear also is tufted in gigantea, and more or less so in macroura.
List of Named Forms
I. RATUFA MACROURA MACROURA, Pennant
1769. Ind. Zool. I, PI. I.
Highlands of Ceylon.
Synonym: ceyloinciis, Erxleben, 1777, Syst. Regn. An. p. 416. Ceylon.
leiiiiinti, Blyth, 1S51, Journ. Asiat. Soc. Bengal, XX, p. 165.
Mountains, Ceylon.
zeylauicus, Ray, 1693, Syn. Qiiadr. p. 215 {fide Trouessart).
RATUFA 38s
2. RATUFA MACROURA MELANOCHRA, Thomas & Wroughton
1915. Joum. Bombay Nat. Hist. Soc, XXIV, p. 36.
Kottawa, Southern Ceylon.
3. RATUKA MACROURA ALBU'KS, Blyth
1859. Joum. Asiat. Soc. Bengal, XXVIII, p. 287.
Locality unknown.
4. R.ATUI-A MACROURA DANDOLENA, Thomas & Wroughton
1915. Joum. Bombay Nat. Hist. Soc, XXIV, p. 36.
W'ellawaya, Uva, Lowland Ceylon. (This race is also known from
S. India.)
5. RATUFA MACROURA SINHALA, Phillips
1931. Ceylon Journ. Sci. Sec. B, XVI, p. 215.
Nikawewa, near Kantalai, Eastern Province, Ceylon.
6. RATUFA INDICA INDICA, Er.\k-ben
1777. Syst. Regn. An. p. 420.
Bombay Presidency.
Synonym: purpureiis, Zimmermann, 1777, Spec. Zool. Geogr. Quad,
p. 518. Bombay.
(?) elphinstonei, Sykes, 1831, Proc. Zool. Soc. London,
p. 103. Deccan.
(?) malabarica, Schinz, 1845, Syn. Mamm. 1 1, p. 32. Malabar.
7. RATUFA INDICA SUPERANS, Ryky
1913. Joum. Bombay Nat. Hist. Soc, XXII, p. 436.
Wotekolli, South Coorg, India.
8. RATUFA INDICA BENGALENSIS, Blanford
1897. Joum. Bombay Nat. Hist. Soc, XI, p. 303, PI. B, fig. 2.
Locality not precisely specified.
9. R.\TUFA INDICA CENTR.-\L1S, Ryley
1913. Joum. Bombay Nat. Hist. Soc, XXII, p. 436.
Hoshangabad, Central Provinces, India.
10. RATUFA INDICA M.AXIMA, Schreber
1784. Saugth. IV, p. 7S4, PI. CCXXII, B.
Malabar, India.
11. RATUFA INDICA DEALB.^TA, Blanford
1897. Joum. Bombay Nat. Hist. Soc, XI, p. 299, PI. .A, fig. i.
Sural Dangs, India.
12. RATUF.A BICOLOR BICOLOR, Sparrmann
1778. Gotheb. Wet. Seversk. Handl. i, p. 70.
Anjer, West Java.
Synonym: major. Miller, 191 1, Proc. Biol. Soc. Washington, XXIV,
p. 28. Anjer, West Java.
(?) albiceps, Desmarest, 1817, Nouv. Diet. Hist. Nat.. X,
p. 105. Java.
javensis, Zimmermann, Geog. Ges., II, 1780, p. 342.
0) leschenaulti, Desmarest, Mamm., 1820, p. 335.
(?) humeralis, Coulon, Mem. Soc. Sci. Nat. Neuchatel, 1835,
I, p. 122.
13. R.^TUFA BICOLOR BALIENSI.S. Thomas
1913. .Ann. Mag. Nat. Hist. 8, XI. p. 506.
Tjetoekambawanp, Bali.
25 — Livirij,' Rodents — I
386 RATUFA
14. RATUFA BICOt.OR PALLIATA, Miller
igo2. Proc. Acad. Nat. Sci. Philadelphia, LIV, p. 147.
Indragiri River, E. Sumatra.
15. RATUFA BICOI.OR LAENATA, Miller
1903. Proc. U.S. Nat. Mus., XXVI, p. 449.
Pulau Tuangku, Banjak Islands, W. Sumatra.
16. RATUFA BICOLOR BATUANA, Lyon
1916. Proc. U.S. Nat. Mus., LII, p. 445.
Tana Bala, Batu Islands, W. Sumatra.
17. RATUF.^ BICCILOR SMITHI, Robinson & Kloss
1922. Ann. Mag. Nat. Hist, q, IX, p. 89.
Langbian Peaks, South Annani.
18. RATUFA NOTABILIS NOTABILIS, Miller
1902. Proc. Acad. Nat. Sci. Philadelphia, p. 150.
Lingga Island, Rhio-Lingga Archipelago.
19. R,\TUFA NOTABILIS INSIGNIS, Miller
1903. Smiths. Misc. Coll., XLV, p. 4.
Pulau Sugi, Rhio-Lingga Archipelago.
20. RATUFA NOTABILIS BUL.\NA, Lyon
1909. Proc. U.S. Nat. Mus., XXXVI, p. 482.
Pulau Bulan, Rhio-Lingga Archipelago.
21. RATUFA NOTABILIS CARIMONENSIS, Miller
1906. Proc. U.S. Nat. Mus., XXXI, p. 257.
Great Karimon Island, Rhio-Lingga Archipelago.
22. RATUFA NOTABILIS CONDURENSIS, Miller
1906. Proc. U.S. Nat. Mus., XXXI, p. 258.
Pulau Kundur, Rhio-Lingga Archipelago.
23. R.i^TUFA NOTABILIS CONFINIS, M.ller
1906. Proc. U.S. Nat. Mus., XXXI, p. 259.
Sinkcp Island, Rhio-Lingga Archipelago.
24. RATUFA NOTABILIS CONSPICUA, Miller
1903. Smiths. Misc. Coll., XLV, p. 5.
Pulau Bintang, Rhio-Lingga Archipelago.
25. RATUFA EPHIPPIUM EPHIPPIUM, Mulkr
1838. Tijds. Nat. Gesch. Physiol., V, p. 147.
S.-F,. Borneo, low country.
26. R.^TUFA EPHIPPIUM COTHURNATA, Lyon
1911. Proc. U.S. Nat. Mus., XL, p. 93.
Mount Palung, near Sukadana, W. Borneo.
27. RATUFA EPHIPPIUM BARAMENSIS, Bonhote
1900. Ann. Mag. Nat. Hist. 7, V, p. 496.
Baram district, Sarawak, Borneo.
28. RATUFA l-PHIPPIUM SANDAKANENSIS, Bonhote
1900. Ann. Mag. Nat. Hist. 7, V, p. 497.
Sandakan, British N. Borneo.
RATUFA 387
29. RATUFA EPHIPPIUM GRISKICOLLIS, Lyon
igri. Proc. U.S. Nat. Mus., XL, p. 94.
Panebangen Island, W. Borneo.
30. R-VrUKA EPHIPPILM VIT'IWTA, Lyon
191 1. Proc. U.S. Nat. Mus., XL, p. 94.
Pulau Laut, S.-E. Borneo.
31. RATUFA EPHIPPIUM VITTATULA, Lyon
191 1. Proc. U.S. Nat. Mus., XL, p. 95.
Pulau Sebuku, S.-E. Borneo.
32. RATUFA EPHIPPIUM BUNGURANENSIS, Thomas & Hartert
1894. Nov. Zool. I, p. 658.
Bunguran Island, Natunas.
33. RATUFA EPHIPPIUM SIRHASSENENSIS, Bonhote
1900. .'^nn. Mag. Nat. Hist. 7, V, p. 498.
Sirhassen Island, Natunas.
34. R,ATUFA EPHIPPIUM N.\NOGIGAS, Thomas & Hartert
1895. Nov. Zool., II, p. 491.
Pulau Laut, N. Natunas.
35. RATUFA EPHIPPIUM POLIA, Lyon
1906. Proc. U.S. Nat. Mus., XXXI, p. 585.
Billiton Island, between Sumatra and Borneo.
36. RATUFA EPHIPPIUM BANCANA, Lyon
1906. Proc. U.S. Nat. Mus., XXXI, p. 587.
Banka Island, off Sumatra.
37. RATUFA EPHIPPIUM LUMHOLZI, Ldnnberg
1925. Ann. Mag. Nat. Hist, g, XVI, p. 514.
Pipoh Boelengan, E. Central Borneo.
38. RATUFA EPHIPPIUM DULITENSIS, Lonnberg & Mjoberg
1925. .■Vnn. Mag. Nat. Hist. 9, XVI, p. 514.
Mount Dulit, Borneo.
39. R.VrUFA AFFINIS AFFIMS, Raffles
1822. Trans. Linn. Soc, XIII, p. 258.
Singapore.
40. R.ATUFA AFFIMS HYPOLEUCA, Horsfield
1824. Zool. Res. in Java, p. 165.
Bencoolen, Sumatra.
41. R.\TUFA AFFINIS C.ATEiLANA, Lyon
1907. Proc. U.S. Nat. Mus., XXXII, p. 443.
Kateman River, S.-E. Sumatra.
42. R.-\TUFA AFFINIS JOHORENSIS, Robinson & Kloss
191 1. Joum. Fed. Malay States Mus., IV, p. 244.
Padang Tuan, Segamat, N.-W. johore.
43. R.\TUFA AFFINIS AURIVENTER, Is. Geoffroy
1832. Mag. Zool. CI. I, pi. V.
"Java" (in error): substitute Malacca.
44- R.-\TUFA AFFINIS .^RUSINUS, Lyon
1907. Proc. U.S. Nat. .Mus., XXXII, p. 442.
.Aru Bay, N.-E. Sumatra.
388 RATUFA
45. RATUKA AFFINIS PVRSUNUTA, Miller
1900. Proc. Washington Acad. Sci., 11, p. 75.
Trang, Siamese Malaya.
46. RATUFA AFFINLS FEMORALIS, Milkr
1903. Proc. U.S. Nat. Mus., XXVI, p. 447.
Pulau Tuangku, Banjak Islands, off W. Sumatra.
47. RATUF.'^ AFFINIS NIGRI-.SCFNS. Miller
1903. Proc. U.S. Nat. Mus., XXVI, p. 44S.
Pulau Mansalar, near Tapanuli Bay, W. Sumatra.
48. RATUFA AFFINLS BALAE, Miller
1903. Smiths. Misc. Coll., XLV, p. 7.
Tana Bala, Batu Islands, W. .Sumatra.
49. RATUF.-X AFFINIS MASAE. Miller
1903. Smiths. Misc. Coll., XLV, p. 8.
Tana Masa, Batu Islands, W. Sumatra.
50. R.ATUFA AFFINIS PINIENSIS, Miller
1903. Smiths. Misc. Coll., XLV, p. 7.
Pulau Pinie, Batu Islands, W. Sumatra.
51. RATUFA AFFINIS BANGUEVI, Chasen & KIoss
1932. Bull. Raffles Mus. 6, p, 22.
Banguey Island, N. Borneo.
52. RATUFA AFFINIS INTERPOSITA, Kloss
1933. Bull. Raffles Mus. 7, p. 2.
Selangor, Malaya.
53. RATUFA AFFINIS FRONTALIS, Kloss
1933. Bull. Raffles Mus. 7, p. 2.
Perak, Malaya.
54. RATUFA GIGANTEA GIGANTEA, Macclelland
1839. Proc. Zool. Soc. London, p. 150.
Assam.
55. RATUFA GIGANTEA LUTRINA, Thomas & Wroughton
1916. Journ. Bombay Nat. Hist. Soc, XXIV, p. 226.
West bank of Upper Chindwin, Upper Burma.
56. RATUFA GIGANTEA MACRUROIDES, Hodgson
1849. Journ. Asiat. Soc. Bengal, XVIII, p. 775.
Bengal.
57. RATUFA GIGANTEA FELLI, Thomas S: Wroughton
1916. Journ. Bombay Nat. Hist. Soc, XXIV, p. 226.
Yin, Lower Chindwin, Burma.
58. R.A.TUFA GIGANTEA HAINANA, Allen
1906. Bull. Amer. Mus. Nat. Hist., XXII, p. 472.
Cheteriang, Hainan.
59. RATUFA GIGANTEA STIGMOSA. Thom.is
1923. Journ. Bombay Nat. Hist. Soc, XXIX, i, p. 86.
Doi .Sritepe, Chiengmai, Siam.
RATUFA 389
60. RATUFA PHAKOPEPLA PHAEOPEPLA, Miller
1913. Smiths. Misc. Coll., LXI, no. 21, p. 25.
Sungei Balik, S. Tenasserim.
(Robinson & Kloss evidently consider this species doubtfully dis-
tinguishable from melanopepla.)
61. RATUFA PHAICOPEPLA MARANA, Thomas & Wroughton
1916. Joum. Bombay Nat. Hist. Soc, XXIV, p. 227.
Mount Popa, Burma.
62. RATUFA PHAEOPEPLA LEUCOGENVS, Kloss
1916. Proc. Zool. Soc. London, p. 43.
Lem Ngop, S.-E. Siam.
63. RATUP'A PHAEOPEPLA SINUS, Kloss
1916. Proc. Zool. Soc. London, p. 44.
Koh Kut Island, S.-E. Siam.
64. RATUFA MELANOPEPLA MELANOPEPLA, Miller
1900. Proc. Washington Acad. Sci., II, p. 71.
Telibon Island, Trong, Siamese Malaya.
65. RATUFA MELANOPEPLA PENINSULAE, Miller
1913. Smiths. Misc. Coll., LXI, 21, p. 25.
Lay Song Hong, Trong, Siamese Malaya.
66. RATUFA MELANOPEPLA DECOLORATA, Robinson & Kloss
1914. Ann. Mag. Nat. Hist. 8, XIII, p. 227.
Koh Samui Island, Bandon Bight, Siamese Malaya.
67. RATUFA MELANOPEPLA CELAENOPEPLA, Miller
19:3. Smiths. Misc. Coll., LXI, no. 21, p. 26.
Domel Island, Mergui Archipelago.
68. RATUFA MELANOPEPLA FRETENSIS, Thomas & Wroughton
1909. Ann. Mag. Nat. Hist. 8, IV, p. 535.
Pulau Langkawi, Malaya.
69. R..\TUFA MEL.\NOPEPLA PENANGENSIS, Robinson & Kloss
191 1. Joum. Fed. Malay States Mus., IV, p. 242.
Telok Bahang, Penang Island.
70. RATUFA MEL.\NOPEPLA TIOMANENSIS, .Miller
1900. Proc. Washington .^cad. Sci., II, p. 216.
Pulau Tioman, E. coast Malay Peninsula.
71. R.'^TUFA MELANOPEPLA AN.\MB.\E, Miller
1900. Proc. Washington .Acad. Sci., II, p. 215.
Pulau Jiniaja, Anamba Islands.
72. R-ATUFA MELANOPEPLA .\NGU.STICEPS, Miller
1901. Proc. Washington .^cad. .Sci., Ill, p. 130.
Pulau Lingung, Natuna Islands.
Section C. Specialized Indo-M.\l.\y.\n Gener.a, all clearly distinct from
Sciurus and immediate allies. In this section, which is not a natural
group as regards relationships, but rather holds several very distinct
390 MENETES
offshoots from the more normal Sciiinis branch, I nickide Meiietes,
Laiisciis, Glyp/iolcs, Rhcithrosciurus, Rhinosciurus and llyosiiuriis.
Genus 13. MENETES, Thomas
1 90S. Menetes, Thomas, Journ. Bombay Nat. Hist. Soc, XVIII, no. 2, p. 244.
Type Species. — Sciurus bcidmorci, Blyth.
Range. — Northern portion of Malayan region; Siam, Annam, Camhodia,
Burma, Tenasserim.
Number of Forms. — Ten.
Characters. — Skull long and narrow, with markedly elongated rostrum;
postorbital process medium in size or relatively small. In-
fraorbital foramen and zygomatic plate rather variable; in some specimens the
part ot the zygomatic plate behind the intraorbital toranien is very narrow,
and the foramen is w'ell open ; in others, there is an approach to the condition
found in more normal genera. A well-developed masseter knob is present.
The parietal ridges may join. The cheekteeth approach the type of the African
Paraxenis, but the pattern tends to wear down very quickly. P. 3 present, strong.
The central depression in the upper main teeth remains in adult as a well-
marked re-entrant fold, but the anterior and posterior folds tend to wear out,
so that the tooth takes on a more or less horseshoe-like shape. Lower teeth
quickly wearing down to a two-lobed structure from a pattern originally like
that oi Piinixenis; the central depression becomes quickly reduced, and remain-
ing as a deep narrow pit, often isolated in the centre of the tooth in old age.
Posterointernal cusp and its adjoining ridge strong. The upper incisors rather
shortened; the lower ones long.
Tail shorter than head and body. Digits with the arrangement characteristic
of arboreal Squirrels. Black and white flank-stripes present, usually strong;
in 71/. b. decoidtus the pattern takes on a superficial resemblance to Tiuiiicis,
including a black mid-dorsal stripe. Mammae 6 (Thomas).
Forms seen : herdmorei, coiisuhiris, decoiatus, iiioeresceiis, inoidiotei, peninsularis ,
rufescens, umbrosus.
List of Named Forms
1. MENI;T1;S herdmorei BERDMOREI, Blyth
1849. Journ. Asiat. Soc. Bengal, XVIII, p. 603.
Thounffveen district. Lower Burma.
2. MENETI:S BERDMOREI AMOTUS, Miller
1913. Smiths. Misc. Coll., LXI, no. 21, p. 24.
Domel Island, Mergui Archipelago.
3. MHNETKS BERDMOREI KORATENSIS, Gyldenstolpe
1917. Kungl. Svenska. Vet. Akad. Handl., LVII, no. 2, p. 39.
Sakerat, near Korat, E. Siam.
4. MENETES BERDMOREI MOUHOTEI, Gray
1861. Proc. Zool. Soc. London, p. 137.
Cambodia.
Synonym: pyrrocephaliis, Milne-Edwards, 1867. Rex-. Mag. Zool. 2,
XIX, p. 225. Cochm China.
MENETES— LARISCUS 391
5. MENETES BERDMOUEI DECORA'IX'S, Thomas
1914. Journ. Bombay Nat. Hist. Soc, XXIII, p. 24.
Mount Popa, Burma.
6. MENETES BERDMOREI MOERESCENS, Thomas
1914. Journ. Bombay Nat. Hist. Soc, XXHI, p. 25.
Bali, near Nhatrang, Annam.
7. MENETES BERDMOREI CONSULARIS, Thomas
1914. Journ. Bombay Nat. Hist. Soc, XXHI, p. 24.
Nan, North Siam.
8. MENETES BERDMOREI UMBROSUS, KIoss
1916. Proc Zool. Soc. London, p. 49.
Koh Chang Island, S.-E. Siam.
9. MENETES BERDMOREI RUFESCENS. KIoss
1916. Proc. Zool. Soc. London, p. 50.
Koh Kut Island, S.-E. Siam.
10. MENETES BERDMOREI PENINSULARIS, Robinson & KIoss
1919. Joum. Nat. Hist. Soc. Siam, HI, no. 4, p. 375.
Ban Kok Klap, Nakon Sritamaret, Peninsular Siam.
Genus 13. I^ARISCUS, Thomas & Wroughton
1867. Lari.^, Gray, Ann. Mag. Nat. Hist. 3, XX, p. 276. (Not of Scopoli.)
1909. Lariscus, Thomas & Wroughton, Proc. Zool. Soc. London, p. 389.
Type Species.— 5c/«n« insignis, Cuvier.
Range. — Southern part of the Malay Peninsula, Sumatra, Java, Borneo,
and neighbouring small islands.
Number of Forms. — Fourteen.
Characters. — Cheekteeth hypsodont, almost completely simplified in
pattern. The lower series are roughly two-lobed, and even
in the very young the pattern is nearly lost. Thus, when cut, the central
depression is already narrow and much reduced. Upper cheekteeth with a
vague pattern reminiscent of Funisciunis sometimes traceable, but usually
completely simple; P. 3 present. A short sagittal ridge may be present in old
age. Rostrum tending to be long. Bullae rather small as a rule. Zygomatic
plate near the Sciunis type; infraorbital foramen normal.
Tail relatively short, usually about sixty per cent of length of head and body.
Digits as in normal arboreal genera. The type and allies have three black stripes
present; hosei is rather more brightly coloured, and has four black stripes.
Mammae, type species 6 (Thomas).
Forsyth ^lajor in 1893 (Proc. Zool. Soc. London, p. 1S5) wrote fully on the
dentition of this genus. He states that the simplification of the teeth is probably
due to the food, comparing these Squirrels with certain Bats, which feed on
juicy fruits whose contents need not be chewed, and differ in a similar manner
from their allies {" Macroglossi, Pteropus scciptilatiis, Epomophori, compared with
other Fteropi"). He remarks that the species oi Lariscus are Ground-squirrels.
392 LARISCUS
Forms seen: divcrsiis, foriiicatus, hosei, insi»nis, jalorensis, javamis, mendion-
alis, niobe, siberii, vulcanis.
Robinson & Kloss, 1918, considered all forms other than hosei as races of
insignis, though niobe is sometimes considered as a distinct species.
List of Named Forms
insignis Group
1. LARISCUS INSIGNIS INSIGNIS, F. Cuvier
1 82 1. Hist. Nat. Marnrn. (ii) 34, pi. 233.
Sumatra.
2. L.^RISCUS INSIGNIS JALORENSIS, Bonhote
1903. Fascic. Malay. Zool. i, p. 25.
Jalor, N. Malay Peninsula.
Synonym: peninsidae, Miller, 1903, Smiths. Misc. Coll., XLV, p. 25.
Khow Sai Dow, Trong, Siamese Malaya.
3. LARISCUS INSIGNIS MERIDIONALIS, Robinson & Klois
1911. Joum. Fed. Malay States Mus., IV, p. 172.
Changi, Singapore Island.
4. L.^RISCUS INSIGNIS FORNIC.ATUS. Robinson
1917. Journ. Fed. Malay States Mus., VII, p. 102.
Tioman Island, E. coast Malay Peninsula.
5. L.ARISCUS INSIGNIS DIVERSUS, Thomas
1898. Ann. Mag. Nat. Hist. 7, II, p. 248.
Baram district, Borneo.
6. LARISCUS INSIGNIS CASTANEUS, Miller
1900. Proc. Acad. Sci. Washington, II, p. 217.
Pulau Siantau, Anamba Islands.
7. LARISCUS INSIGNIS S.^TURATUS, Chasen
1934. Bull. Raffles. Mus. 9, p. 99.
Rhio Archipelago, Malaya; Bintang Island.
S. L.ARISCUS INSIGNIS NIOBE, Thomas
1898. Ann. Mag. Nat. Hist. 7, II, p. 249.
Pajo, highlands of W. Sumatra.
g. L.ARISCUS INSIGNIS SIBERU, Chasen & Kloss
1928. Proc. Zool. Soc. London, p. 827.
Siherut, Mentawei Islands, W. Sumatra.
10. L.ARISCUS INSIGNIS VULCANUS, Kloss
1921. Joum. Fed. Malay States, Mus., X, p. 233.
Ongop Ongop, Idjen Massif, 5,700 ft., Eesoeki, E. Java.
11. LARISCUS INSIGNIS JAVANUS, Thomas & Wroughton
1909. Proc. Zool. Soc. London, Abstr., p. 19, id. torn, cit., p. 389.
Buitenzorg, W. Java.
12. LARISCUS INSIGNIS OBSCURUS, Miller
1903. Smiths. Misc. Coll., XLV, p. 23, pi. i, fig. 2.
South Pagi Island, W. Sumatra.
LARISCUS— GLYPHOTES— RHEITHROSCIURUS 393
13. LARISCUS INSIGNIS ROSTRATUS, Miller
1903. Smiths. Misc. Coll., XLV, p. 24.
Tana Bala, Batu Islands, W. Sumatra.
hosei Group
14. LARISCUS HOSEI, Thomas
1892. .^nn. Mag. Nat. Hist. 6, XX, p. 215, 216.
Mount Dulit, Baram district, Borneo.
Genus 14. GLYPIIOIES, Thomas
1898. Glyphotes, Thomas, Ann. Mag. Xat. Hist. 7, II, p. 251.
Type Species. — Glyphotes simus, Thomas.
Range. — Borneo.
Number of Forms. — One.
Characters. — A small Squirrel with peculiar and specialized incisors.
Muzzle short and broad; nasals short. Postorbital process
small. Zygomatic plate slanting upwards somewhat vertically (though not
comparable to Xatmusciiirus section); infraorbital foramen rather well open, the
part of the zygomatic plate behind it rather reduced. Mandible weak, w4th
coronoid process very low, and condylar process slender. Upper incisors very
broad, but not thickened anteroposteriorly ; their lower tips tending to cur\'e
away from each other. Lower incisors similar, but their upper portions more
strongly divergent from each other. Cheekteeth |, the pattern evidently not
abnormal; P. 3 small.
Externallv with no special features; tail rather narrow, relatively long;
flank-stripes present (white over black).
Forms seen : simus. (Only the type skull and skin.)
List of Named Forms
1. CJLYPHOTES SIMUS, Thomas
1898. .A.nn. Mag. Nat. Hist. 7, II, p. 251.
Kina Balu, N. Borneo.
Genus 15. RHEITHROSCIURUS, Gray
1867. RHEITHROSCIURUS, Gray, .Ann. Mag. Nat. Hist. 3, XX, p. 272.
Type Species. — Sciunis macrotis. Gray.
Range. — Borneo.
Number of Forms. — One.
Characters. — Skull and teeth abnormal, the incisors greatly thickened from
before backwards, the toothrow much reduced, in these
characters paralleling the .African .-Vnomaluroid genera Zenkerella and Idiuriis.
Incisors with a strong subapical notch present. Their anterior faces clearly
Fig. 97. Rheithrosciurus macrotis, Gray.
B.M. No. 27.8.6.5, <J; X I.
Fig. 98. Rheithrosciurus macrotis, Gray.
B.M. No. 27-8.6. 5, o; X I.
RHEITHROSCIURUS— RHINOSCIURUS 395
marked with very many narrow parallel grooves, a structure traceable also in
I'limius and Marmota, though in these genera not so well marked. The incisors
are much compressed. Toothrow considerably reduced, the premolars of both
upper and lower series smaller than the molars. All teeth brachyodont, basin-
shaped, nearly simplified in pattern; those of the upper series near Lariscus,
and with traces of pattern nearly obliterated in all skulls seen; the lower teeth
square, with a more or less well-marked small cusp at each corner.
The hinder part of the mandible is more rounded than usual, the angular
process higher, well ridged below, the coronoid not reduced, but not higher than
the condylar, the bone running from one to the other nearly straight. Palate
depressed betw'een toothrows, ending abruptly just behind them, the posterior
termination forming three sides of a square with the hamular processes. Ros-
trum much elongated; zygomatic plate slanting abruptly upwards, flat, not
heavily ridged, placed rather far back; but orbit as in normal Sciurinae. Infra-
orbital foramen forming a long canal, its anterior opening far in front of P. 4.
Frontals broad; postorbital processes moderately well developed.
Size large; about 23 inches head and bodv length (or more?); ear large, with
extremely enlarged ear-tufts. Tail excessivelv thick and bushy, relatively long.
Fur crisp and harsh. Digits of hindfoot as in normal Tree-squirrels. JManus
with normally D.4 longer than D.3, though evidently there may be some
variation in this character.
Mr. W. Frost told me that these animals are Ground-squirrels.
Remarks. — One of the most specialized and distinct genera in the group.
Forms seen : macrotis.
List of Named Forms
1. RHEITHROSCIURUS MACROTIS, Gray
1856. Proc. Zool. Soc. London, p. 341, pi. XLVI.
Sarawak, Borneo.
Genus 16. RHINOSCIURUS, Gray
1843. Rhinoscr'RUS, Gray, List. Mamm.. p. 195. (According to Tate, 1935, .^rner.
Mus. Nov., 807, R. tupaioides. Gray, is nom. nud. and the name Rliinosciurus should
date from Blyth, 1855, with type 5. laticaudatus, Miiller & Schlegel.)
Type Species. — Rhinosciurus tupaioides. Gray.
Range. — Southern portions of the Malay Peninsula, Sumatra, Borneo, and
adjacent islands.
Number of Forms. — Seven.
Characters. — Skull highly abnormal, with immensely elongated rostrum;
upper incisors much reduced, very narrow, nearly vestigial;
upper cheekteeth rapidly wearing down and simplifying. The rostrum in its
length is quite unique in the family, except Hyosciurus, in which it is even
longer. Postorbital process short. Bullae considerably enlarged. Zygomatic
396 RHINOSCIURUS
plate with its upper border ridged but short, and the portion behind the infra-
orbital foramen, which is rather well open, much narrowed. Palatal foramina
far in front of toothrows.
Upper incisors appearing hardly functional; lower incisors long, not much
reduced. Upper cheekteeth originally with rather complicated pattern, cusps
and ridges well marked, but the pattern quickly wearing down, the main de-
pression remaining at first as a wide outer fold, later simplifying and wearing
away altogether. Lower cheekteeth with the same elements as the upper series
as regards the change of pattern brought about by wear; posterointernal cusp
originally well marked; the central depression seems in this genus to take the
form of an outer re-entrant fold rather than an inner one, differing from other
genera in this respect. P. 3 large, well developed.
Hindfoot narrow, but arrangement of digits as usual, D.4 slightly longer
than D.3. Forefoot with arrangement of digits as usual. Tail bushy, consider-
ably shorter than head and body length. Of the habits of this genus Robinson
& Kloss write : "They are strictly terrestrial and very shy . . . their diet, judging
from numerous specimens examined, is principally insectivorous, consisting of
large ants and beetles. The tongue is very long, and remarkably protrusile,
and it is probable that gritty matter taken up with the insects by means of this
organ accounts for the rapid wear of the teeth."
The genus appears to show certain resemblances in the infraorbital foramen,
original pattern of upper teeth, and tendency to elongation of rostrum, to
Menetes, which genus is, however, much less specialized.
But I think in this genus we have probably a parallel in evolution to the
remarkable Murine genus Rhvnchoinys. In Rhvncliomys there is the same elonga-
tion of muzzle, while the upper incisors are even more reduced; but the cheek-
teeth in this case have become so reduced as to be nearly invisible. Thomas
thought that this Rat was insectivorous; it appears that Rhinosciurus in many
respects is going well on the same road, in fact had some teeth, for instance the
premolars, become suppressed, the parallel between these two unrelated genera
would be nearly complete.
Forms seen : leo, laticaudtitiis, tiipaioides, robiiisoni, rhionis.
List of Named Forms
1. RHINOSCIURUS LATICAUDATUS LATICAUDATUS, Muller & Schlegel
1839. Vcrhandl. Nat. Ciesch. p. 100. pi. XV, figs. I, II, III.
Pontianak, Borneo.
2. RHINOSCIURUS LATICAUDATUS SATURATUS, Robinson & Kloss
1919. Joum. Fed. Malay States Mas., VII, p. 274.
Barisan Range, W. Sumatra.
3. RHINOSCIURUS LATICAUD.ATUS TUPAIOIDES. Gray
1843. List. Mamm., p. 195.
Singapore.
Synonvm: peracer, Thomas & Wroughton, 1909, .Ann. Mag. Nat. Hist.
8, III, p. 440. Perak.
Fig. 99. Rhinosciurus laticaudatus tupaioides, Gray.
B.M. No. 9.4.1.228; X 2.
Fig. 100. Rhinosciirls laticaudatus tupaioides, Gray.
B.M. No. 9.4.1.228; x 2.
398 RHINOSCIURUS— HYOSCIURUS
4. RHINOSCIURUS LATICAUDATUS LEO, Thomas & Wroughton
1909. Ann. Mag. Nat. Hist. 8, III, p. 440.
Changi, Singapore Island.
5. RHINOSCIURUS LATICAUD.ATUS RHIONIS, Thomas & Wroughton
1909. Ann. Mag. Nat. Hist. S, III, p. 441.
Karimon, Rhio-Lingga Archipelago.
6. RHINOSCIURUS L,ATICAUD.ATUS ROBINSONI, Thomas
190S. Journ. Fed. Malay States Mus., II, p. 104.
Tioman Island, E. Malay Peninsula.
7. RHINOSCIURUS LATICAUDATUS INCULTUS, Lyon
1916. Proc. U.S. Nat. Mus., LII, p. 444.
Pulau Tuanku, Banjak Islands, W. Sumatra.
Genus 17. HYOSCIURUS, Tate & Archbold
1935. HYOSCIURUS, Tate & Archbold, Amer. Mus. Nov. 807, p. 2.
Type Species. — Hyosciurus heinrichi, Tate & Archbold.
R.\NGE. — Celebes.
Number oe Forms. — Two.
This genus was not represented in the British Museum when this work was
originally written, but a fine series of a dozen has been obtained in 1939 by
Mr. Frost from the Molengraff Range, Mid Celebes.
Char.'\cters. — Skull \\ ith extremely elongate rostrum, if anything more so
than in Rhinosciurus. Postorbital process small. In the
adult, the temporal ridges fuse to form a short but strong sagittal ridge. Infra-
orbital foramen forming a long canal. Zygomatic plate slanting upwards
anteriorly, rather flat, and much less projecting forwards than in Rhinosciurus.
Nasals projecting anteriorly far forwards over the incisors. The upper incisors
are not reduced, but are thick and strong (compare Rhinosciurus). Bullae
medium-small. Palatal foramina far in front of toothrows; palate normal.
Mandible stronger than in Rhinosciurus, the angular portion inflected to a
degree (not extremely so, for instance, not comparing with that of Cynomys).
Lower incisors robust, not extremely long. Cheekteeth J; molars quite
normal, Sciurine in pattern, witliout the peculiarities of Rhinosciurus. M.3 is
rather small. In the oldest specimen seen, a male with pattern of teeth ob-
literated and a strong sagittal ridge to the skull, there is not the slightest sign
of the extreme deterioration in the molars which takes place in Rhinosciurus
with wear.
Tail very short. Claws enormous, particularly those of the forefoot, but
even so, less strongly enlarged than in Sperinuphilopsis.
Forms seen: heinrichi.
List of Named Forms
I. HYOSCIURUS HEINRICHI HEINRICHI, Tate & Archbold
1935. .^mer. Mus. Nov. Soi, p. 2.
Latimodjong Mountains, S. Celebes.
HYOSCIURUS— HELIOSCIURUS 399
2. HYOSCIURUS HKIMUCHI ILKILE, Tate & Archbold
1936. Amer. Mus. Nox. 846, p. i,
Ile-ile, N. Celebes; 1700 m.
Mr. Frost states that it is a burrowing form, living underground, or more
or less, and that the natives know it as a species of Rat.
Section D. African Arboreal Genera. All except Heliosciurus are clearly
distinguishable from Sciurus on dental characters, and there is a tendency-
present for the lower molars to lose the central depression characteristic
of normal Squirrels, and for these teeth to become transversely ridged,
as in the upper series. In Protoxerus and allies the infraorbital foramen
is normally unusually large, and forms no canal.
Genus 18. HELIOSCIURUS, Trouessart
1880. Heliosciurus, Trouessart, le Naturaliste, II, no. 37, p. 292.
IQ16. Aethosciurus, Thomas, Ann. Mag. Nat. Hist. 8, XVII, p. 271. Sciurus
poensis, Smith. Valid as a subgenus.
Type Species. — Sciurus gambianus, Ogilby.
Range. — Africa: Sudan, Abyssinia, Kenya, Uganda, Tanganyika; Senegal,
Gambia, Sierra Leone, Liberia, Ivory Coast, Gold Coast, Nigeria,
Cameroons, Fernando Po, Congo, Angola, Rhodesia, Nyasaland, Mozambique.
Number of Forms. — About fifty-two.
Remarks. — The genus was originally given generic rank by Thomas in 1909
on the single character that, compared with Sciurus, P.3 is
absent. This is not a valid character. To the genus Sciurus in the same paper
were referred three African Squirrels, S. puensis, S. lucifer, and 5. ruwenzori.
In 1916 Thomas referred these to a new genus Aethosciurus, on the grounds that
the baculum differed from Sciurus (vulgaris), though only poensis had been
e.xamined.
Hollistcr, 1919 (U.S. Nat. Mus. Bull. 99, p. 9), pointed out that the teeth of
Heliosciurus and Aethosciurus agreed with each other and differed from those of
Sciurus vulgaris in certain details, and treated the two groups as a single genus,
remarking that it was not wise to give Aethosciurus generic rank simply on account
of the small extra premolar, which was also known to be present occasionally
in typical Heliosciurus.
But the dental characters pointed out by Hollister, while constant, and separate
from Sciurus vulgaris, agree with certain Asiatic forms, such as Callosciurus.
There are then only two characters which seem to me to separate the present
genus clearly from Callosciurus. First the baculum, which according to Pocock
is absent in Heliosciurus (apparently a very rare feature in the Order, but known
elsewhere in the .American Tamiasciurus in the present familv), and minute in
Aethosciurus (poensis). But it must he admitted that only punctatus, rufohrachium,
and poensis were examined or mentioned in Pocock's paper; so that it mav be
that this character will later be found to be invalid ; though it must be stated that
400 HELIOSCIURUS
all typical Heliosciurus are apparently so closely related that Ingoldby has
suggested that all forms should he referred to one species only.
Secondly, the zygomatic plate, which seems to me to be constantly more
strongly ridged, and with an extremely powerful masseter knob present, in
typical Heliosciurus and in Aethosciurus ruwenzorii. The other species referred
to Aethosciurus agree with Faraxerus and Funisciurus in the curiously shortened
zygomatic plate, with the ridge stopping abruptly over the infraorbital foramen,
and not approaching the superior border of rostrum. Further, ruwensorii has
a vestigial P. 3, while this tooth is quite well developed and relatively large in all
other Aethosciurus seen (as in Paraxerus). The baculum is minute also in
Funisciurus and Paraxerus, or those of this genus which Pocock examined.
Apart from poensis and ruwenzorii, I very much suspect that the other species
named for Aethosciurus. namely hicifer, vexillaris, will be found when a repre-
sentative collection comes to hand, to belong to Paraxerus. The lower cheek-
teeth, which afford the only character by which Aethosciurus mav be separated
from Paraxerus, appear to me to be very suggestive of Paraxerus in all skulls
examined, but all seen are much too worn for me to be able to say. "Aetho-
sciurus" bvatti is definitely based on a Paraxerus, so far as the tvpe skull shows,
and is here transferred to that genus, ^ut poensis, though agreeing in zygomatic
plate formation with Funisciurus and Paraxerus, has definitely the unspecialized
teeth found in Heliosciurus and must remain in this genus; and the same remarks
apply to ruzcenzorii, which is probably a primitive Heliosciurus s.s. in which the
minute premolar has not yet become suppressed.
Characters. — Skull often with parietal ridges, which may tend to join. Post-
orbital process usually rather well developed. Bullae of
moderate size. Palate normal. Infraorbital foramen usually rather well open,
the upper part of the zygomatic plate prominently ridged, approaching the
Xerus type though less extreme, slanting upwards far forwards, and there is a
very strong masseter knob present. The upper incisors are in rare individual
cases with a faint groove traceable. Upper cheekteeth like Sciurus, but with the
main ridges strongly convergent internally, particularly the third ( = the second
principal) ridge, which runs almost from the outer corner to the middle of the
inner part of the tooth. The inner side of AI.i and .M.2 nearly square. The
anterior cusp of P. 4 is extremely well developed, projecting forwards, the
depression immediately behind it well marked. Lower cheekteeth with elements
not very different from Sciurus, but with a narrow transverse valley extending
from first outer main cusp to the anterointernal cusp; this structure is present
in some of the Indo-I\Ialavan Squirrels. It evidently marks the beginning of the
type of tooth found in Paraxerus and Funisciurus . The central depression is
not obliterated, and is usually present as an important feature of the teeth.
External characters as in normal Tree-squirrels. Tail long. Back not striped.
The subgenus //f//o.'if«/nw contains the^f/w/)/!^;;;,? group only, all the members
of which are referred to a single species by Ingoldby (Proc. Zool. Soc. London,
1927, p. 471).
The subgenus Aethosciurus at the moment consists of three groups: ruiven-
zorii, P. 3 minute, zvgomatic plate more as in typical Heliosciurus; poensis, P. 3
HELIOSCIURUS 401
rather large; zygomatic plate as in Paraxerus, size smaller than is usual in the
genus, head and body less than 200 mm. (St. Leger); and the lucifer group,
zygomatic plate and P. 3 as in poensis, but size larger, over 200; this group will
probably ultimately be found to be referable to the genus Paraxerus. lucifer
may be remarked on as a species with an attractive colour pattern.
Forms seen : acticola, annulatus, atibryi, hongensis, hrauni, canaster, caurinus,
coenosus, daucinus, ele«ans, emissus, i^ambianus, liardvi, isabellinus, kajfensis,
keniae, lateris, leonemis, libericus, loandiiis, lualabae, lucifer, maculatus, madogae,
multicolor, mutabilis, nyansae, obfuscatus, otnensis, pasha, poensis, punctatus, rho-
desiae, rufobrachium, ruwenzorii, semliki, senescens, undulatus, vexillaris, vulcanius.
List of N.\med Forms
Subgenus Heliosciurus, Trouessart
(Revised by Ingoldby, Proc. Zool. Soc. London, 1927, p. 471)
1. HELIOSCIURUS GAMBI.^NUS GAMBIANUS, Ogilby
1835. Proc. Zool. Soc. London, p. 103.
Gambia.
Synonym: (?) annulatus, Desmarest, 1822, Mamm. ii, p. 338. This
species often regarded as unidentifiable.
annularis, Schinz, 1845, Syn. Mamm. Bd, II, p. 14.
albina. Gray, .Ann. Mag. Nat. Hist. 3, XX, p. 329, 1867,
nom. nud.
2. HELIOSCIURUS GAMBIANUS SENESCENS, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. 544.
Thies, Senegal.
3. HELIOSCIURUS GAMBIANUS LIMB.^TUS, Schwarz
1916. Wiesbaden Jahrb. ver Natk. 68, p. 65.
E. Cameroons.
4. HELIOSCIURUS G.AMBI.ANUS BONGENSIS, Heuglin
1877. Reis. Nord. Ost. Afr., II, p. 59.
Bahr-El-Ghazal, Sudan.
5. HELIOSCIURUS G.\MBIANUS CANASTER, Thomas & Hinton
1923. Proc. Zool. Soc. London, p. 256.
Jebel Marra, Darfur.
6. HELIOSCIURUS GA.MBIANUS MULTICOLOR, Ruppell
1835. Neue Wirbelth, p. 38, pi. 13.
Valleys of Kulla and east slope of coast range, .Abyssinia.
7. HELIOSCIURUS GAMBI.ANUS L.ATERIS, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. 102.
Lado, Sudan.
8. HELIOSCIURUS GAMBIANUS ELEGANS, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. 103.
Mount Elgon, Kenya.
9. HELIOSCIURUS GAMBIANUS COENOSUS, Thomas
1909. .Ann. Mag. Nat. Hist. 8. TV, p. 104.
19° 30' E. on River Ubangui, Congo.
26 — Living Rodents — I
402 HELIOSCIURUS
10. HELIOSCIURUS GAMBIANUS OMENSIS, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. 104.
Lower Omo River, near Lake Rudolf, E. Africa.
11. HELIOSCIURUS G.AMBIANUS MADOGAE, Heller
lyii. Smiths. Misc. Coll. 56, no. 17, p. i.
Unia, 50 miles north of Nimula, Uganda.
12. HELIOSCIURUS CJAMBIANUS KAEFENSIS, Neumann
igo2. Siu. Ber. Ges. Nat. Fr. Berlin, p. 57.
.Anderatscha, Kaffa, Abyssinia.
13. HELIOSCIURUS GAMBLANUS ABASSENSIS, Neumann
1902. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 57.
South of Lake Abassi, Abyssinia.
14. HELIOSCIURUS GAMBIANUS RHODESIAE, Wroughton
IQ07. Manch. Mem. Lit. Phil. Soc, no. 5, p. 15.
Plateau west of Mchinga Escarpment, N. Rhodesia.
15. HELIOSCIURUS GAMBIANUS LOANDICUS, Thomas
1923. Ann. Mag. Nat. Hist. 9, XI, p. 521.
N'dola Tando, Northern Angola.
16. HELIOSCIURUS GAMBIANUS MUTABILIS, Peters
1852. Monatsber. Ak. Wiss. Berlin, p. 273.
Boror, Portuguese E. Africa.
Synonym: shiremis. Gray, 1867, Ann. Mag. Nat. Hist. 3, XX, p. 327.
Shire River, Nyasaland.
17. HELIOSCIURUS GAMBIANUS BEIRAE, Roberts
1913. Ann. Transv. Mus., IV, p. 78.
Beira, Portuguese E. Africa.
18. HELIOSCIURUS GAMBIANUS CHIRINDENSIS, Roberts
1913. Ann. Transv. Mus., IV, p. 78.
Chirinda Forest, S.-E. Mashonaland.
19. HELIOSCIURUS GAMBIANUS UNDULATUS, True
1892. Proc. U.S. Nat. Mus., XV, p. 465.
Kilimanjaro, Tanganyika.
Synonym: iindulattis mancitzi, Miiller, 1911, Zool. .Anz. 37, p. 82.
Kilimanjaro.
20. HELIOSCIURUS GAMBIANUS DAUCINUS, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. loi.
Mombasa, Kenya.
21. HELIOSCIURUS GAMBIANUS DOLOSUS, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. 100.
Mafia Island, Tanganyika.
22. HELIOSCU'RUS GAMBIANUS SHINDI, Heller
1914. Smiths. Misc. Coll., LXIIl, no. 7, p. 7.
Mt. Uniengo, Taita Hills, Kenya.
23. HELIOSCIURUS GAMBIANUS PUNCTATUS, Temminck
1853. Esq. Zool. Cote de Guine, p. 138.
Guinea Coast.
HELIOSCIURUS 403
24. HELIOSCIURUS GAMBIANUS SAVANNIUS, Thomas
1923. Ann. Mag. Nat. Hist. 9, XI, p. 521.
Beoumi, Ivory Coast.
25. HELIO.SCIURUS GAMBIANUS KENIAE, Neumann
1902. Sitz. Ber. Oes. Nat. Fr. Berlin, p. 176.
Mount Kenya.
26. HELIOSCIURUS GAMBIANUS RUFOBRACHIUM, Waterhouse
1842. Ann. Mac. Nat. Hist, i, X, p. 202 (published November).
Fernando Po.
■Synonym: rtifobrachiatiis, Waterhouse, 1842, Proc. Zool. Soc. London,
p. 12S (published January', 1843).
aubryi, Milne-Edwards, 1867. Rev. Zool., XIX, p. 228.
Gaboon.
27. HELIOSCIURUS GAMBIANUS PASHA, Schwann
1904. Ann. Mag. Nat. Hist. 7, XIII, p. 72.
Bellima, Monbuttu, N.-E. Congo.
28. HELIOSCIURUS GAMBIANUS BENGA, Cabrera
1917. Bol. Real. Soc. Espanola, 17, p. 517.
Cabo San Juan, .Spanish Guinea.
2<j. HELIOSCIURUS GAMBIANUS ISABELLINUS, Gray
1867. Ann. Mag. Nat. Hist. 3, XX, p. 329.
Lower Niger.
30. HELIOSCIURUS GAMBIANUS LEONENSIS, Thomas
1923. Ann. Mag. Nat. Hist. 9, XI, p. 523.
Sierra Leone.
31. HELIOSCIURUS GAMBIANUS EMISSUS, Thomas
1923. Ann. Mag. Nat. Hist. 9, XI, p. 520.
S.-E. Nigeria.
32. HELIOSCIURUS GAMBL^NUS ACTICOLA. Thomas
1923. Ann. Mag. Nat. Hist. 9, XI, p. 525.
Fernando Po.
33. HELIOSCIURUS G.\MBIANU3 C.'\URINUS, Thomas
1923. Ann. Mag. Nat. Hist. 9, XI, p. 523.
Gunnal, Portuguese Guinea.
34. HELIOSCIURUS GAMBIANUS HARDYI, Thomas
1923. Ann. Mag. Nat. Hist, g, XI, p. 519.
Beoumi, N. Ivory' Coast.
35. HELIOSCIURUS GAMBIANUS OBFUSCATUS, Thomas
1923. Ann. Mag. Nat. Hist. 9, XI, p. 526.
Oban district, S.-E. Nigeria.
36. HELIOSCIURUS GAMBIANUS NUCULATUS, Temminck
1853. Esq. Zool. Cote de Guin6, p. 130.
"Guinea." Probably Gold Coast.
Synonym: aschantiensis, Neumann, 1902, Sitz. Ber. Ges. Nat. Fr.
Berlin, p. 175. Ashanti, Gold Coast.
uaterhottsii, Grav, Ann. Mag. Xat. Hist. 3, XX, p. 328,
1867.
404 HELIOSCIURUS
37. HELIOSCIURUS GAMBIANUS NYANSAE, Neumann
1902. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 56.
Kwa Kitoto, Kavirondo, Uganda.
38. HELIOSCIURUS GAMGIANUS SEMLIKII, Thomas
1907. Ann. Mag. Nat. Hist. 7, XIX, p. 120.
Beni, Semliki, Congo.
39. HELIOSCIURUS GAMBIANUS MEDJIANUS, Allen
1922. Bull. Amer. Mus. Nat. Hist., XLVII, p. 46.
Mcdje, Ituri Forest, Congo.
40. HELIOSCIURUS GAMBIANUS RUBRICATUS, .Allen
1922. Bull. Amer. Mus. Nat. Hist., XLVII, p. 47.
Near Lubila River, 50 miles S.-W. of Avakubi, Congo.
41. HELIOSCIURUS GAMBIANUS ARRHENII, Lonnberg
1917. Stockholm Vet. Akad. Handl. 58, 2, p. 68.
Masisi, near Lake Kivu.
42. HELIOSCIURUS GAMBIANUS LUALABAE, Thomas
1923. Ann. Mag. Nat. Hist. 9, XI, p. 520.
Lodja, Upper Lukenge River, S. Congo.
43. HELIOSCR'RUS G.AMBIANUS LIBERICUS, Miller
1900. Proc. Washington Acad. Sci., II, p. 633.
Mount Coflfee, Liberia.
44. HELKJSCIURUS GAMBIANUS BRAUNI, St. Leger
1935. Nov. Zool. XXXIX, p. 252.
Fazcnda Congulu, Amboim district, Angola.
Subgenus AetliDsciurus, Thomas
ruKensorii Group
45. HELIOSCIURUS RUVVENZORII RUWENZORII, Schwann
1904. Ann. Mag. Nat. Hist. 7, XIII, p. 71.
Luimi Valley, Ruwenzori.
46. HELIOSCIURUS RUWENZORII VULCANIUS, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. 476.
Volcanoes north of Lake Kivu, Belgian Congo.
puensis Group
47. HELIO.SCIURUS POENSIS POENSIS, Smith
1830. S. Afr. Quart. Joum., 2, p. 128.
Fernando Po.
Synonym: olivaceus, Milne-Edwards, 1S67. Rev. Mag. Zool., XIX,
p. 228.
affinis, Rhoads, 1896, Proc. Acad. Philadelphia, p. 521.
4S. HELIOSCIURUS POEiNSIS MUSCULINUS. Temmmck
1853. Esq. Zool. Cote dc Guine, p. 141.
"Guinea." Probably Gold Coast.
4y. HELIOSCIURUS POENSIS SUBVIRIDESCENS, Le Conte
1857. Proc. Acad. Philadelphia, p. 11.
Gaboon.
HKLIOSCIURUS— PARAXERUS 405
Iticifer Group
(For remarks on generic status of this group see p. 400.)
50. Hr.LIOSCIURUS LUCIFER, Thomas
1897. Proc. Zool. Soc. London, p. 430.
Kombe Forest, Masuku Range, N. Nyasa.
51. HKLIOSCIURUS VEXILLARIUS, Kershaw
1923. Ann. Mag. Nat. Hist. 9, XI, p. 591.
Usambara, Tanganyika.
incertae sedis
52. HKLIOSCIURUSC-) BAYONII, Bocage
1890. Jom. Sci. Lisbon, II, p. 3.
Braganca, Angola.
(A Funisciiirus according to G. M. Allen, 1939.)
Genus 19. PARAXERUS, Major
1893. Par.\xerus, Forsyth Major, Proc. Zool. Soc. London, p. 189.
1918. Tamiscus, Thomas, Arm. Mag. Nat. Hist. 9, I, p. 33. Sciunis emini, Stuhlmann.
Type Species. — Sciurus cepapi, Smitli.
Range. — Eastern and South .\frica: Sudan, Somaliland, Kenya, Uganda,
Tanganyika, Zanzibar, Congo, Rhodesia, Mozambique; South-
west Africa, Bechuanaland, Zululand.
Number of Forms. — About forty-four.
Characters. — Cheekteeth semihypsodont, the upper molars with the three
depressions, particularly the second, tending to take the form
of re-entrant folds, the pattern clear and definite, and apparently usually long
retained. M.3 with two depressions, the second very broad, or occasionally in
this tooth three depressions traceable. P. 3 present, well marked; P. 4 with no
prominent anterior cusp. In the lower teeth, the posterointernal cusp is strong,
the teeth are more or less transverselv ridged, with three depressions separating
four ridges; the second, which corresponds to the main depression of less
specialized Sciurinae, appearing as a broad inner re-entrant fold, with a small
outer re-entrant fold opposite to it. The cusps, particularly the anterointernal,
moderately high. In old age, the teeth are more or less simplified to a two-
lobed structure.
Skull essentially as in Funisciiirus (next to be described), except that the
rostrum does not tend to become elongated.
External characters as in normal Tree-squirrels.
The boehtni group were referred to a genus Tamiscus by Thomas, which
Hollister regarded as a subgenus, on account of their dorsal stripes, the infra-
orbital foramen said to be less open (but the difference is very small; there
appears to be less difference to me between Paraxerus and " Tamiscus" than
between individual specimens of Menetes berdmorei); the "molars less hvpso-
dont, the crowns more abruptly marked off from the roots . . . the large internal
4o6 PARAXERUS
root narrow, well spaced from its neighbours on each side, and abruptly
broadens out above at crown." The incisors are more proodont, but within
other genera this is a very variable character; compare, for instance, Callosciunis.
This division may be of subgeneric value, but to me Tomiscas seems no more
than a specific group of Paraxerus.
Forms seen : alexamiri, f]iii;iistus, animosus, antoniiie, aniscensis, buehiiii,
hrldgciinini, byutti, capitis, cvpapi, e/cctiis, einim, exgeaniis, flavii'ittis, frerei,
ganiirui, gazellac, ihcamis, jacksoni, lastii, Iiiiniris, tiiosstimbiciis, ochraceus, ornatus,
pcillititus, pcrcivali, pliahicna, c/iiotiis, sindi, soccniiis, spoiisiis. siiaheliciis, swynncr-
tuni, tuiiae, vukanoruin, viilei.
I am inclined provisionally to divide this genus into four groups:
hoehmi group : smallish striped squirrels; usually four black stripes bordering
three lighter ones, general effect 7V;;«/<«-like; hoehmi is coloured rather
differently from the other species referred to the group; the stripes can
become faint; the genus " Tumiscus" of Thomas.
flavivittis group: Atlantoxerus-\\kQ forms; usually pale, with thick white
flank-stripe present.
palliatus group : usually larger than cepapi group, becoming about maximum
size for genus; tail red or orange; belly red; sometimes head red. P.
bridgemani is a type which tends to be intermediate between this group
and the cepapi group.
cepapi group: usually smaller than the last; dull-coloured squirrels with no
red or orange markings, so far as seen. Includes ochraceus.
Not allocated: hvatti, hitherto referred to Aethosciiinis, is a Paraxerus as
regards the dental formation of the type skull. I have not seen the sub-
species described by Allen & Loveridge.
This arrangement must be regarded as provisional.
List of Named Forms
cepapi Group
I. PARAX1;RUS cepapi cepapi. Smith
1836. App. Report Explor. S. Africa, p. 43.
Marico Ri\'er, Rustenburg district. Trans\'aal.
Synonym: nnitahilis, Huet, 1880, Nouv. Arch. Mus., p. 143 (not of
Peters).
(?) siiperciliaris, Wagner, Schreb. Saugth. Suppl., III.
1843, p. 212.
z. PARAXERUS CEPAPI CHOBIENSIS, Roberts
1932. Ann. Transv. Mus., XV, p. g.
Kabulabula, Chobi River, N. Bechuanaland.
3. PARAXERUS CEPAPI MAUNENSIS, Roberts
1932. Ann. Transv. Mus., XV, p. g.
Maun, Ngamiland.
PARAXERUS 407
4. PARAXERUS CEPAPI SOCCATUS, Wroughton
1909. Ann. Mag. Nat. Hist. 8, III, p. 515.
Hewe River, N. Angoniland, Nyasaland.
5. PARAXKRUS CEPAPI PHALAENA, Thomas
1926. Proc. Zool. Soc. London, p. 296.
Hetween Ukuambi and Ondong^va, Ovamboland.
6. PARAXERUS CEPAPI SINDI, Thomas & Wroughton
1908. Proc. Zool. Soc. London, p. 543.
Tete, Lower Zambesi.
7. PARAXERUS CEPAPI KALAHARICUS, Roberts
1932. Ann. Transv. Mus., XV, p. 10.
Mabeleapudi, Kalahari.
S. PARAXERUS CEPAPI AURIVENTRIS, Roberts
1926. Ann. Transv. Mus., XI, p. 250.
Magudi, Portuguese E. Africa.
9. PARAXERUS CEPAPI QUOTUS, Wroughton
1909. Ann. Mag. Nat. Hist. 8, III, p. 516.
Katanga, Congo.
10. PARAXERUS CEPAPI YULEI, Thomas
1902. Proc. Zool. Soc. London, p. 120.
Muezo, near Lake Mweru.
11. PARAXERUS OCHRACEUS OCHR.\CEUS, Huet
1880. Nouv. Arch. Mus., p. 154, pi. VII, fig. 2.
Bagamoyo, near Dar-es-Salaam, Tanganyika.
12. PARAXERUS OCHRACEUS ARUSCENSIS, Pagenstecher
1885. Jahrb. Hanib. Wiss, Aust. 2, p. 42.
Pangani River near the coast and Aruscha, Mt. Meru, Tanganyika.
Synonym: (?) ^au/i, Matschie, 1894, Sitz. Ber. Ges. Nat. Fr. Berlin,
p. 256. Tanganyika.
13. PAR.AXERUS OCHRACEUS SALUTANS, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. 106.
Dar-es-Salaam, Tanganyika.
14. PARAXERUS OCHR.\CEUS ELECTUS, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. 106.
Elgeyo, Kenya Colony.
15. PARAXERUS OCHRACEUS ANIMOSUS, DoIIman
1911. Ann. Mag. Nat. Hist. 8, VIII, p. 655.
Mount Nyiro, Kenya Colony.
16. PARAXERUS OCHRACEUS PERCIVALI, Dollman
igii. Ann. Mag. Nat. Hist. 8, VIII, p. 653.
Marsabit, Kenya Colony.
(According to G. M. .-Mien the correct name for this subspecies is
P. 0. affiiiis, Trouessart, Cat. Mamm., Viv. Foss. pt. 3, p. 406, 1897.)
Synonym: ochraceus atigustus, Dollman, 191 1, Ann. Mag. Nat. Hist. 8,
VIII, p. 654. Marsabit, Kenya.
17. PARAXERUS OCHRACEUS IC^HARI, Heller
191 1. Smiths. Misc. Coll. LVI, no. 17, p. 2.
Meru Boma, north-east of Mount Kenya.
4oS PARAXERUS
i8. PARAXKRLS OCHRACEL'S JACKSONI, dc Winton
1897. Ann. Mag. Nat. Hist. 6, XIX, p. 574.
Kikuyu, Kenya Colony.
Synonym: jacksoiti capitis, Thomas, 1909, Ann. Mag. Nat. Hist. S, IV,
p. 105. Nairobi Forest.
It). PARAXERUS OfHRACEUS GANANA, Rhoads
1S96. Proc. Acad. Sci. Philadelphia, XLVIII, p. 522.
Ganana River at Bar Madu, Abyssinia.
palliatiis Oroup
20. PARAXERUS BRIDGEMANI, Dollman
1914. Ann. Mag. Nat. Hist. cS, XIV, p. 152.
Panda, Portuguese E. Africa.
21. PARAXERUS PALLIATUS PALLIATUS, Peters
1852. Monatsber. .Akad. Wiss. Berlin, p. 273.
Mozambique.
22. PARAXERUS PALLIATUS ORNATUS, Gray
1864. Proc. Zool. Soc. London, p. 13, pi. i.
Zululand.
23. PARAXERUS PALLIATUS SUAHELICUS, Neumann
1902. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 17S.
Tanganyika coast, opposite Zanzibar.
24. PARAXERUS PALLI.ATUS FRICREI, Gray
1S73. Ann. Mag. Nat. Hist. 4, XII, p. 265.
Zanzibar.
25. P.ARAXERUS PALLIATUS LASTII, Thomas
1906. Ann. Mag. Nat. Hist. 7, XVIII, p. 297.
Zanzibar.
(According to St. Leger probably not distinguishable horn frerei.)
26. PARAXERUS PALLIATUS SWYNNERTONI, Wroughton
1908. Ann. Mag. Nat. Hist. 8, I, p. 305.
Chirinda Forest, N. Rhodesia.
27. PAR.AXERL'S PALLIATUS TAN.AE, Neumann
1902. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 178.
Tana River, Kenya Colony.
zS. PARAXERUS PALLIATUS BARAWENSIS, Neumann
1902. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 178.
Somaliland.
2'). PARAXERL'S SPONSUS SPONSUS, Thomas & Wroughton
1907. Proc. Zool. Soc. London, p. 292.
Inhanibane, Zululand.
30. P.\R.AXERUS SPONSUS TONGENSIS, Roberts
1931. .\nn. Transv. Mus., XIV, p. 229.
Mangusi Forest, N. Zululand.
flavivittis Group
31. PARAXERUS FLAVIVITTIS FLAVIVITTIS, Peters
1852. Reise Mossamb., I, taf. XXIX.
Mossimba, near Mozambique.
PARAXERUS 409
32. PARAXERUS FLAVIVITTIS MOSSAMBICUS, Thomas
igig. Ann. Mag. Nat. Hist. 9, IV, p. 31.
Lumbo, Portuguese E. Africa.
33. PARAXERUS FLAVIVITTIS EXGEANUS, Hinton
1920. Ann. Mag. Nat. Hist. 9, V, p. 311.
Kilwa Kisiwani, Tanganyika.
34. PARAXERUS FLAVIVITTIS IBEANUS, Hinton
1920. Ann. Mag. Nat. Hist. 9, V., p. 312.
Mombasa, Kenya.
boehmi Group
35. P.\RAXERUS BOEH.MI, Reichenow
1886. Zool. Anz., IX, p. 315.
Marungu, S. Congo.
36. PARAXERUS EMINI EMINI, Stuhlmann
1894. Mit. Emin. Pasha Herz. Afrika, p. 320.
Upper Semliki River, Belgian Congo.
Synonym: emini ugandae, Neumann, 1902, Sitz. Ber. Ges. Nat. Fr.
Berlin, p. iSo. Uganda.
37. PARAXERUS EMINI GAZELLAE, Thomas
1918. Ann. Mag. Nat. Hist. 9, I, p. 34.
Meridi, Bahr-el-Ghazal.
38. PAR.AXERUS VULC.\NORUM VULCA.XORUM, Thomas
1918. .\nn. Mag, Nat. Hist. 9, I, p. 35.
Buhamba, near Lake Kivu, Belgian Congo.
39. PARAXERUS VULCANORUM LUNARIS, Thomas
1918. Ann. Mag. Nat. Hist. 9, I, p. 36.
Mubuku Valley, Ruwenzori East.
40. PAR.AXERUS VULCANORUM TANGANYIKAE, Thomas
1918. Ann. Mag. Nat. Hist. 9, I, p. 36.
10 miles west of Baraka, Burton Gulf, Lake Tanganyika.
41. P.\RAXERUS ALEX,.\NDRI, Thomas & Wroughton
1907. .Ann. Mag. Nat. Hist. 7, XIX, p. 376.
Gudima, R. Iri, Upper Welle.
42. PAR.AXERUS ANTONIAE, Thomas & Wroughton
1907. Ann. Mag. Nat. Hist. 7, XIX, p. 377.
Ponthierv'ille, Upper Congo.
Not allocated to group
43- PARAXERUS BYATTI BYATTI, Kershaw
1923. Ann. Mag. Nat. Hist. 9, XI, p. 592.
Moshi, Kilimanjaro.
44. PAR.\XERUS BYATTI LAETIS, Allen & Loveridge
1933. Bull. Mus. Comp. Zool. Har\ard, LXXV, no. 2, p. 96.
Ukinga Mountains, north of Lake Nyasa, Tanganyika.
4IO
FUNISCIURUS
Genus 20. FUNISCIURUS, Trouessart
1880. FUNISCIURUS, Trouessart, le Naturaliste, II, no. 37, p. 293.
Type Species. — Sciiinis /emniscatus, Le Conte.
Range. — Africa : Gambia, Sierra Leone, Ivory Coast, Gold Coast, Nigeria,
Cameroons, Fernando Po, Gaboon, Congo, Angola, Tanganyika,
Ruwenzori, S.-W. Africa.
Number of Forms. — Thirtv-four.
P'uNisciuRus PYRRHOPUS LEONis, Thomas.
B.M. No. 1Q38.6.10.9, ?; .< 2.
Characters. — Siiull wealcly ridged, the parietal ridges usually not joining.
Postorbital process moderate. Palate normal. Zygomatic
plate \vell ridged, but short, the ridge stopping abruptly above the upper border
of the infraorbital foramen, not approaching the superior border of the rostrum.
Masseter knob weak or absent. Infraorbital foramen sometimes with a ridge
curving upwards from its upper border and joining the forepart of ridge of
zygomatic plate. Rostrum tends to become elongated. Cheekteeth more
specialized than in Paraxerus, or for that matter almost all other Squirrels;
tending to become completely flat-crowned, or nearly so; semihypsodont. P. 3
Fig. 1 02. FuNisciuRus pyrrhopus leonis, Thomas.
B.M. No. 1938.6.10.9, $; X 2.
Fig. 103. FUNISCIURUS pyrrhopus leonis, Thomas.
Cheekteeth; B.M. No. 1938.6. 10.9, $; X 8.
412 I-X'NISCIURUS
well de\eIoped. Lower molars like those of Anomaliinis or Erethizon in general
appearance; three wide inner re-entrant folds, and one wide outer re-entrant
fold present. Upper molars fiat in adult, the three depressions more or less
appearing as re-entrant folds; the first sometimes curving round and joining
the third; the second depression the widest. M.3 usually with elements as the
other teeth, and often relatively small.
External characters as in normal Tree-squirrels. According to Pocock,
baculum minute in this genus and in Paraxerus, of those he examined.
Forms seen : cikkii, cuieiytltnis, aiiricuhitus, bandurum, heatits, bovdii, carrtt-
thersi, c/irvsippiis, cotigiciis, erytliros^enys, flavinus, interior, isabella, lemniscatus,
leonis, leucustigiita, maiidingu, mayumbicus, mystax, nigrensis, tiiveatiis, oc/irogaster,
oenone, olirelliis, oliricie, peinbertoni, pvrrhopiis, raptoniiii, siibstriatiis, talboti,
langanyikac.
Three groups are recognizable among the material examined, though these
must be treated as provisional :
Itmniscatus group: back longitudinally striped, as in Paraxerus boelimi group;
usually four black stripes bordering tiiree paler ones.
congicus group: back without longitudinal stripes; usually pale forms
reminiscent of Atlantuxirus or Paraxerus flavivittis gnjup; with a well-
marked whitish flank-stripe each side; only the race interior is darker in
colour.
pyrrliopus group : all others. Normally darker than congicus group (which
perhaps should be referred to present group); limbs and head red in
prrrhopiis, usually, which in most cases has a narrow light flank-stripe
present; auriculatus has some red colouring on head and rump, but not
on the limbs, and the flank-stripe is present; carruthersi is a dark form,
without stripes. But intermediate forms appear to exist between these
types.
List of N.;\med Forms
lemniscatus Group
1. FUNISCIURUS LEMNISC.VrU.S LEMNISC-^TUS, Le Conte
1857. Proc. Acad. Nat. Sci. Philadelphia, p. 11.
Rio Muni, Gaboon.
Synonym: sharpei. Gray, 1873, Ann. Mag. Nat. Hist. 4, XII, p. 265.
Gaboon.
2. FUNISCIURUS LEMNISCATUS IS,\BELL.\, Gray
1S62. Proc. Zool. Soc. London, p. 180, pi. XXIV.
Cameroon Mountain.
,1. PX'NISCIURUS MAYUMBICUS, Kershaw
1923. Rev. Zool. Afr., XI, 4, p. 363.
Ganda Sundi, Mayumbe Province, Lower Congo.
4. FUNISCIURUS POOL II, Jentink
igo6. Notes Leyd. Mus., XXVIII, p. 139.
Stanley Falls, Upper Congo.
FUNISCIURUS 413
congicus Group
5. FUNISCIURUS CONGICUS CONGICUS, Kuhl
1820. Beitr. Zool., p. 66.
Canboca, N. Angola.
Synonym: praetextus, Wagner, Schreb. Saug. Suppl. 3, p. 316, 1843.
6. FUNISCIURUS CONGICUS OLIVELLUS, Thomas
1904. .^nn. Mag. Xat. Hist. 7, XIII, p. 410.
Cunga, N. .Angola.
7. FUNISCIURUS CONGICUS FU.^VINUS, Thomas
1904. .\nn. Mag. Nat. Hist. 7, XIII, p. 411.
Capongombi, S. Angola.
8. FUNISCIURUS CONGICUS OENONE, Thomas
1926. Proc. Zool. Soc. London, p. 297.
Cunene Falls, Ovamboland.
9. FUNISCIURUS CONGICUS I.NTERIOR, Thomas
1916. Ann. Mag. Nat. Hist. 8, XVIII, p. 236.
Inkongo, South Congo.
pyrrhopiis Group
10. FUNISCIURUS PYRRHOPUS PYRRHOPUS, F. Cuvier
1833. Hist. Nat. Mamm. IV (66), p. 2.
Gaboon.
SjTionym: ruhripes, du Chaillu, i860, Proc. Boston Soc. Nat. Hist., VII,
p. 366. Gaboon. (A valid race according to G. M.
.\llen, 1939.)
erytkrops. Gray, 1867, Ann. Mag. Nat. Hist. 3, XX, p. 330.
11. FUNISCIURUS PYRRHOPUS LEONIS, Thomas
1905. Ann. Mag. Nat. Hist. 7, XV, p. 79.
Bo, Sierra Leone.
12. FUNISCIURUS PYRRHOPUS AKK.A, de Winton
1899. .■Xnn. Mag. Nat. Hist. 7, IV, p. 356 (footnote).
Tingasi, Monbuttu, N.-E. Congo.
Synonym: emini, de Winton, 1895, Ann. Mag. Nat. Hist. 6, XVI, p. 197.
Not of Stuhlmann.
Ktntoni, Neumann, 1900, Zool. Jahrb., 13, p. 547.
13. FUNISCIURUS LEUCOSTIG^LA LEUCOSTIGMA, Temminck
1853. Esq. Zool. Cote de Guine, p. 133.
Gold Coast.
14. FUNISCIURUS LEUCOSTIGNU TALBOTI, Thomas
1909. .^nn. Mag. Nat. Hist. 8, IV, p. 478.
Oban, S.-E. Nigeria.
15. FUNISCIURUS LEUCOSTIGMA NIVE.-\TUS, Thomas
1923. Ann. Mag. Nat. Hist. 9, XI, p. 522.
Beoumi, 12 miles east of Bandama, Ivory Coast.
16. FUNISCIURUS PEMBERTONI, Thomas
1904. .\nn. Mag. Nat. Hist. 7, XIV, p. 201.
Dondo. Cuanza River, .Angola. (A race of F. pvrrhopus according to
G. M. .Allen.)
414 FUNISCIURUS
17. FUNISCIURUS RAPTORUM, Thomas
1903. Ann. Mag. Nat. Hist. 7, XI, p. 80.
Forcados, Lower Nigeria.
iS. FUNISCIURUS ANERYTHRUS ANERYTHRUS, Thomas
iSyo. Proc. Zool. Soc. London, p. 447.
Buguera, west of Lake Albert, Congo.
19. FUNISCIURUS ANERYTHRUS BANDARUM, Thomas
1915. Ann. Mag. Nat. Hist. 8, XVI, p. 146.
Baniingui River, Upper Shari.
20. FUNISCIURUS ANERYTHRUS NIAPU, Allen
IQ22. Bull. Amer. Mus. Nat. Hist., XLVII, p. 52.
Niapu, Belgian Congo.
21. FUNISCIURUS AURICULATUS .\URICULATUS, Matschie
iSgi. Archiv. Naturg., I, 3, p. 353.
Kribi, Cameroons.
22. FUNISCIURUS AURICULATUS BE.ATUS, Thomas
igio. Ann. Mag. Nat. Hist. 8, V, p. ig6.
Benito River, French Congo.
23. FUNISCIURUS AURICULATUS BOYDI, Thomas
1910. Ann. Mag. Nat. Hist. 8, V, p. ig6.
Mussaka, Lower Mongo River, east of Cameroon Mountain.
24. FUNISCIURUS AURICULATUS OLIVIAE, Dollman
191 1. Ann. Mag. Nat. Hist. 8, VHI, p. 733.
Oban, S.-E. Nigeria.
25. FUNISCIURUS MYSTAX MYSTAX. de Winton
1898. Ann. Mag. Nat. Hist. 7, II, p. 9.
Benito River, French Congo.
26. FUNISCIURUS MYSTAX OCHROGASTER, Cabrera & Ruxton
1926. Ann. Mag. Nat. Hist. 9, XVII, p. 597.
Luluabourg, Kasai, S. Congo.
27. FUNISCIURUS LEUCOGENYS, Waterhouse
1S42. Ann. Mag. Nat. Hist. X, p. 202.
F\-rnando Po.
Synonym: erylliyogenys, Waterhouse, 1843, Proc. Zool. Soc. London,
p. 129, 1842.
28- FUNISCIURUS SUBSTRIATUS, de Winton
1899. Ann. Mag. Nat. Hist. 7. IV, p. 357.
Kintampo, Gold Coast.
29. FUNISCIURUS MANDINGO MANDINGO, Thomas
1903. Ann. Mag. Nat. Hist. 7, XI, p. 79.
Nianimaru, Gambia.
30. FUNISCIURUS MANDINGO NIGRENSIS, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. 478.
Abutschi, Lower Niger.
31. FUNISCIURUS CARRUTHERSI CARRUTHERS], Thomas
1906. Ann. Mag. Nat. Hist. 7, XVIII, p. 140.
Ruwenzori East.
FUNISCIURUS— PROTOXERUS 415
32. FUNISCIURUS CARRUTHERSI TANGANYIKAE, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. 477.
Usumbura, north end of Lake Tanganyika.
33. FUNISCIURUS C.ARRUTHKRSI CHRYSIPPUS, Thomas
1923. .Ann. Mag. Nat. Hist. 9, XI, p. 522.
Wabembe, north-west of Lake Tanganyika.
34. FUNISCIURUS C.ARRUTHFRSI BIRUNGENSIS, Gyldenstolpe
1927. Arkiv. Zoologi, Band 19B, no. 6, p. i.
Mount Karissimbi, Birunga volcanoes, East Congo.
Genus 21. PROTOXERUS, Forsyth Major
1893. 1'rotoxerus, Forsyth Major, Proc. Zool. Soc. London, p. 189.
Type Species. — Sciurus stangeri, Waterhouse.
R.\NGE. — Africa: Kenya, Uganda; Gold Coast, Nigeria, Cameroons,
Fernando Po, Gaboon, Congo, Angola.
Number of Forms. — Fourteen.
Ch.\racters. — Infraorbital foramen large, round, and at ma.ximum develop-
ment for the family, not forming a canal, and apparently
well open enough to transmit a small strand of muscle. The portion of the
zygomatic plate behind it considerably narrowed. Zygomatic plate strongly
ridged, but less so than in Xerus, though it approaches the type found in that
genus. Frontals broad, postorbital process moderately developed; rostrum
short, broad. Parietal ridges not hea\'y, but joining in adult. Palate normal.
Bullae not reduced. Toothrows not reduced. Cheekteeth |, relatively brachy-
odont, in structure suggesting the Xerus type; the usual four ridges and three
depressions traceable in the upper series, but the whole effect usually rather
simple. The centre depression rather broad, the anterior and posterior ones
narrow. Forsyth Major suggested that this was the type of tooth from which
the normal Xerus dentition was derived. Central depression in lower molars not
becoming obliterated, but in most seen the pattern is not clear. Upper and lower
incisors considerably thickened anteroposteriorly, their surfaces plain ; there is
a certain tendency towards this character in all genera of African Tree-squirrels.
Size large, up to 310 mm. head and body. Tail densely bushy, often rather
longer than head and body. Digits with the arrangement characteristic of
arboreal types. Ventral surface of body poorly furred, often tending to be nearly
naked, a rare feature in the Order.
Forms seen : bea, caliurus, centricola, dissonus, eborivorus, loandae, moereiis,
nigeriae, notabilis, signatus, stangeri, temmincki.
List of Named Forms
I. PROTOXERUS STANGERI STANGERI, Waterhouse
1S42. Proc. Zool. Soc. London, p. 127.
Fernando Po.
Synonym: nordhoffi, du Chaillu, i860, Proc. Boston Soc. Nat. Hist.,
VII, p. 363. Gaboon.
4i6 PROTOXERUS— MYRSILUS
z. PROTOXl-RUS STANGERl CALLIURUS, Peters
1S74. Monatsber. Akad. Wiss. Berlin, p. 707.
Cameroons.
3. PRdTOXKRUS STANGERl UISSONL'S, Thomas
1923. Ann. Mag. Nat. Hist. 9. XI, p. 527.
Bitye, Ja River, Cameroons.
4. PR()Tt)Xi:RUS STANGERl NTGERIAE. Thomas
1Q06. .Ann. Mag. Nat. Hist. 7, XVIH, p. 296.
Abutschi, Lower Niger.
5. PROTOXERUS STANGERl TEMMINCKI, Anderson
1879. Zool. Yunn, p. 229 (note).
ChiM Coast.
Synonym: C(iH;Vf/>s, Temminck, 1S53, Esq. Zool. Cote de Guine, p. 127,
Gold Coast. Not of Gray, 1842.
6. PROTOXERUS STANGERl EBORIVORUS, du Chaillu
1S60. Proc. Boston Soc. Nat. Hist., VH, p. 363.
Gaboon.
Synonym: (?) stibalbidtis, du Chaillu, same reference, p. 365.
7. PROTOXERUS STANGERl PERSONATUS, Kershaw
1923. Rev. Zool. Afr., XI, 4, p. 364.
A'lakia Ntete, Lower Mayumbe, Congo.
8. PROTOXERUS STANGERl TORRENTIU.M, Thomas
1923. .Ann. Mag. Nat. Hist. 9, XI, p. 529.
Stanley Falls, Congo River.
q. PROTOXICRUS STANGERl NOTABILIS. Thomas
1923. Ann. Mag. Nat. Hist. 9, XI, p. 528.
Avakubi, Ituri Forest, Belgian Congo.
10. PROTOXERUS STANGERl MOERENS, Thomas
1923. Ann. Mag. Nat. Hist. 9, XI, p. 527.
Lobi, near Angu, Uelle River, Belgian Congo.
11. PROTOXERUS STANGERl SIGNATUS, Thomas
1910. Ann. Mag. Nat. Hist. 8, V, p. 85.
Lodja, near Upper Lukenye River, S. Congo.
12. PROTOXERUS STANGERl LOANDAE, Thomas
1906. Ann. Mag. Nat. Hist. 7, XVIII, p. 296.
Canhoca, N. Angola.
13. PROTOXERUS ST.'\NGERI CENTRICOLA, Thomas
1906. Ann. Mag. Nat. Hist. 7, XVIH, p. 297.
Entebbe, Uganda.
14. PROTOXERUS STANGERl BEA, Heller
1912. Smiths. Misc. Coll., LIX, no. 16, p. 2.
Lukosa River, Kakamega Forest, Kenya.
Genus 22. MYKSILUS, Thomas
1909. Myrsilus, Thomas, Ann. Mag. Nat. Hist. 8. Ill, p. 470.
Type Species. — SciurKs auhinnii. Gray.
MYRSILUS— EPIXERUS 417
Range. — West Africa : Liberia, Ashanti.
Number of Forms. — Two.
CuAR.'VCTFR.';. — E.sscntial sivull characters, including the large infraorbital
foramen, as Prutoxerus, but skull higher in frontal region, and
nasals slanting downwards anteriorly. Zygomatic plate rather weaker. In the
four skulls examined, the cheekteeth have much more clearly marked ridges
and depressions than the majority of Protoxerus. P. 3 present, very small.
Incisors of similar type to those of Protoxerus.
E.xternally differing from Protoxerus in the normally furred belly, and the
narrower tail, which is less densely bushy. Smaller than Protoxerus.
Remarks. — This genus is not very clearly distinguishable from Protoxerus,
but probably must stand, at any rate provisionally, until more
material comes to hand. The well-furred belly and the more strongly ridged
cheekteeth (constant .') are the main distinguishing characters.
Forms seen : auhitmii.
List of Named Forms
1. MYRSILUS AUBINNII AUBINNII, Gray
1873. .'\nn. Mag. Nat. Hist. 4, XII, p. 65,
Fantee, Ashanti, \V. .Africa.
2. MYRSILUS AUBINNII SALAE, Jentink
1881. Notes Leyden Mus., Ill, p. 63.
Liberia.
Genus 23. EPIXERUS, Thomas
igog. Epi.\ERUs, Thomas, Ann. Mag. Nat. Hist. 8, III, p. 472.
Type Species. — Sciurus wilsoni, du Chaillu.
Range. — West Africa : Gold Coast, Cameroons, Gaboon.
Number of Forms. — Two.
Char.\cters. — Closely related to Protoxerus; skull longer and narrower;
infraorbital foramen of siinilar tvpe, but much less open in
adult skulls examined, its appearance slitlike, though scarcely forming a canal.
Frontals relatively narrower, and muzzle longer. Bullae considerably reduced.
Palate extending very slightly more behind toothrows. Toothrows strongly
reduced. In all skulls seen, the depressions tend to take the form of re-entrant
folds. Cusps low. Lower cheekteeth with the elements apparently near Xerus;
a well-marked outer re-entrant fold appearing between the outer main cusps.
P.3 absent. Incisors plain, much thickened anteroposteriorly.
Extcrnallv large; tail thicklv bushv, rather longer than the head and bodv;
belly poorly turrcd, nearly naked often, as in Proto.xerus; digits of arboreal type.
Remarks. — Not clearly very separable from Proto.xerus.
Forms seen : wilsoni, ebii.
-7 — I.i\inj^ Rodents- I
4i« EPIXERUS— XERUS
List of Named Forms
1. HPIXERL'S WILSONI, du Chaillu
iS6o. Proc. Boston Soc. Nat. Hist., VII, p. 364.
Gaboon.
2. EPIXERUS EBII, Tcmminck
1S53. Esq. Zool. Cote de Guine, p. iig.
Forests of Guinea, most abundant at Dabocrom.
Section E. Xeri's. African and Palaearctic terrestrial Squirrels presenting
the following features: lachrymal iisuallv much enlarged; bullae usually
more enlarged than is normal; palate always extending considerably
behind the toothrows; claws prominent to extremely enlarged; fur inost
often bristly.
Genus 24. XERL'S, Ilemprich &: Ehrenberg
1833. Xerus, Hemprich & Ehrenberg, Symb. Phys. Manim. i, te.xt to pi. IX.
1834. Geoscurus, Smith, S. Afr. Quart. Joum., ii, p. 12S. Sciiiriis capeiisis, Kerr.
Valid as a subgenus.
igoq. EiXERi's, Thomas, Ann. Mag. Nat. Hist. S, HI, p. 473. Sciiinis erythropiis,
Geoffroy. Valid as a subgenus.
Type Species. — Xerus brac/notis, Hemprich & Ehrenberg ^=5r»/;i« riitilus,
Cretzchmar.
R.WGK. — Africa: Sudan, Sahara, Abyssinia, Somaliland, Kenya, Uganda;
Senegal, Sierra Leone, Gaboon ; South-west Africa, South Africa.
Ni'MBER OF F'ORMS. — Nineteen.
Cii.AR.xcTERS. — Palate produced posteriorly considerably behind leyel of last
molars; bullae large, round and inflated; lachrymal conspicu-
ously enlarged. Zygomatic plate much narrower, particularly above, yery
strongly ridged along anterior border; intraorbital foramen with well-deyeloped
masseter knob. Parietal ridges, though not well marked, joining in old age.
Hamular processes thick, long, joining bullae. Postorbital process short,
directed backwards. Incisors so far as seen without trace of grooying (compare
Atlantoxerus).
Cheekteeth \ in Xerus s.s. and Geosciurus; the extra premolar present,
minute, in Euxerus. The depressions are well marked; the cusps and ridges not
high; originally there are three depressions between four ridges; in old age, the
posterior ridge and depression tend to become obliterated. M.j smaller and
rather simpler than M.i and i\L2. Lower cheekteeth with well-developed
posterointernal cusp, and the central depression of each tooth more or less
compressed, and restricted, not taking up the greater part of the tooth, as it does
in most genera. With wear the depression takes the form of a wide re-entrant
fold. The anterointerna! cusp, as usual the highest, but the cusps not particu-
larly developed. Cheekteeth strongly hypsodont in Euxerus and Geosciurus;
less so in typical Xerus.
Fig. 104. Xerus rutills rutilus, Cretzchmar.
B.M. No. 97.8.9.11, (J; X ij.
Fig. 105. Xerus rutilus rutilus, Cretzchmar.
B.M. No. 97.8.9M, S; > ij-
420 XERUS
Fur in the loriii cil sliort bristles, wiiich cover the head, hinbs and more or
less the whole body. D.3 in the forefoot definitely longer than D.4, which is
longer than D.2 and D.5. Hindfoot rather long, the three central digits longer
than the outer two, D.3 very slightly the main digit. Tail biishv, nearly as long
as head and body. Ear short.
In the typical subgenus, there is no side-stripe, but in both the other sub-
genera there is a white stripe on each flank.
Eiixeriis retains tlie extra small premolar, which is said to be lost early;
and has a narrow skull, and reduced postorbital processes.
Gcosciiinis, from South Africa, has a much broader skull than is normal in
the genus; the skull is not unlike that of Spcniiophilupsis except that the
bullae are more evenly inflated, the interparietal is not so clearly marked,
and the parietal ridges tend to join. The hindfoot is as a rule relatively
larger than in allies.
These two groups are currently given full generic rank, but the time, I think,
has come not to recognize genera on the least or vaguest excuses. The three
groups are essentially congeneric in all main characters; much more nearly
allied to each other than are some of the numerous subgenera now referred to
Citellus in North America.
The following table should indicate that it is no longer necessary to retain
Eiixcriis and Geosciunis as full genera, in that the characters of the three groups
intergrade to a considerable degree.
Xerus Eiixrrns Geosciunis
Head and Body Length 230-250 (St. Leger) 240-300 250-290
Flank-stripe Absent Present Present
Cheekteeth More brachyodont Hypsodont Hypsodont
P. 3 Absent Present Absent
Skull Rather broad Narrow Broad
Forms seen : agadiiis, capensis, chadensis. dabivjalla, dorsalis, crythrupiis,
fuh'ior, iuternus, lacustris, leiicoiiinbn'iiiis, llniitaiieiis, iinii'sliis, micrudun, namaquen-
sis, piiiuips, nijijions, nitiliis, satiiratiis, steplunnciis.
Li.sT OF Named Form.s
Subgenus Xerus, Hemprich & Ehrenberg
1. XERl'S RL'TILl'S RUTH, U.S. Crctzchmar
1826. Rijppell .Atlns. p. 59, pi. 24.
Eastern slope nf .Abyssinia.
Synonym: hrachyolis, Himprich & IChrenbers;, 1S32, Symp. Rhys., i,
text to pi. I.X.
(?) ahcssiniciis, Gmelin, Syst. Nat. i, p. 141). 1788.
fiisciis. Huet; 18S0. Nouv. .Arch. Mus. N. H. Paris, 2. 3, 139.
XERUS 421
2. XKRUS RUTILUS DABAGALLA, HcuKlin.
1861. Nov. Act. Acad. Leop. Car. Nat. Cur., XXVIII, p. 4, Tab. 2.
Probably Eritrea.
3. XKRUS RUTILUS INTENSUS, Thomas
1904. Ann. Mag. Nat. Hist. 7, XIV, p. 100.
Gerlogubi Wells, Soinalil.md.
4. XERUS RUTILUS STEPHANICUS, Thomas
iyo6. Ann. Mag. Nat. Hist. 7, XVHI, p. 301.
Lake Stephanie, .Abyssinia.
5. XKRUS RUTILUS RUFIFRONS, Dollman.
191 1. Ann. Mag. Nat. Hist. 8, VH, p. 518.
N. Guaso Nyiro, Kenya.
Synonym: (?)^arus, Milne-Edwards, 1867, Rev. Mag. Zool. 229,
"Gaboon" error; Somaliland. Not of Linnaeus.
f). XERUS RUTILUS SATURATUS, Neumann
1900. Zool. Jahrb. Syst., XHI, p. 546.
Kibwezi, Kenya.
7. XERUS RUTILUS DORSALIS, Dollman
1911. .Ann. Mag. Nat. Hist. 8, VH, p. 519.
Lake Baringo, Kenya.
Subgenus Euxerus, Thomas
8. XERUS KRVTHROPUS ERYTHRfJPUS, Geoffrey
1803. Cat. Mamni. p. 178.
W. Africa; possibly Senegal.
Synonym: albovittatus, Desmarest, 1817, Nouv. Diet. Hist. Nat., X,
p. no.
(?) simplex. Lesson, 1S36, Hist. Nat. Mamm., V, p. 402.
Senegal.
marabutus. Lesson, Comp. BufTon, 2, Paris, i, 467, 1838.
prestigialor. Lesson, same reference, p. 468.
lessoiiii, Fitzinger, 1867, Sitz. K. .Ak. Wiss. Wien. math,
nat. CI. 55, I, 488.
9. XERUS ERYTHROPUS MOESTUS, Thomas
19 10. Ann. Mag. Nat. Hist. 8, V, p. 419.
Daru, Sierra Leone.
10. XERUS ERYTHROPUS AGADIUS, Thomas & Hinton
1921. Nov. Zool., XXVIII. p. 6.
Agades, .Air, Sahara.
11. XERUS ERYTHROPUS CHADENSIS, Thomas
1905. .Ann. Mag. Nat. Hist. 7, XV, p. 387.
Yo, Lake Chad.
12. XERUS ERYTHROPUS LACUSTRIS, Thomas
1905. .Ann. Mag. Nat. Hist. 7, XV, p. 388.
Masindi, Unyoro, Uganda.
13. XERUS ERYTHROPUS LIMITANEUS, Thomas S: Hinton
1923. Proc. Zool. Soc. London, p. 255.
Zalingei, Darfur, Sudan.
422 XERUS— ATLANTOXERUS
14. XF.Rl'S ERYTHRdrrS LKUCOUMBRINL'S. Ruppcll
1S35. Ncue Wirb. Fauna Abyss. Saugeth. p. 38.
Abyssinia or Sudan.
15. XERUS ERYTHROPUS MICRODON, Thomas
1905. .\nn. Mag. Nat. Hist. 7, XV, p. 389.
Kitui, Kenya.
Synonym: microdot fithior, Thomas, 1905, Ann. Mag. Nat. Hist. 7,
XV, p. 389. Fort Hall, Kenya.
Subgenus Gcosciurus, Smith
ih. XERUS INAURIS INAL'RIS, Zinimcriiian
17S0. Geogr. Geschichte, 2, 344.
Kaffirland, 100 miles north of Cape of Good Hope.
Synonym: /ciuillaiiti, Kuhl, 1820, Beit. Zool. 67.
setosiis. Smuts, 1832, Enum. Mamm. Cap. 33.
rapensis, Kerr, 1792, Linn. Anim. Kingd. 266.
oinginiatufs, Shaw, Gen. Zool. 2, pt. i, 147, 1801.
(hc/iinslilcits, Gmelin, Syst. Nat. i, p. 151, 1788.
ajricanus, Shaw, Gen. Zool. 2, pt. i, 172, 1801.
17. XERITS INAURIS NAMAQUENSIS, Lichtenstcin
1793. Cat. Rer. Nat. p. 2.
Orange River, Namaqualand.
iS. XERUS PRINCEPS. Thomas
1929. Proc. Zool. Soc. London, p. 106.
Otjitundua, Central Kaokoveld.
The incisors ;ire white in A", capcnsis, but normal (yellow) in X. princeps.
Genus 25. ATLANTOXERUS, Forsyth Major
1893. AtlaJs'toxeris, Forsyth Major, Proc. Zool. Soc. London, p. 189.
Type Species. — Sciurus gctuhis, Linnaeus.
R.ANGE. — Palaearctic section of Africa (North-west): Morocco, Algeria:
"All the Grand Atlas from the Atlantic coast between the Uad
Tensift and Uad Sus, at extreme east of the chain extending to the middle
Atlas and the Algerian Sahara" (G. M. Allen).
Number of Forms. — One.
Characters. — Like Xerus, but fur not definitely bristly, though short and
stiff. D.4 appears in the manus to be relatively longer, so
that rarely D.3 and D.4 may be subequal. A prominent white stripe on each
flank, and sometimes a mid-dorsal stripe may be indicated. Skull flatter than is
normal in Xcrus; upper incisors frequently with traces of a groove.
Parietal ridges poorly marked, evidently not tending to join. P. 3 present,
fairly well developed; dentition essentially as Xerus. Lachrymal not specially
enlarged.
ATLANTOXERUS— SPERMOPHILOPSIS 423
REMARKS.^Thc smaller lachrymal, and fur which is not bristly, may be used
for retaining this genus. It is evidently a primitive member of
the Xerus section, with the characters of that section at least development. It
is the only Squirrel in Africa north of the Sahara.
Forms seen : getulus.
List of Named Forms
I. ATLANTOXERUS GETULl'S, Linnaeus
1758. Syst. Nat. i, loth Ed. p. 64.
Agadir, Morocco.
Synonym: Irkittatus, Gray, 1842, Ann. Mag. Nat. Hist., X, p. 264.
There is at the British Museum a skull labelled "barbarus." The reference
to this name has not been traced.
Genus 26. SPERMOPHILOPSIS, Blasius
1884. Spermophilopsis, Bla.sius, Tageblatt 57ten. Versamml. Deutsch Naturf. Magde-
burg, 5, pp. 322-325.
Type Species.^ — Spertnophilus leptodactylus, Lichtenstein.
Range. — Afghanistan and Russian Turkestan (from Caspian Sea to Semir-
echyia district) (Vinogradov).
Number of Forms. — Three.
Characters. — In essential cranial characters this genus is clearly a member
of the Xerus section. It differs from all Sciuridae seen in the
extreme development of the claws, except Hyoscitirus from Celebes. The
frontals are very broad indeed, the braincase behind the postorbital process
wide. Postorbital process not very large. In none of the skulls is there the
slightest sign of the parietal ridges coming together; the interparietal is well
marked. Occipital region relatively weak. Infraorbital foramen well open,
barely forming canal. Bullae not evenly inflated, though larger than is usual.
Palate rather shorter posteriorly than Xerus.
P. 3 minute. In all the skulls seen, the upper cheekteeth are more or less
flat-crowned and well on the way towards complete simplification, but all are
much worn. The teeth are strongly hypsodont. One main outer re-entrant
fold is traceable in all teeth, with sometimes a short fold in front of it. The
teeth in extreme age seem to wear down to a simple ring-shape. Lower cheek-
teeth of all seen with one short outer and one long inner fold; or tending to
become completely simple at later development. The dentition appears to
indicate a nearer relationship w ith the Xerus section rather than w ith the Citellus
section.
D.3 of forefoot the longest. Claws of all digits of hindfoot, and the four main
digits of forefoot, excessively long, thick and powerful. Pollex very short, but
clawed. Sole thickly haired in those examined. Summer pelage rough, almost
bristly as in Xerus; winter pelage long, silky. Mammae 6 or 8 (Obolensky).
Fig. io6. Spermophilopsis leptodactyhs leptodactvlI'S, Lichtenstein.
B.M. No. 0.4.3.23, V; ■ li.
Fig. 107. Spermophilopsis leptodactylis leptodactvlis, Lichtenstein.
B.M. No. 9.4. 3. 23, . ; ■ li.
SPERMOl'IHLOI'SIS— SCIUROTAMIAS 425
Tail strongly reduced, not much longer than hindfoot. Cheek-pouches present
(\inogradov).
Forms seen: leptodactyltis.
The genus is revised by Obolensky, 1927, C.R. Acad. Sci. Leningrad, p. 188.
List of Named Forms
1. SPERMOPHILOPSIS LEPTODACTYLUS LEPTODACTi'LUS, Lichtenstein
1823. Eversmann. Reise, p. 119.
Karata, 140 versts north-west of Bokhara.
Synonym: turcomanus, Eichwald, Reise i, p. 305, 1834.
2. SPERMOPHILOPSIS LEPTODACTYLUS SCHUMAKOVI, Satunin
1908. Mitt. Kauk. Mus. p. 255.
Kushka, S. Transcaspia.
3. SPERMOPHILOPSIS BACTRIANUS, Scully
1888. Joum. .Asiat. Soc. Bengal, LVI, p. 70.
Khamiab, North Afghanistan.
Considered by Vinogradov, 1933, Rodents of U.S.S.R., as probably a sub-
species of leptodactylus.
Section F. T.\mias and Allies. Semiterrestrial Squirrels with narrow
rather flattened skull, relatively small postorbital process, cheek-pouches
present, digits 3 and 4 of manus about equal to each other in length (neither
constantly longer than the other), dentition of Sciurus type.
Genus 27. SCIUROTAML-VS, Miller
1901. SciUROT.VMi.'vs, Miller, Proc. Biol. Soc. Washington, XIV, p. 23.
1922. RVPESTES, Thomas, .Ann. Mag. Nat. Hist. 9, X, p. 398. {Rupestesforrestt, Thomas.)
Valid as a subgenus.
Type Species. — Sciurus dazidianus, Milne-Edwards.
Range. — China: Moupin, Szechuan, Kansu, Shensi, Shansi, Chihli: Yunnan.
Number of Forms. — Five.
Characters. — Skull much like Asiatic Tamias, hut infraorbital foramen not
abnormallv open, though more so than is usual in Sciurinae.
Tail thickly bushy, no colour pattern; and differing " in the direction of Sciurus
in the reduction of the capacity of the cheek-pouches." Skull long, relatively
narrow, with smooth braincase, much reduced postorbital process, and with
posterior portion of skull not depressed posteriorly, agreeing in this respect
with Tamias. but differing from Sciurus. Zygomatic plate, as in Tamias, short
and little tilted upwards, the ridges of its superior border weak. Infraorbital
foramen scarcely forming canal; masseter knob weak or absent. L'pper cheek-
teeth as in Tamias; P. 3 present, though vestigial. Lower cheekteeth not essen-
tially different from Tamias; palate normal.
426 SCIUROTAMIAS— TA.MIA.S
Tail about thrtc-quartcrs of head antl body length, bushy. D.3 and D.4
in manus subequal in length. Ilindtoot rather broad, with sole hairy; arrange-
ment of digits as in Taniias, with tendency for D.4 to be slightly longer than
D.3. Fur thick and soft.
Upper incisors rather short.
Kiipesles described as a full genus by Thomas is not distinguishable in
cranial and dental characters in anv way that could be considered as of generic
value. P. 3 is absent. The hindfoot is narrow, with naked sole, and an addi-
tional sole pad halfway between heel and digital pad at base of hallu.x. Fur and
digit arrangement as in normal Sciurotainias. Three pairs of mammae (Thomas).
Further specimens would be welcome to make the exact status of this species
clear; at the moment there are no characters which distinguish it as a genus
except the form of the hindfoot, which mav be very variable elsewhere within
a genus.
Forms seen: daridiainis, consohrinus, Jonesti.
List of Named F'orms
Subgenus Sciiirutainias, Miller
1. SCIUROTAMIAS DAVIDIANUS D.WIDIANUS, Milm--Ed«ards
1867. Rev. Mag. Zool. p. iq6.
X. China, mountains near Pekin.
2. SCIUROTAMIAS DAVIDIANUS TH.-\YKR1, U. Allen
1913. Mem. Mus. Comp. Zool. 40, p. 231.
Washan, W. Szechuan, China.
3. SCIUROTAMIAS DAVIDIANUS CONSOBRINUS, Milne-EdwarJs
1.S6S. Rcch. Hist. Nat. Mamni. p. 305.
Moupin, Szechuan, China.
4. SCIUROTAMIAS DAVIDIANUS OWSTONI. Allen
igog. Bull. Amer. Mus. Nat. Hist., XXVI, p. 42g.
Tai-Pa-Shiang Mts., Shen-Si, China.
Subgenus Riipcstcs, Thomas
:;. SCIUROTAMIAS FORRESTI, Thomas
ig22. Ann. Mag. Nat. Hist, g, X, p. 3g8.
Mekong-Yangtze Divide, 27 N., Yunnan, China.
Genus 28. TAMIAS, Illiger
iNii. Tamias, Illiger, Prod. Syst. Manim. et Avium, p. 83.
1880. EtiTAMiAS, Trouessart, Cat. IVIanim. Viv. et Koss. RuJentia, in Bui. .Sue. Etudes
Sci. d'Angers. 10, p. 86. (Sciiiriis striatus asiaticus, Gmelin.) \'alid as a subgenus.
ig2g. Neotami.^s, Howell, North .Amer. Fauna, No. 26, p. 52. (Eiitamias nierriami,
.Allen.) \'alid as a subgenus.
Type Species. — Sciurus striatus, Linnaeus.
TAMIAS
427
Range. — Holarctic: U.S.S.R., eastwards from River Dwina and Kama,
northwards from 58 N. in European Russia, also widely distri-
buted in wooded and wood-steppe districts of Siberia and Far East (\'ino-
jjradov); Altai, Ussuri; .Manchuria, Chihli, Korea, Shansi, Shensi, Kansu,
Szechuan; Japan, Kurile Islands; Yukon, Mackenzie, Ontario, Alberta, British
Fig.
"Yl-
108. TA^^.^s dorsalis dorsalis, Baird.
B.M. No. 88.9.24.1, o; X 2j.
Columbia, Washington, Oregon, Idaho, Montana, Wyoming, Wisconsin,
California, Lower California, Nevada, Utah, Arizona, Colorado, New Mexico,
Texas, New York, South Carolina; south into Northern Mexico.
NtMBER OF FoR.MS. — About eighty-one.
Remarks. — This genus is usually split into two, Tamias and Etitamias, the
former containing the type species only.
The subgenus Eulamias was originally proposed by Trouessart with the single
character that the premolars are ; instead of J as in typical Tamias. Merriam
in 1897 (Proc. Biol. Soc. Washington, XI, p. 1S9), gave Eutamias full generic
rank, stating, "it will be observed that the name Eutamias, proposed bv Troues-
sart in 1880 as a subgenus of Tamias is here adopted as a full genus. This is
428 TAMIAS
because ot tlic conviction tiiat tiie superficial resemblance betw een the two groups
is accidental parallelism, in no way indicative of affinity. In fact the two groups,
it my notion of their relationship is correct, had very different ancestors,
Tamias being an oflshoot of the ground-squirrels of the subgenus Ictidomys of
Allen, and Eiitainias of the subgenus Aniinospcrmophihis, Merriam." At the same
time he gives no characters which would separate the tw'o "genera."
Howell in 1929 (Revision of the Chipmunks, North .\mer. Fauna, No. 52,
p. I, 1929), erected a new subgenus for .American " Kiiliiiiiids," and keys the
three groups thus:
101). Tamias dorsalis dorsalis, Baird.
B.\I. No. SS.9.24.1. S; ■: zl.
Upper premolars two; dorsal stripes unequally spaced (median bordered
on either side by a much broader band). Genus T.amias
Upper premolars four; dorsal stripes equally spaced (all of approximately
equal width). Genus Eutamias
Anteorbital foramen suborbieular; postorbital processes broad at
base. Subgenus Eutamias
Anteorbital foramen narrowly oval; [lustorbital processes narrow at
base. Subgenus Ncotamius
This key convinces me that all these forms must be referred to one genus
only. The characters given to separate "Eutamias'' from Tamias are based
only on the absence or presence of the functionless premolar, and on the colour
pattern. If colour pattern is to be used as a generic character, it seems Citellus
suslicus will require a new name when compared with C. citellus, etc.
The Asiatic Chipmunk is intermediate between typical Tamias and the
TAMIAS 429
smaller American forms in many characters. Comparing Neotamias with
Eutamias, Howell, writing of the latter, states, "the ears are broad, rounded, of
medium height, much as in Tumias; postorhital broad at base, tapering to a
point, much as in Tamias; interorliital constriction slight, as in Tamias; upper
molariform toothrows slightly convergent posteriorly, as in Tamias." He also
states, "F.utamias of Asia resembles Tamias of North America and differs from
American Eutamias in a number of characters, notably the shape of the ante-
orbital foramen, the postorbital process, the breadth of the interorbital region,
the development of the lambdoidal crest, and the shape of the external ears.
On the other hand, American Eutamias agrees with the .Asiatic members of the
genus in the shape of the rostrum, the well-defined striations of the upper
incisors, the presence of the e.xtra peg-like premolar, and in the pattern of the
dorsal stripes."
It becomes clear that these forms agree in far too many essential characters
for the Eutamias group to be retained longer as a distinct genus.
Ch.^r.\cters. — (Subgenus Eutamias; Palaearctic). Skull lightly built,
narrow, and with no prominent ridges for muscle attachment,
except the upper border of the zygomatic plate, which is ridged superiorly,
though not tilted upwards as much as is usual owing to the skull being flatter
than in most Sciuridae. Postorbital process small and v.eak, though broader
than in subgenus iXeotamias. Bullae relativelv large. Palate broad, normal, not
continued far backwards; lachrymal not much enlarged. Infraorbital foramen
forming no canal, large, round and apparently well open enough to transmit a
strand of muscle. The part of the zygomatic plate behind it is narrow; a small
masseter knob is usually present. Mandible with angular portion somewhat
pulled inward.
Cheekteeth of Sciurus type; P. 3 present, but vestigial. The ridges and
depressions well marked originally, but evidently tending to wear down rather
early. The lower teeth are not unlike those of Citellus, but without the great
height of the cusps characteristic of normal members of that genus, and the
posterointernal cusp more developed ; the ridge connecting the two outer main
cusps weaker. Upper incisors with traces of many minute grooves, as in
Marmota.
Externally, tail relatively long, though rather shorter than the head and
body, as a rule; not densely bushy, though fully haired. In the manus, D.4 and
D.3 are usually roughly equal to each other in length. It has been stated
(Winge, Weber) that D.3 is the longest digit ; but out of two hundred and
twenty-seven skins examined (including .Asiatic and American species of the
genus), only tiftv-two had the middle digit slightly longer than the fourth; and
sometimes D.4 may be slightly the longer; in some skins of Asiatic Tamias, the
digit lengths vary in the two hands. Sole usually partly haired. Ilindfoot
with the outer digits shorter than D.3 and D.4, with a tendency for D.4 to be
slightly the longest; hallux rather reduced. The back typically with five black
stripes, bordering four white ones. There is some variation in colour pattern;
in (Neotamias) dorsalis, only one mid-dorsal stripe is clearly marked.
Neotamias, containing the American forms with a roughly similar colour
430 TAMIAS
pattern and P. 3 present, have the infraorbital foramen rather less open and less
rounded, but still of large size compared with most Sciuridae. The postorbital
process is narrower at the base, and lighter. The skull is less heavily built.
But as shown above, Asiatic Tamias connects these forms with the typical
subgenus. The baculum is said to differ from subgenus Eutamias.
Tamias s.s. has P. 3 absent, and there are only two white flank-stripes each
side, though the five black ones of the other forms are present. The rostrum is
less abruptly constricted near the base, and narrowing evenly from base to tip;
the upper incisors have the grooves shallow or absent. Further, Howell states
that the palate is relatively longer (but the differences in British Museum
material seems to amount to very little); and that the bullae are relatively
smaller. The tail is usually rather shorter. This subgenus is confined to
eastern North America.
Tamias and Eiitaiiiias contain a single species each. Ncotainias is divided
bv Howell into five specific groups:
The alpiinis group: small, skull 30-3-3 1'" in length; interorbital region broader
than in other small species; (coloration rather pale).
The minimus group: small-medium in size; hindfoot 26-35 mm.; skull length
28-7-34-.
The aniociiiis group: evidentlv not very clearly distinguishable from the iiiiiiimiis
group (hindfoot 29-5-35; skull length 3i-3-35-'i); "certain forms in the
two groups . . . inhabiting widely separated areas are so closely similar
both in external and cranial characters that many specimens are difficult
to identify without recourse to the locality label" (Howell).
The qiiadrivitiiitus group: size mediimi; hindfoot 32-36; skull larger than races
of miniums or amoenus, 34-5-36-8.
The toKiisendii group: large; hindfoot 34-39; skull 36-8-40-8.
Forms seen: alpinus, amoenus, "asiaticus" = sibiricus, horealis, hiilUri, calli-
peplus, cinereicollis, consohriniis, dorsalis, fratcr, hiiidsi, intercessor, inyoensis,
lineatus, lysteri, nierriami, minimus. iu\<;lectus, ochrogenxs, ordinalis, orientalis,
panamintinus, pricei, quadrimaculatiis, quiitiriTittatus, sencscens, senex, spcciosus,
striatus, townseiidi.
List of Named Forms
Subgenus Neotamias, Howell
alpinus (jroup
1. T.^ML^^S .-M.PINUS, AU-rriam
1893. Proc. Biol. Soc. Washington, VIIl, p. 137.
Bis Cottonwood Meadows, soutli of Mount Whitney, Tulare County,
California.
minimus Group
2. TAML^S MINIMl'.S MINIMtJ.S, Bachman
1S39. Joum. Acad. Nat. .Sci. Philadelphia, VIII, pt. i, p. 71.
Near CJreen River City, Sweetwater County, Wyoming.
TAMIAS 431
3. TAMIAS MINIMUS PICTUS, Allen
1890. Bull. Anier. Mus. Nat. Hist., Ill, p. 115.
Kclton, lioxelder County, Utah.
Synonym: mininuis melamirus, Merriam, 1890, N. A. Fauna, No. 4,
p. 22. Snake R., Blackfoot, Bingham County, Idaho.
4. TAMIAS MINIMUS GRISKSCENS, Howell
1925. Journ. Mamm. Baltimore, 6, p. 52.
Farmer, Douglas Count>', Washington.
5. TAMIAS MINIMUS CARYl, Merriam
1908. Proc. Biol. Soc. Washington, XXI, p. 143.
Medano Ranch, San Luis Valley, Costilla County, Colorado.
6. TAMIAS MINIMUS PALLIDUS, Allen
1874. Proc. Boston Soc. Nat. Hist., XVI, p. 289.
Camp Thome, near Glendive, Dawson County, Montana.
7. TAMIAS MINIMUS CACODEMUS, Carj-
1906. Proc. Biol. Soc. Washington, XIX, p. 89.
Sheep Mountain, Big Bad Lands, South Dakota; Fall River County.
8. TAMIAS MINIMUS CONFINIS, Howell
1925. Journ. Mamm. Baltimore, 6, p. 52.
Crescent Lake, Oneida County, Wisconsin.
(). TAMIAS MINIMUS CONSOBRINUS, .^llen
1890. Bull. Amer. Mus. Nat. Hist., Ill, p. 112.
Wasatch foothills, 18 miles east of Salt Lake City, Utah.
Synonym: clarus, Bailey, 1918, Proc. Biol. Soc. Washington, XXXI,
p. 31. Swan Lake Valley, Yellowstone National Park.
lectus, Allen, 1905, Mus. Brooklyn Inst. Arts & Sci.: Sci.
Bull. I, p. 117. Beaver Valley, Beaver Counn-, Utah.
10. TAMIAS MINIMUS OPERARIUS, Merriam
1905. Proc. Biol. Soc. Washington, XVIII, p. 164.
Gold Hill, Boulder County, Colorado.
11. TAMIAS MINIMUS .ATRISTRI.^TUS, Bailey
IQ13. Proc. Biol. Soc. Washington, XXVI, p. 129.
Penasco Creek, Sacramento Alountains, Lincoln County (12 miles east
of Cloudcroft), New Mexico.
12. TAMIAS MINIMUS ARIZONENSIS, Howell
1922. Joum. Mamm. Baltimore, 3, p. 178.
Prieto Plateau, Blue Range, Greenlee County, Arizona.
13. TAMIAS MINIMIS OREOCETES, Merriam
1897. Proc. Biol. Soc. Washington, XI, p. 207.
Summit Mountain. Flathead County, Montana.
14. TAMIAS .MINI.MUS BOREALIS, .-Mien
1877. Monogr. N. .Amer. Rodents, p. 793.
Fort Liard, Mackenzie, Canada.
Synonym: neglectus. .-Mien, 1890, Bull. Amer. Mus. Nat. Hist. Ill, p. 106.
Mouth of Montreal River, Lake Superior, Canada.
15. T.\M1AS MINI.MUS CANICEPS, Osgood
1900. Nonh Amer. Fauna, No. 19, p. 28.
Lake Lebargc. Yukon, Canada.
43:: TAMIAS
i6. TAMIAS MIMMUS JACKSOM, Houcll
1Q25. Jiiurn. Mamni. Baltimore, 6, p. 53.
Crescent Lake, Oneida County, Wisconsin.
17. TAMIAS MINIMUS SCRUTATOR, Hall & HatHild
1934. Univ. Calif. Publ. Zool. 40, p. 321.
Near Blanco Mountain, White Mountains, Mono County, California.
anioemis Group
iS. TAMIAS AMOKNUS AMOENUS, Allen
1890. Bull. Amer. Mus. Nat. Hist., Ill, p. 90.
Fort Klamath, Klamath Coimty, Oregon.
Synonyni: aniot'niis propi}itiiiiis, Anthony, 1913, Bull. .Amer. Mus. Nat.
Hist., XXXH, p. 6. Ironside, Malheur Coimty, Oregon.
iQ. TAMIAS AMOKNUS OCHRACEUS, Howell
1925. Journ. Mamni. Baltimore, 6, p. 54.
Studhorse Canyon, Siskiyou Mountains, California.
20. TAMIAS AMOENUS MONOENSIS, Grinnell & Storer
1916. Univ. Calif. Publ. Zool. 17, p. 3.
Warren Fork, Mono County, California.
21. TAMIAS AM(JENUS LUTEIVENTRIS, Allen
1890. Bull. Amer. Mus. Nat. Hist., Ill, p. loi.
Chief Mountain Lake (Waterton Lake), Alberta, 3 J miles north of
U.S. .A. -Canada boundary.
22. TAMIAS AMOENUS VALLICOLA. Howell
1922. Journ. Mamm. Baltimore, 3, p. 179.
Bass Creek, near Stevensville, Ravalli County, Montana.
23. TAMIAS AMOENUS CANTCAUDUS, Merriam
1903. Proc. Biol. Soc. Washington, XVI, p. 77.
Spokane, Spokane County, Washington.
24. TAMI,\S AMOENUS AFFINIS, Allen
1S90. Bull. .Amer. Mus. Nat. Hist., Ill, p. 103.
Ashcroft, British Columbia.
25. TAMIAS AMOENUS LUDIBUNDUS, Holhster
1911. Smiths. Misc. Coll., LVL no. 26, p. i.
Yellowhead Lake, British Columbia.
26. T.AMIAS AMOENUS FELIX, Rhoads
1895. Amer. Nat. 29, p. 941.
Church Mountain, Mount Baker Range, British Columbia (New
Westminster district).
27. TAMI.AS AMOENUS CAliRINUS. Mcrriam
1898. Proc. Acad. Nat. .Sci. Philadelphia, p. 352.
Near head of Soleduc Ri\er, Olympic Mountains. Clallam Coimty,
W'ashington.
2.S. TAMIAS PANAMINTINUS PANAMINTINUS, Merriiim
1893. Proc. Biol. Soc. W'ashington, VIII, p. 134.
Johnson Canyon, Panamint Mountains, Inyo County, California.
TAMIAS 433
29. TAMIAS I'ANAMINTINUS JUNIPERUS, Burt
1Q31. Joum. Mamm. Baltimore, 12, p. 298.
Charleston Mountains, Clark County, Nevada.
quadrivittatus Group
30. TAMIAS QUADRIVITTATUS QUADRIVITTATUS, Say
1823. Long's Exp. Rocky Mountains, 2, p. 45.
.Arkansas River, Colorado, about 26 miles below Canyon City, Pueblo
County.
Synonym : ijitadrivittaliis gracilis , Allen, 1 890, Bull. Amer. Mus. Nat. Hist.,
Ill, p. 99. San Pedro, Socorro County, New Mexico.
quadrivittatus animosus, Warren, 1909, Proc. Biol. Soc.
Washington, XXII, p. 105. Las Animas County,
Colorado; Irwin's Ranch.
31. TAMIAS QUADRIVITTATUS HOPIENSIS, Merriam
1905. Proc. Biol. Soc. Washington, XVIII, p. 165.
Keam Canyon, Navajo Count>-, Arizona.
32. TAMIAS QUADRIVITTATUS INYOENSIS, Merriam
1897. Proc. Biol. Soc. Washington, XI, p. 208.
White Mountains, Inyo County, California.
33. TAMIAS QUADRIVITTATUS FR.'^TER, Allen
1890. Bull. Amer. Mus. Nat. Hist., Ill, p. 88.
Donner, Placer County, California.
34. TAMIAS QUADRIVITTATUS SEQUOIENSIS, Howell
1922. Joum. Mamm. Baltimore, 3, p. 180.
Mineral King, east fork of Kaweah River, Tulare County, California.
35. TAMIAS QUADRIVITTATUS SPECIOSUS, Merriam
1890. Bull. Amer. Mus. Nat. Hist., Ill, p. 86.
Head of White Water Creek, San Bernardino Mountains, San Bernar-
dino County, California.
36. TAMIAS QUADRIVITTATUS NEVADENSIS, Burt
193 1. Joum. Mamm. Baltimore, 12, p. 299.
Sheep Mountains, Clark County, Nevada.
37. TAMIAS CALLIPEPLUS, Merriam
1893. Proc. Biol. Soc. Washington, VIII, p. 136.
Mount Pinos, Ventura County, California.
38. TAMIAS PALMERI, Merriam
1897. Proc. Biol. Soc. Washington, XI, p. 208.
Charleston Peak, Clark Countj', Nevada.
39. T.\MIAS ADSITUS, Allen
1905. Mus. Brooklyn .-Xrts & Sci.: Sci. Bull, i, p. 118.
Briggs Meadows, Beaver Mountains, Millard Count\-, Utah.
40. TAMIAS UMBRINUS, Allen
1890. Bull. .Amer. Mus. Nat. Hist.. III. p. 96.
Black Fork, Uinta Mountains, Utah.
28 — Living Rodents — 1
434 TAMIAS
41- TAMIAS RUFICAUDUS RUFICAUDUS, Howell
IQ20. Proc. Biol. Soc. Washinuton, XXXIII, p. gi.
Upper St. Mary Lake, Glacier County, Montana.
42. TAMIAS RUFICAUDUS SIMULANS, Howdl
1922. Journ. Mamrn. Baltimore, 3, p. 179.
Coeur d'Alene, Kootenai County, Idaho.
43- TAMIAS CINKREICOLLIS CINERFICOLI.IS, Allen
iSyo. Bull. Arner. Mus. Nat. Hist. Ill, p. 94.
San Francisco Mountain, Coconino County, .-Xrizona.
44- TAMIAS CINICREICOLLIS CINEREUS, Bailey
1913. Proc. Biol. .Soc. Washington, XXVI, p. 130.
Magdalena Mountains, Socorro County, New Mexico.
45. TAMEAS CINEREICOLLIS CANIPES, Bailey
1902. Proc. Biol. Soc. Washington, XV, p. 117.
Guadaloupe Mountains, El Paso County, Texas.
4'.- T.\MIAS BULLERI BULLERI, Allen
1SS9. Bull. Amer. Mus. Nat. Hist., II, p. 173.
Sierra de Valparaiso. Zacatecas, Mexico.
47. TAMIAS BULLI;R1 DURANGAE, Allen
1903. Bull. Amer. Mus. Nat. Hist., XIX, p. 594.
Arroyo de Bucy, N.-W. Durango, Mexico.
Synonym: nexus, Elliot, 1905, Proc. Biol. Soc. Washington, XVIII, p. 233.
Coyotes, Durango, Mexico.
4S. TAMIAS BULLERI SOLIVAGUS, Howell
1922. Journ. Mamm. Baltimore, 3, p. 179.
Sierra Guadalupe, Coahuila, Mexico.
townsendii Group
40- TAMI.-\S TOWNSENDII TOWNSENDII. Bachman
1839. Journ. Acad. Nat. Sci. Philadelphia, VIII, pt. i, p. 68.
Lower Columbia River, Oregon.
.Synonym: hindsi. Gray, 1S42, Ann. Mag. Nat. Hist.. X, p. 264. Fort
Vancouver, Washington.
toumsendii littoralis, Elliot, 1903, Field Columb. Mus. pub.
74, zool. ser. 3, p. 153. Marshfield, Oregon.
50. TAMIAS TOWNSENDII C(JOPERI, Baird
1855. Proc. Acad. Nat. Sci. Philadelphia, p. 334.
Klickitat Pass, Cascade Mountains, Washington; Skamania County.
51. TAMIAS TOWNSENDII OCHROGENYS, .Mernam
1897. Proc. Biol. Soc. Washington, XI, p. 195.
Mendocino, Mendocino C(.)unty, California.
52. TAMIAS TOWNSENDII SISKIYOU. Howell
1922. Journ. Mamm. Baltimore, 3, p. 180.
Near summit of White Mountain, Siskiyou County, California.
53. TAMIAS TOWNSENDII SENEX, Allen
1890. Bull. .^mer. Mus. Nat. Hist., Ill, p. 83.
Summit of Donner Pass, Placer County, California.
TAMIAS 435
54. TAMIAS TOVVNSF.NDII SONOMAK, Grinnell
1915. Univ. Calif. Publ. Zool. 12, p. 321.
Guemcville, Sonoma County, California.
55. TAMIAS ALLliNl, Howell
1922. Joum. Mamm. Baltimore, 3, p. 181.
Inverness, Marin County, California.
Synonym: hindsi, Merriam, Proc. Biol. Soc. Washington, XI, p. 194,
1897. Not of Gray.
56. TAMIAS QUADRIMACULATUS, Gray
1867. .\nn. Mag. Nat. Hist. 3, XX, p. 435.
East of Michigan Bluff, Placer County, California.
Synonym: macrorhabdotes , Merriam, 1886, Proc. Biol. Soc. Wash-
ington, III, p. 25. Blue Canyon, Placer County, California.
57. TAMIAS MERRIAMI MERRIAMI, Allen
i88g. Bull. .^mer. Mus. Nat. Hist., II. p. 176.
San Bernardino Mountains, San Bernardino County, California.
Synonym: merriami mariposae, Grinnell & Storer, 1916, Univ. Calif.
Publ. Zool. 17, p. 4. El Portal, Mariposa County,
California.
58. TA.MIAS MERRI.\MI PRICEI, Allen
1895. Bull. Amer. Mus. Nat. Hist., VII, p. 333.
Portola, San Mateo County, California.
59. T.^MIAS MERRIAMI KERNE.NSIS, Grinnell & Storer
1916. Univ. Calif. Publ. Zool. 17, p. 5.
Fay Creek, Kern County, California.
60. TAMI.\S MERRIAMI OBSCURUS, Allen
1890. Bull. .Amer. Mus. Nat. Hist., Ill, p. 70.
San Pedro Martir Mountains, Lower California, Mexico.
61. T.AMIAS MERRI.-\MI MERIDIONWLIS, Nelson & Goldman
1909. Proc. Biol. Soc. Washington, XXII, p. 23.
.\guaje de San Esteban, north-west of San Ignacio, Lower Cahfomia.
62. T.\.MIAS DORS.\LIS DORSALIS. Baird
1855. Proc. Acad. Nat. Sci. Philadelphia, VII, p. 332.
Fort Webster, Gila River, Grant County, New Mexico.
SraonjTn: canescens, Allen, 1904, Bull. Amer. Mus. Nat. Hist., XX,
p. 208. Guanacevi, Durango, Mexico.
63. T.\MIAS DORS.\LIS UT.\HENSIS. .Merriam
1897. Proc. Biol. Soc. Washington, XI, p. 210.
Ogden, Weber County, Utah.
64. TAM1.\S DORSALIS GRINNELLI. Burt
1931. Joum. Mamm. Baltimore, 12, p. 300.
Sheep Mountains, Clark County, Nevada.
Subgenus Eutamias, Trouessart
65. TAMIAS SIBIRICUS SIBIRICUS, La.xmann
1769. Sibirisches Briefe, p. 69.
Barnaul, Siberia.
Synon>Tn: asiaticus, Gmelin, 1788, Syst. Nat. p. 150. Barnaul, Siberia.
pallasi, Baird, 1856, Ann. Rep. p. 55 (fide Trouessart).
43^' TAMIAS
b6. TAMIAS S1BIRICL:S ALTAICUS, Hollister
1912. Proc. Biol. Soc. Washington, XX, p. 183.
Tapucha, Altai Mountains, Siberia.
67. TAMIAS SIBIRICUS ORIKNTALIS. Bonhotc
iSyg. .■Xnn. Mat. Nat. Hist. 7, IV, p. 3S5.
Sungatscha River, Upper Ussuri, East Siberia.
bS. TAMI.\S SIBIRICUS UTHENSIS, Pallas
i<S3i. Zoograph. Rosao-Asiat. i, p. i8g.
Uda River, N.-E. Siberia.
6i). TAMIAS SIBIRICUS ORDINALIS, Thomas
IQ08. Abstr. Proc. Zool. Soc. London, p. 44; Proc. Zool. Soc. London, p. 968.
Vu-lin-fu, Shensi, China.
70. TAMIAS SIBIRICUS UMBR(.)SUS, Howell
1927. Joum. Washington Acad. Sci., XVH, p. 80.
140 miles south of Lanchowfu, vicinity of .Archuen, Minshan Moun-
tains, Kansu, China.
71. T.^MIAS SIBIRICUS INTERCESSOR, Thomas
iqoS. Abstr. Proc. Zool. Soc. London, p. 44; Proc. Zool. Soc. London, p. 969.
Ning-\vu-fu, Shensi, China.
12. TAMI.AS SIBIRICUS ALBOGULARIS, Allen
1909. Bull. Amer. Mus. Nat. Hist., XXVI, p. 429.
Tai-pa-shiang, Shensi, China.
73. TAMIAS SIBIRICUS SENESCENS, Miller
1898. Proc. Acad. Nat. Sci. Philadelphia, p. 349.
15 miles west of Peking, China.
74. TAMIAS SIBIRICUS OKAD.AE, Kuroda
1932. Journ. Mamm. Baltimore, 13, p. 58.
Mt. Chachanupuri, Kunashiri Island, South Kurile Islands.
75. TAMIAS SIBIRICUS LINEATUS, Siebold
1824. Spic. Faun. Japon in Diss. Hist. Nat. Japon, p. 13.
Japan.
76. TAMIAS SIBIRICUS JACUTENSIS, Ognt-v
1935. Wiss. Ber. Moskaeur Staats. Univ. 4, p. 93.
Yakutsk, East Siberia.
Subgenus Tamias, Illiger
77. TAMIAS STRIATUS STRIATUS, Linnaeus
175S. Syst. Nat. Ed. 10. i, p. 64.
Unknown; probably near Savannah River, South Carolina.
Synonym: americaiius, Gmelin, Syst. Nat. i, p. 150, 1788.
78. TAMIAS STRIATUS FISHERI, Howell
1925. Journ. Mamm. Baltimore, 6, p. 51.
Merritts Corners, Ossining, New York.
79. TAMIAS STRIATUS LYSTERI, Richardson
1829. Fauna Boreali-Americana, i, p. 181,
Penetanguishene, Georgian Bay, Ontario, Canada.
TAMIAS— CITELLUS 437
So. TAMIAS STRIATUS GRISKUS, Mearns
1891. Bull. Amer. Mus. Nat. Hist., Ill, p. 231.
Fort Spelling, Hennepin County, Minnesota.
81. TAMIAS STRIATUS VKNISTL'S, Bangs
1896. Proc. Biol. Soc. Washington, X, p. 137.
Stilvvell, Adair County, Oklahoma.
Section G. Marmota Section. Ground-squirrels with the following
characters. Palate not produced far behind last molars, and lachrymal
not enlarged. Infraorbital foramen forming a canal, postorbital process
usually well developed, and D.3 in manus so far as seen constantly
longer than D.4. Tendency present for upper main cheekteeth to
become much constricted on inner side; usually strongly hypsodont.
Genus 29. CITELLUS, Oken
1816. CiTELLUS, Oken, Lehrbuch der Zoologie, pt. 3, vol. 2, p. 842.
1817. Anison"vx, Rafinesque, .^mer. Monthly Mag. 2, i, 45. (Anisonyx brachyurus,
Rafinesque- Arctomys coliinibuiuiis, Ord.)
1825. Spermophius, F. Cuvier, Dents des Mamm. 160, 161, pi. LV, p. 255.
1844. CoLOBOTis, Brandt, Bull. CI. Phys. Math. Acad. Imp. Sci. St. Petersb. II, no. 23,
24, pp. 365, 366. {Spermophilus fulvus, Lichtenstein.)
1844. OrosPERMOPHiLts, Brandt, Bull. CI. Phys. Math. Acad. Imp. Sci. St. Petersb.
vol. 2, p. 379. (Sciurus grammiirus. Say.) Valid as a subgenus.
1877. ICTIDOMVS, .Allen, Monogr. Nth. Amer. Rodentia, p. 821. {Sciurus tridecem-
lineatiis, Mitchill.) Valid as a subgenus.
1893. Xerospermophilus, Merriam, Proc. Biol. Soc. Washington, VII, p. 27. (Sper-
mophilus mohavensis, Merriam.) Valid as a subgenus.
1892. Ammospermophills, Merriam, Proc. Biol. Soc. Washington, VII, p. 27. (Tamias
leucurus, Merriam.) Valid as a subgenus.
1 90 1. Callospermophills, Merriam, Proc. Biol. Soc. Washington, XI, p. i8g. {Sciurus
lateralis. Say.) Valid as a Subgenus.
1907. IcTlDOMoiDEs, Meams, Mamm. Mex. Boundary, U.S. pt. i, p. 328. {Sciurus
mexicanus, Erxleben.)
1927. Urocitellus, Obolensky, C. R. Acad. Leningrad, p. 192. {Spermophilus evers-
manni, Brandt.)
1938. NoTOCiTELLUS, Howell, North Amer. Fauna, No. 56, p. 162. {Spermophilus
ammlatus, Audubon & Bachman.) Valid as a subgenus.
1938. PoLiociTELLUS, Howell, North Amer. Fauna, No. 56, p. 133. {Arctomys
franklinii, Sabine.) Valid as a subgenus.
Type Species. — Mus citellus, Linnaeus.
Range. — Holarctic: Silesia, Bohemia, Galicia, Hungary, Rumania, Bulgaria,
Greece, Turkey, Asia Minor, Caucasus ; the whole of southern
European Russia, north to Rivers Kama and Oka ; most of South-western
Siberia (Kazakstan area), south into Persia, north to Ural River, Irtish River,
Semipalatinsk, etc.; Altai; Zungaria; Transbaikalia, East Siberia (Okhotsk,
Verhoiansk, .\nadyr districts, etc.); Mongolia, Manchuria, Shansi, Shensi,
Kansu; Alaska, Mackenzie, Saskatchewan, Alberta, British Columbia; Washing-
ton, Oregon, Idaho, Montana, Wyoming, Minnesota, California, Nevada, Utah,
Arizona, New Mexico, Texas, southwards to Central Mexico.
Fig. iio. Citellus citellis citellus, Linnaeus.
B.M. No. 8.9.10.7, ?; X 2.
Fig. III. Citellis citellus citellus, Linnaeus.
B.M. No. 8.9.10.7, ?; X 2.
Number of Forms.-
Remarks.-
CITELLUS
-About a hundred and fortv-four.
439
KiG.
112. CiTELLUS CITELLUS.
Cheekteeth: X 5.
-The American forms of this genus have been recently revised
by Howell, 193S (North Amer. Fauna, No. 56). This revision
has long been needed, and completes the list of all large Nearctic genera revised.
The genus is made much larger by Howell, by the inclusion of the groups
hitherto regarded as distinct genera, Callospermophilus, Ammospermophiliis and
Otospertnopliilus. I had endeavoured to keep these three names standing as
valid genera, and except in the case of _
Ammospermophilus had found it no easy
matter. Thev are all evidently much better
regarded as subgenera of Citelbis, being at
once distinguishable from Tamias, with
which they were formerly classed, by the
character of the infraorbital foramen which
is quite normal, whereas in Tamias it forms
no canal and is relatively ven,^ large. But
their inclusion in Citelltis makes it desir-
able to make some of the other Sciurine
genera, as, for instance, Xerus, wider than
they are at the moment currently accepted.
Howell very rightly states that in many
cases the genera recognized to-day are little
better than specific groups.
Obolensky revised the Palaearctic Citelltis in 1927 (C. R. Acad. Sci. Lenin-
grad, p. 188). These were arranged in three subgenera, Citellus {citelltis and
suslicus groups as here understood), soles haired, tail one-fifth to a third head
and body length, interorbital region relatively narrow; Colobotis (fulvus and
pygmaeus groups as here understood), with bare soles, tail length and inter-
orbital region about as in Citelltis s.s.; and Urocitellus {eversmanni group), with
haired soles, tail a third to a half head and body length, interorbital region
relatively broad. Howell regards these groups as belonging to Citellus s.s. The
dentition of all Palaearctic forms examined agrees with Howell's Citelltis s.s.;
and I am of the opinion that Obolensky's names must be disregarded; the
Palaearctic Citellus form with the North American t)-pical subgenus of Howell
a natural group as far as I have had occasion to examine.
Ch.\r.\CTERS. — {Citellus s.s.). Upper cheekteeth noticeably constricted on
inner side, so that the teeth appear roughlv three-sided
instead of rounded or nearly square as in most Sciurinae. Skull with slender
postorbital processes, which always appear well developed in the typical group.
Certain constriction is usually apparent in front of the postorbital processes;
braincase not flattened. Palate extending ver\- slightly behind toothrows, which
are not or scarcely convergent posteriorly. Infraorbital foramen often rather
well open; masseter knob at its lower border prominent. Zygomatic plate with
upper border strongly ridged, e.xtending upwards far forwards, and with upper
part much narrowed. Sagittal ridge rarely tornied by parietal ridges in material
440 CITELLUS
examined ; if present, never as conspicuous or heavy as in Marmota or Cynomys,
in British Museum material, though a skull ot parrxii figured hy Howell appears
to have a rather conspicuous ridge present. Mandihle with angular portion as
a rule strongly pulled inwards.
Upper cheekteeth with inner side strongly hypsodont; P. 3 very little
reduced, and functional; main teeth with three strong outer cusps present;
the ridges from the second and third cusps high, converging to the inner cusp.
The anterior ridge, from the front cusp, is short. The main ridges separated
by deep depressions. M.3 with only one well-marked main ridge, the posterior
part of the tooth flatter, and with less defined elements. Lower teeth vv'ith two
very high anterior cusps, the anterointernal one the highest, the two cusps
joined hy a ridge; and a moderately high posteroexternal cusp; the three main
cusps of the teeth surrounding a deep depression. Posterointernal cusp vestigial
or absent. M.3, as in the upper series, tends to be the largest tooth. The two
outer main cusps in the lower teeth joined by a short ridge.
External characters somewhat variable, but always in the whole genus so
far as I have seen characterized by the fact that the digits are arranged in the
manner of terrestrial forms, D.3 being in the manus the main digit. Hindfoot
with the two outer digits, particularly the hallux, shorter than the three central
ones. Pollex much reduced. Ear small. Tail always considerably shorter than
head and body; from about half this measurement to little longer than hindfoot.
Claws as a rule prominent; particularly in such forms as parryii; never so
developed as Spermophilopsis.
Mammae 10, 12 or 14 in Palaearctic species (Obolensky), 10, 12 or 8 accord-
ing to Howell in Nearctic species, cheek-pouches present.
In the Palaearctic, I provisionally recognize five species groups; tne evers-
manni (characters indicated above, p. 439); the citclliis group {characters as
above); the stislicus group, like citellus but with a well-marked spotted colour-
pattern, this much more developed than in anv other Palaearctic species seen;
the pygmacus group, like citellus group but with naked soles; and the fiilrus
group, which has a much heavier skull and dentition than is normal in the genus.
Eight subgenera are recognized by Howell as occurring in America. He
keys them as follows:
Molars relatively hypsodont; parastyle ridge on M.i and M.2 joining
protocone with an abrupt change of direction.
Metaloph of P. 4 continuous. Citellus s.s.
Metaloph of P. 4 not continuous. Ictidomys
Molars relatively brachyodont, parastyle ridge on M.i and M.2 rising
evenly to join protocone, without abrupt change of direction.
Anterior upper premolar simple, less than one-fourth size of P. 4.
Upper incisors relatively stout, distinctly recurved.
Braincase rounded on upper surface.
Supraorbital foramen open. Otospermophilus
Supraorbital foramen closed. Notocitfllus
CITELLUS 441
Braincase flattened on upper surface. Ammospermophilus
Upper incisors relatively slender, not distinctly recurved.
Postorbital process long and slender; rostrum longer.
Callospermoph I LUS
Postorbital process short and stout; rostrum shorter.
X EROSPERMOPHl LUS
Anterior upper premolar more than a quarter size of P. 4,
hearing two cusps and a functional cutting edge. Poliocitellus
Subgenus IcTlDOMY-S. This includes trulecemlineatus, also according to Howell
the species mexicanus and spilosoma. The braincase is relatively
narrower than Citellus; P. 3 is relatively much smaller, and the upper
incisors are said to be shorter and stouter.
C. tridecemlineatus has the most specialized colour-pattern of any
living Squirrel. The postorbital process appears to be much smaller
than in Citellus s.s. C. mexicanus has a spotted colour-pattern; and
spilosoma, of which very few are in the British Museum, is described as
spotted, though apparently in some forms the spots may be faint. Tail
60-80 per cent head and body length (Howell).
Subgenus Poliocitellus. This is based on franklinii only; a plain non-striped
species which in many respects appears to me to resemble the
tridecemlineatus group (e.g. cranial characters). P. 3 is more reduced
than in Citellus s.s. The dentition is, however, not like that of tvpical
Citellus, but more transitionary to the Tamias or Sciurus-\\k& type found
in all the remaining subgenera. The tail is more than half head and
body length. The zygomata are less expanded than in normal
Citellus.
Subgenus Otospermophilus. Rather large forms, in which the postorbital
process is relatively large, and a sagittal crest present in all adult
skulls seen. The teeth have no tendency to the internal constriction
of typical Citellus (in the upper series). Zygomatic plate as in normal
Citellus. Externally more Sciurus-\\ke; ears moderately large ; tail about
two-thirds head and body length, bushy ; upper incisors stout. The
cheekteeth are not very different from those of Sciurus. P. 3 is small.
Subgenus Notocitellus. Based on aimulatus, heretofore placed in Oto-
spermophilus. Not seen, not represented in the British Museum.
Tail described as more than two-thirds head and bodv length.
Supraorbital foramen closed (differing from Otospermophilus in this
character). P. 3 relatively small.
Subgenus Ammospermophilus. This group seems to me to be the most
distinct of all Howell's subgenera. Bullae large and inflated in all
skulls examined. Palate usually ending in long and conspicuous spinous
process. Dental characters not essentially different from Sciurus or
Tamias. Externally very reminiscent of the .\frican Eu.xerus, except
442 CITELLUS
tor the much shorter tail, which is said to be carried over the hack
when the animal is rimnins:!, and is about half head and body length.
Fur sometimes slightly bristly. .\ white stripe on each flank. Ears
small. Postorbital process slender; incisors stout; infraorbital foramen
narrower than in normal Cilellus (agreeing with Otospermophilus).
Subgenus Xerospermophilus. This is based on mohavensis and the tereti-
coudiis group. Very few have been seen. The claws are described as
long, sharp ; the sole is heavily haired ; ears very low ; tail 40-60 per
cent head and body length. Molars (said to be) near Otospennopliihis.
Subgenus C.\llospermophilus. Postorbital process rather prominent; cranial
characters near Otospermop/iiliis, except that sagittal crest, so far as seen,
is usuallv absent, the masseter knob moderate or small, the postorbital
process rather lighter; upper incisors relatively more slender. Tail
usually more than half head and body length. Ear low. Form Tamias-
like, with the white flank-stripes bordered usually by four black ones;
but no mid-dorsal stripe.
Forms seen: heeclievi, bcldiiigi, bcriiardimis, bunmcri, hrevicauda, caniithersi,
chrvsodeinis, citellus, cinenisccns. concolur, dauricus, douglasi, elegaus, ervthro-
genxs, eversiiianni, fislieri, jrankliiiii, fidvus. gradojevici, grtimmiirus, harrisi,
herbicola, jacutensis, kodiakciisis, lateralis, leiiciinis, macrouriis, mexicanus, mollis,
mongolicus, mugosaricus, tielsoni, urcgoinis, oxianiis, pallidiis, parthianus, parrvii,
peninsulac, planicola, pvgmaeus, ramosiis, richardsoni, rufescens, spilosoma, suslicus,
tereticatidus, touiistiuli, tridecemlineatus, umbratiis, rariegatiis, vimdtis, tvortmani,
xanthopixmmis.
The differences in the four groups recognized by Howell in American
Citellus s.s. refer mainly apparently to size and colour. C. washitigtoiii is
described as a spotted species, in colour near the Old World guttatus (suslicus).
Members of the parrvii group are, according to Howell's key, also spotted or
mottled.
In his revision some new forms are described, which, however, are not listed
here as, in all other genera, forms described to 1936 only are included.
List of N.^med Forms
Subgenus Citellus, Oken
Palaearctic Species
incertae sedis
1. CITELLUS FL.WESCENS, Pallas
177S. Nov. Spec. Quadr. Glir. Ord. p. 122.
Locality not known.
Not seen; not allocated to group
2. CITELLUS .^TRIC.-\PILL.\. Orlov^
1927. Mat. Contra fauna L. Volga, i, p. 92.
Lower Volga, U.S.S.R.
^ This appears to be preoccupied b>' alncapltlus, Bryant. No. 108 of this list. I therefore
rename it buiominatus.
CITELLUS 443
fulvus Group
3. CITELLUS FULVUS FULVUS, Lichtcnstein
1823. Eversmann Reise. p. 119.
River Kuwandzaliur, cast of Mugodsharski Mountains, north of Sea of
Aral, Siberia.
4. CITELLUS FULVUS OXIANUS, Thomas
191S. Ann. Mag. Nat. Hist. 8, XV, p. 422.
50 miles south-west of Bokhara.
5. CITELLUS FULVUS PARTHIANUS, Thomas
1915. Ann. Mag. Nat. Hist. 8, XV, p. 423.
Meshed, N.-E. Persia.
6. CITELLUS FULVUS CONCOLOR, Geoffroy
1834. Belanger, Voyages, p. 151.
Sultenia, near Kazvin, N.-W. Persia.
7. CITELLUS FULVUS HYPOLEUCUS, Satunin
1909. Ann. Mus. Zool. St. Petersb. 14, p. i.
Kutshun, Central Persia.
pygmaeus Group
8. CITELLUS RUFESCENS RUFESCENS, Keyserling & Blasius
1840. Wirbelth. Europas, p. 42.
Ural \Iountains, Russia.
Synonym: undulatus, Eversmann, 1840, Bull. Nat. Moscou, p. 35 (fide
Trouessart).
9. CITELLUS RUFESCENS ERYTHROGENi^S, Brandt
1841. Bull. .Acad. Sci. St. Petersb. p. 41.
Altai.
10. CITELLUS RUFESCENS UNG.'^E, Martino
1923. Ann. Mus. Zool. Petrograd, 24, p. 23.
Kirghiz Steppes.
11. CITELLUS PALLIDICAUD.V Satunin
1902. .\nn. Mus. Zool. St. Petersb., VII, p. 5.
Mongolian Altai; Cholmu Noor, Ullyn-Bulyk, River Baidarak.
12. CITELLUS PYGMAEUS PYGMAEUS, Pallas
1778. Nov. Spec. Quadr. Glir. Ord. p. 122.
Bet\veen Emba and Ural Rivers.
13. CITELLUS PYGMAEUS BREVICAUDA, Brandt
1843. Bull. Acad. Sci. St. Petersb., I, 23, p. 364.
Altai.
Synonym: intermedius, Brandt, I.e. p. 378 (fide Trouessart).
14. CITELLUS PYGM.\EUS MUGOSARICUS, Lichtenstein
1823. Eversmann Reise, p. 19.
Mugodshary Mountains, Kirghisia.
15. CITELLUS PYGMAEUS HERBICOLA, Martino
1916. .Ann. Mus. Zool. Petrograd, 21, pp. 269-301.
North Kirghisia.
444 CITELLUS
.6. CITKI.LLS PVGMAEUS SEPTKNTRIONALIS. Ubolcnsky
1927. C. R. Acad. Sci. Leningrad, p. 190.
Ferapontovka, Samara.
17. CITELLUS PYGNL^EL'S ORLOVL N™ Name
To replace pallidiis, Orlov & Feniuk
1927, Mat. Contr. Faun. Lower Volga, i, p. 63. Not of Allen. 1S77.
Kalmouk Steppes, near Astrakhan, South Russia.
18. CITELLU.S PYGMAEUS PLANICOLA, Satunin
1 90S. Mitt. Kauk. Mas. p. 46.
Karanogai Steppes, Kizljar, Caucasus.
iq. CITELLUS PYGMAEUS MUSICUS, M6n6tri6s
1S32. Catal. Rais. p. 21.
Elburz, Caucasus.
20. CITELLUS PYGMAEUS SATUNINI, Sviridinko
1922. Bull. Mus. Georgie, i, p. 69.
Daghestan, Caucasus.
21. CITELLUS PYGMAEUS BRAUNERI, Martino
1920. Notes Crimea Soc. Naturalists, 3.
Ecatermoslav, Crimea, South Russia.
22. CITELLUS PYGMAEUS NIKOLSKII, Heptner
1934. Folia Zool. Hydrob. 6, p. 20.
Aral Lake Shore.
23. CITELLUS PYGMAEUS KAZAKSTANICUS, Goodwin
1935. .Amer. Mus. Nov. 769, p. 5.
Kazakstan, Central Asia; Tuz Bulak, 150 miles north of Kizilorda
(Perovsk).
24. CITELLUS PYGMAEUS CARRUTHERSI, Thomas
1912. Ann. Mag. Nat. Hist. 8, IX, p. 393.
Barlik Mountains, N.-W. Dzungaria.
25. CITELLUS RELICTUS, Kashkarov
1923. Trans. Sci. Soc. Turkest. p. 185.
Talass-Alatau, Namanghan, Ferghana.
citellus Group
26. CITELLUS CITELLUS CITELLUS, Linnaeus
1766. Syst. Nat. I, 1 2th I-:d. p. 80.
Austria.
27. CITELLLS CITELLUS GRADOJEVICI, Martino
1929. Journ. Mamm. Baltimore, 10, p. 76.
Djerdjclija, Macedonia.
2S, CITELLUS CITELLUS ISTRICUS, Calmescu
1934. Zeitschr. fur Saugetierk, 9, p. 106.
E. Rumania; Munteni.
2Q. CITELLUS XANTHOPRYMNUS X.ANTHOPRYMNUS, Bennett
18^5. Proc. Zool. .Soc. London, p. 90.
Erzerum, Asia Minor. This species is probably not more than a sub-
species uf C^. citellus.
CITELLUS 445
30. CITELLUS XANTHOPRYMNUS SCHMIDTI, Satunin
1908. Mitt. Kauk. Mus. IV, p. 28.
Transcaucasia.
31. CITELLUS .JiLASCHANICUS AL.J\SCHANICUS, Buchner
1888. Wiss. Res. Przewalski Central-.-\sien Reisen: Zool. Th. I, Saugeth., p. 11.
South Alashan, Mongolia.
32. CITELLUS ALASCHANICUS DILUTUS, Formozov
1927. C. R. .Acad. Leningrad, p. 192.
Ikhe-Bogdo, Mongolian Altai.
33. CITELLUS ALASCFL-SiNICUS OBSCURUS. Buchnei
1888. Wiss. Res. Przewalski Central-.Asien Reisen: Zool. Th. I, Saugeth., p. 17.
Kansu, China.
34. CITELLUS ALASCR^VNICUS SICCUS, G. M. Allen
1925. Amer. Mus. Nov. 163, p. 3.
Shansi, China, lo miles west of Taijoianfu.
35. CITELLUS D.A.URICUS DAURICUS, Brandt
1843. Bull. Acad. St. Petersb. p. 379.
South Transbaikalia.
36. CITELLUS DAURICUS MONGOLICUS. Milnc-Edwards
1867. .Ann. Sci. Nat. p. 376.
Manchuria; Pekin.
37. CITELLUS DAURICUS UMBRATUS. Thomas
1908. Proc. Zool. Soc. London, p. 970.
Taboul, 100 miles north-west of Kalgan, Mongolia.
38. CITELLUS DAURICUS RAMOSUS, Thomas
1909. Ann. Mag. Nat. Hist. 8, IV, p. 501.
Fan Chia Tun, Kirin Province, Manchuria.
suslicus Group
39. CITELLUS SUSLICUS SUSLICUS, Guldenstaedt
1770. N. Comm. Ac. Sc. Petr. xiv, pt. i, p. 389.
Voronezh, Russia.
S>Tion>Tn; guttatuliis, Schinz, 1895, Synop. Mamm., II, p. 70.
leiicopictus, DonndorfT, 1792, Zool. Beytrage, i, p. 486.
40. CITELLUS SUSLICUS GUTT.ATUS, Pallas
1770. N. Comm. Ac. Sc. Petr. xiv, pt. i, p. 506.
Locality not known.
41. CITELLUS SUSLICUS AVERINI, Migutin
1927. Proc. Nat. Hist. Soc. Kharhov, p. 50, pt. 2.
Russka Lesonia, 18 km. north of Kharkov, Russia.
42. CITELLUS SUSLICUS MERIDIOCCIDENTALIS, Migutin
1927. Proc. Nat. Hist. Soc. Kharkov, 50, pt. 2.
Environs of Odessa, Russia.
everstnanni Group
43. CITELLUS EVERSMANNI EVERS.\L\NN1. Brandt
1841. Bull. Acad. St. Petersb. p. 43.
.Altai Mountains.
Synonym: allaictis, Eversmann, 1841, .Add. Zoog. R. .Asiat. fasc. 2, p. i.
446 CITELLUS
44. CITELLUS EVF.RS^LA^'^'I STRAMINEUS, Obolcnsky
1927. C. R. Acad. Sci. Leningrad, p. 192.
N.-W. Mongolia.
4:;. CITELLUS EVERSMAXM LEUCOSTICTUS, Brandt
1S43. Bull. Acad. Sci. St. Petersb., 11, p. 379.
Okhotsk River, N.-E. Siberia.
46. CITELLUS EVERSMANNI BUXTONI, Allen
1901. Bull. Amer. Mus. Nat. Hist., XIX, p. 139.
Gichiga, west coast Okhotsk ,Sea = efersniaiiiii leucostictus according to
Chaworth-Musters, Ann. Mag. Nat. Hist., 1934, 10, XIII, p. 555.
47. CITELLUS EVERSMANNI TRANSBAICALICUS, Obolensky
1927. C. R. Acad. Sci. Leningrad, p. T92.
Lake Ivan, Transbaikalia.
48. CITELLUS EVERSM.^NNl JACUTENSIS, Brandt
1844. Bull. Ac. Sci. St. Petersb., II, 23-24, p. 378.
Yakutsk, Siberia.
4.,. CITELLUS EVERSM.A.NNI STEJNEGERI, Allen
1903. Bull. Amer. Mus. Nat. Hist., XIX, p. 142.
Kamtchatka.
Nearctic Forms. Revised by Howell, 1938.
townsendii Group
50. CITELLUS TOWNSENDII TOWNSENDII, Bachman
1839. Joum. .A.cad. Nat. Sci. Philadelphia, VIII, p. 61.
Columbia River, about 300 miles above its mouth, near mouth of \\ alia
Walla River, Washmgton.
Synonvm: mollis yakimensts, Merriam, 1898, Proc. Biol. Soc. Washing-
ton XII, p. 70. Mabton, Yakima County, Washington.
51. CITELLUS TOWNSENDII CANUS, Merriam
1898. Proc. Biol. Soc. Washington, XII, p. 7°-
Antelope, Wasco County, Oregon.
52. CITELLUS TOWNSENDII VIGILIS, Merriam
1913. Proc. Biol. Soc. Washington, XXVI, p. 137.
Vale, Malheur County, Oregon.
53. CITELLUS TOWNSENDII MOLLIS, Kennicott
1863. Proc. Acad. Nat. Sci. Philadelphia, p. 157.
Camp Flovd, near Fairlield, Wasatch County, Utah.
Synonvm: stephensi, Merriam, 1898, Proc. Biol. Soc. Washington, XII,
p. 69. Esmeralda County (Queen Station), Nevada.
nashoensis. Merriam, 1913, Proc. Biol. Soc. Washington,
XXVI, p. 138. Carson Valley, Douglas County, Nevada.
lemodon, Merriam, 1913, Proc. Biol. Soc. Washington,
XXVI, p. 136. Murphy, Owyhee Countv-, Idaho.
54. CITELLUS TOWNSENDII .ARTEMISL\E, Merriam
1913. Proc. Biol. Soc. Washington, XXVI, p. 137.
Birch Creek, Fremont County, Idaho.
Synonym: pissimus. Merriam, 1913, Proc. Biol. Soc. Washington,
XXVI, p. 138. Big Lost River, Fremont County, Idaho.
CITELLUS 447
55. CITELLUS IDAHOENSIS. Merriam
1913. Proc. Biol. Soc. Washington, XXVI, p. 135.
Payette, Payette County, Idaho.
washingtoni Group
56. CITELLUS WASHINGTONI, Howell
(1938). North Amer. Fauna, No. 56, p. 69.
Touchet, Walla Walla County, Washington.
Synonym: tojviisemli. Dice, 1919, Joum. Mamm. Baltimore i, p. 18, not
toioiseiidi, Bachman.
57. CITELLUS BRUNNEUS, Howell
1928. Proc. Biol, Soc. Washington, XLI, p. 211.
New Meadows, Adams County, Idaho.
richardsoni Group
58. CITELLUS RICH.ARDSONI RICHARDSONI, Sabine
1822. Trans. Linn. Soc, XIII, p. 589.
Carlton House, Saskatchewan.
5t;, CITELLUS RICHARDSONI ELEGANS, Kennicott
1863. Proc. Acad. Nat. Sci. Philadelphia, p. 158.
Fort Bridger, Uinta County, Wyoming.
60. CITELLUS RICHARDSONI NEVADENSIS, Howell
1928. Proc. Biol. Soc. Washington, XLI, p. 211.
Paradise, Humboldt County, Nevada.
61. CITELLUS ARMATUS, Kennicott
1863. Proc. Acad. Nat. Sci. Philadelphia, p. 158.
Near Fort Bridger, Uinta County, Wyoming.
62. CITELLUS BELDINGI BELDINGI, Merriam
i888. Ann. New York Acad. Sci. 4, p. 317.
Dormer, Placer County, California.
63. CITELLUS BELDINGI OREGONUS, Merriam
1898. Proc. Biol. Soc. Washington, XII, p. 69.
Swan Lake Valley, Klamath County, Oregon.
parryii Group
54. CITELLUS COLUMBIANUS COLUMBI.ANUS, Ord
1815. Guthrie's Geography, 2nd .-Vmer. Ed. vol. 2, p. 292.
Camas prairie, between forks of Clearwater and Kooskooskie, Lincoln
County, Idaho.
Synonym: brachiura, Rafinesque, 1817, .\mer. Monthly Mag., 2, p. 45.
erythrogluteius, Richardson, 1829, Fauna Boreali-Americana
I, p. 161.
columbiamis alberlae, Allen, 1903, Bull. Amer. Mus. Nat.
Hist., XIX, p. 537. Canadian National Park. .-Mberta.
65. CITELLUS COLUMBIANIS RUFICAUDUS, Howell
1928. Proc. Biol. Soc. Washington, XLI, p. 212.
Wallowa Lake, Oregon.
448 CITELLUS
(.6. CITELLUS PARRYIl PARRYIL Richardson
1S25. Appendix to Parry's Second Voyage, p. 316.
Five Hawser Bay, Lyon Inlet, Melville Peninsula, Franklin, Canada.
Synonvni: vmpitra. True, 1S85, Proc. U.S. Nat. Mus., VII, p. 594.
phaeogiiallnis, Richardson, 1829, Fauna Boreali-.Aniericana,
p. 161. Hudson Bay.
keniucotti. Ross, 1861, Canadian Nat. &; Geol. 6, p. 434.
Mackenzie, near Fort Good Hope.
h7. CITELLUS PARRYIl BARROWENSIS, Merriam
1900. Proc. Washington Acad. Sci., II, p. 19.
Point Barrow, Alaska.
Synonym: heringensis, Merriam, 1900, Proc. Acad. Sci. Washington, II,
p. 20. Cape Lisburne, ."Maska.
68. CITELLUS PARRYIl PLESIUS, Osgood
1900. North Amer. Fauna, No. 19, p. 29.
Bennett City, head of Lake Bennett, British Columbia.
6q. CITELLUS PARUYII ABLUSUS, Osgood
1903. Proc. Biol. Soc. Washington, XVI. p. 25.
Nushagak, .Alaska.
Synonym; stoiiei, .Allen. 1903, Bull. .Amer. Mus. Nat. Hist., XIX, p. 537.
Stevana Flats, Alaska Peninsula, Alaska.
70. CITELLUS PARRYIl NEBULICOLA, Osgood
1903. Proc. Biol. Soc. Washington, XVI, p. 26.
Nagai Island, Shumagin Islands, Alaska.
71. CITELLUS P.ARRYII LYRATUS, Hall S: Gilmore
1933. Univ. Calif. Publ. Zool. 28, p. 396.
St. Lawrence Island, Behring Sea.
72. CITELLUS KODIACENSIS, Allen
1874. Proc. Boston Soc. Nat. Hist. 16, p. 292.
Kodiak Island, Alaska.
73. CITELLUS OSGOODI, Merriam
1900. Proc. Washington Acad. Sci., II, p. 18.
F'ort Yukon, Alaska.
Subgenus Ictidomvs, Allen
tridecemlineatus Group
74. CITELLUS TRIDECEMLINEATUS TRIDECEMI.INE.^TUS, Mitchill
1821. Med. Repos. n.s. vol. 6 (21), p. 248.
Central Minnesota.
Svnonym; Iwodi, Sabine, 1822, Trans. Linn. Soc, XIII, p. 590.
75. CITELLIS TRIDECEMLINEATUS TEXENSIS, Merriam
1S98. Proc. Biol. Soc. Washington, XII, p. 71.
Gainesville, Cooke County, Texas.
Synonym: hadius. Bangs, 1899, Proc. New Engl. Club, i, p. i. Stotes-
bury, Missouri.
76. CITELLUS TRIDECEMLINE.ATUS ARENICOLA, Howell
1928. Proc. Biol. Soc. Washington, XLI, p. 213.
Pendennis, Kansas.
CITELLUS 449
77. CITELLUS TRIDECEMLINEATUS PALLIDUS, Allen
1877. Monogr. North Amer. Rodt-nts, p. 872.
Plains of Lower Yellowstone River, Montana.
Synonym: olivaceus, Allen, 1895, Bull. Amer. Mus. Nat. Hist. VII,
P- 337- Custer, Custer County, South Dakota.
78. CITELLUS TRIDECEMLINEATUS ALLENI, Merriam
1898. Proc. Biol. Soc. WashinRton, XII, p. 71.
Bighorn Mountains, Washakie County, Wyoming.
79. CITELLUS TRIDECEMLINE.^TUS HOLLISTERI, Bailey
1913. Proc. Biol. Soc. Washington, XXVI, p. 131.
Elk Valley, Sacramento Mountains, Lincoln County, New Mexico.
So. CITELLUS TRIDECEMLINEATUS MONTICOLA, Howell
1928. Proc. Biol. Soc. Washington, XLI, p. 214.
Marsh Lake, White Mountains, Arizona.
81. CITELLUS TRIDECEMLINEATUS P.'^RVUS, Allen
1895. Bull. Amer. Mus. Nat. Hist. VII, p. 337.
Uncompahgre Indian Reser\ation, N.-E. Utah.
82. CITELLUS MEXICANUS MEXICANUS, Erxleben
1777. Syst. Regn. -Anim. vol. i, p. 428.
South Central Mexico.
83. CITELLUS MEXICANUS PARVIDENS, Meams
1896. Prelim, diagn. new Mamm. Mex. Border U.S., p. i, (Reprint: Proc. U.S. Nat.
Mus. XVIII, p. 443).
Fort Clark, Kinney County, Texas.
spilosoma Group
84. CITELLUS SPILOSOMA SPILOSOMA, Brandt
1833. Proc. Zool. Soc. London, p. 40.
N. Mexico and extreme W. Texas.
85. CITELLUS SPILOSOMA PALLESCENS, Howell
1928. Proc. Biol. Soc. Washington, XLI, p. 212.
La Ventura, Coahuila, Mexico.
86. CITELLUS SPILOSOMA C.ANESCENS, Merriam
1890. North. .Amer. Fauna, No. 4, p. 38.
Wilcox, Cochise County, .Arizona.
Synonym: macrospilotiis, Merriam, 1890, North. Amer. Fauna, No 4,
p. 38. Oracle, Pinal County, Arizona.
aretis, Bailey, 1902, Proc. Biol. Soc. Washington, XV,
p. 118. El Paso, Texas.
87. CITELLUS SPILOSOMA ^LMOR, Merriam
1890. North .Amer. Fauna, No. 4, p. 39.
Albuquerque, Bernalillo County, New Mexico.
Synonym: marginatus . Bailey, 1902, Proc. Biol. Soc. Washington. XV,
p. 118. Alpine, Brewster County, Texas.
88. CITELLUS SPILOSOMA ANNECTENS, Merriam
1893. Proc. Biol. Soc. Washington, VIII, p. 132.
Padre Island, Cameron County, Texas.
29 — Living Rodents — I
450 CITELLUS
S<). CITELLUS SPILOSOALA PRATENSIfi. Merriam
iSgo. North Amcr. Fauna, No. 3, p. 55.
Pine Plateau at north foot of San Francisco Mountain, Coconino
County, Arizona.
Synonym ; obsidiamis, Merriam, 1890, North Amer. Fauna, No. 3, p. 56.
(North-east of San Francisco Mountain, Coconino
County, Arizona.)
go. CITELLUS SPILOSOMA CRYPTOSPILOTUS, Merriam
1890. North Amer. Fauna, No. 3, p. 57.
Tenebito Wash, Painted Desert, Coconino County, Arizona.
91. CITELLUS SPILOSOMA OBSOLETUS, Kennicott
1863. Proc. Acad. Nat. Sci. Philadelphia, p. 157.
Extreme W. Nebraska.
92. CITELLUS PEROTENSIS, Merriam
1893. Proc. Biol. Soc. Washington, VIII, p. 131.
Perote, Vera Cruz, Mexico.
Subgenus Poliocitelliis, Howell
93. CITELLUS KRANKLINII, Sahine
1822. Trans. Linn. Soc. XIII, p. 587.
Vicinity of Carlton House, Saskatchewan, Canada.
Subgenus Otosperniophilus, Hrandt
94. CITELLUS VARIEGATUS VARIEGATUS, Erxlebcn
1777. Syst. Regn. Anim. i, p. 421.
South Central Mexico.
Synonym: buccatus, Lichtenstein, Abh. k. Akad. Wiss. Berlin, 1S27
(1830), p. 117.
niaaiirus, Bennett. 1833, Proc. Zool. Sue, London, p. 41.
Mexico.
flS. CITELLUS VARIEG.^TUS RUPESTRIS, Allen
1903. Bull. Amer. Mus. Nat. Hist. XIX, p. 595-
Rio Sestin, N.-W. Durango, Mexico.
96. CITELLUS VARIEGATUS COUCHII, Baird
1855. Proc. Acad. Nat. Sci. Philadelphia, VII, p. 332.
Santa Catarina, Nuevo Leon, Mexico.
97. CITELLUS VARIEGATUS BUCKLEYI, Slack
1861. Proc. Acad. Nat. Sci. Philadelphia, p. 314.
Packsaddle Mountain, Llano County, Texas.
98. CITELLUS VARIEGATUS GR.'VMMURUS, Say
1S23. Long's Exp. Rocky Mountains, p. 72.
Purgatory River, near mouth of Chacuaco Creek, Colorado, Las Animas
County.
Synonym: juglans, Bailey, 1913, Proc. Biol. Soc. Washington, XXVI,
p. 131. Glenwood, Rio San Francisco, Socorro County,
New Mexico.
99. CITELLUS VARIEGATUS TULAROSAE, Benson
1932. Univ. Calif. Publ. Zool. 38, p. 335.
New Mexico: French's Ranch, 12 miles north-west of Carrizozo,
Lincoln County.
CITELLUS 451
100. CITELLUS VARIEGATUS UTAH, Merriam
1903. Proc. Biol. Soc. Washington, XVI, p. 77.
Foot of Wasatch Mountains, near Ogden, Weber Count\-, Utah.
loi. CITELLUS BEECHEYI lilCF.CHEYI, Richardson
1829. Fauna Boreali-Americana, i, p. 170.
Neighbourhood of San Francisco and Monterey, California.
102. CITELLUS BEECHEYI DOUGLASI, Richardson
1829. Fauna Boreali-.A.mericana, i, p. 172.
Banks of Columbia River, Oregon.
103. CITELLUS BEECHEYI FISHERI, Merriam
1893. Proc. Biol. Soc. Washington, VIII, p. 133.
South Fork, Kern River, Kern County, 3 miles above Onyx, California.
104. CITELLUS BEECHEYI PARVULUS, Howell
1931. Joum. Mamm. Baltimore, 12, p. 160.
Shepherd Canyon, Argus Mountains, California.
105. CITELLUS BEECHEYI NUDIPES, Huey
1931. Trans. S. Diego Soc. Nat. Hist. 7, p. 18.
Hanson Laguna, Sierra Guarez : Lower California.
106. CITELLUS BEECHEYI RUPINARUM, Huey
1931. Trans. S. Diego Soc. Nat. Hist. 7, p. 17.
Catavina : Lower California.
107. CITELLUS BEECHEYI NESIOTICUS, Elliot
1904. Field Columb. Mus. publ. 90, zool. ser. vol. 3, p. 263.
Santa Catalina Island, Santa Barbara Islands, California.
108. CITELLUS ATRICAPILLUS, Bryant
1889. Proc. Calif. Acad. Sci. ser. 2, vol. 2, p. 26.
Comondu, Lower California, Mexico.
Subgenus Notocitellus, Howell
109. CITELLUS ANNULATUS ANNULATUS, Audubon & Bachman
1842. Joum. .'\cad. Nat. Sci. Philadelphia, 8, pt. 2, p. 319.
Unknown ; probably W. Mexico.
no. CITELLUS ANNULATUS GOLDMANI, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 69.
Santiago, Nayarit, Mexico.
111. CITELLUS ADOCETUS, .Merriam
1903. Proc. Biol. Soc. Washington, XVI, p. 79.
La Salada, 40 miles south of Uruapan, Mexico (Michoacan).
Subgenus Ammospermophilus, Merriam
112. CITELLUS H.\RRISII H.ARRISII, .Audubon & Bachman
1854. Quadr. N. .Amer., vol. 3, p. 267.
Unknown; probably S.-W. Arizona.
113- CITELLUS H.ARRISII SAXICOLA, Mcams
1896. Prelim, diagn. Mamm. Mex. border U.S., p. 2. Reprint, Proc. U.S. Nat. Mus.
iS, p. 444, 1896.
Tinajas Atlas, Gila Mountains, Yuma County, Arizona.
452 CITELLUS
114. CITELLUS LELX'URUS LEUCURUS, Mcrriam
18S9. North Amer. Fauna, No. 2, p. 20.
San Gorgonio Pass, Riverside County, California.
Synonym: finulus, Elliott, 1903, Field. Columb. Mus. publ. 87, zool.
ser. 241. Keeler, Inyo County, California.
115. CITELLUS LEUCURUS TERSU.S, Goldman
1929. Joum. Washington Acad. Sci. 19, p. 435.
Arizona: Prospect Valley, Grand Canyon, Hualpai Indian Reser\'ation.
116. CITELLUS LEUCURUS CINNAMOMEUS, iMurriam
iSgo. North Amer. Fauna, No. 3, p. 52.
Echo Cliffs, Painted Desert, Coconino County, Arizona.
117. CITELLUS LEUCURUS PENNIPES, Howell
1931. Journ. Mamm. Baltimore, 12, p. 162.
Grand Junction, Colorado.
118. CITELLUS LEUCURUS PENINSULAE, .-Xllen
1893. Bull. Amer. Mus. Nat. Hist. V, p. 197.
San Telmo, Lower California, Mexico.
IK). CITELLUS LEUCURUS C.ANFIELDAE, Huey
1Q29. Trans. San Diego Soc. Nat. Hist. 5, p. 243.
Punta Prieta, Lower California.
120. CITELLUS LEUCURUS EXTIMLIS, Ntlsou & Goldman
1929. Journ. Wash. Acad. Sci. 19, p. 281.
Lower California; Saccaton, 15 miles north of Cape San Lucas.
121. CITELLUS INTERPRES, Merriam
1890. North Amer. Fauna, No. 4, p. 21.
El Paso, El Paso County, Texas.
122. CITELLUS INSULARIS, Nelson & Goldm.an
1909. Proc. Biol. Soc. Washington. XXII, p. 24.
Esperitu Santo Island, Gulf of California, Mexico.
123. CITELLUS NELSONI, Merriam
1893. Proc. Biol. Soc. Washington, VIII, p. 129.
Tipton, San Joaquin Valley, California, Tulare County.
Synonym; amplus, Taylor, 1916, Univ. Calif. Publ. Zool. 17, p. 15.
Twenty miles south of Los Banos, Merced County,
California.
Subgenus Xerospennophiliis, Merriam
124. CITELLUS MOHAVENSIS. Merriam
18S9. North Amer. Fauna, No. 2, p. 15.
-Mohave River, San Bernardino County, California.
izs- CITELLUS TERETICAUDUS TERET1C.\UDUS, Baird
1S57. Mamm. N. Amer., p. 315.
Old Fort Yuma, Imperial County, California.
Synonym: zocifcrans, Yiuey. 1927, Proc. Biol. Soc. Washington, XXXIX,
p. 29. San Felipe, Lower California.
cremononnis. Elliot, 1903, Field Columb. Mus., publ. 87,
3, p. 243. Furnace Creek, Death Valley, Inyo County,
California.
CITELLUS 453
126. CITELLUS TERETICAUDUS NEGLECTUS, Merriam
1889. North .\mer. Fauna, Xo. 2, p. 17.
Dolan's Spring, Mohave County, Arizona.
Synonym: sonoriensis. Ward, 1891, Amer. Nat. 25, p. 158. Hermosillo,
Sonora, Mexico.
arizonae, Grinnell, 1918, Proc. BioL Soc.Washington, XXXI,
105. Tempe, Maricopa County, Arizona.
127. CITELLUS TERETICAUDUS CHLORUS, Elliot
1903. Field Mus. Columb. Publ. 87, zool. ser. 3, p. 242.
Palm Springs, Riverside County, California.
128. CITELLUS TERETICAUDUS APRICUS, Huey
1927. Trans. S. Diego Soc. Nat. Hist. 5, p. 85.
Lower California, Valle de la Trinidad.
Subgenus Callospermophilus, Merriam
I2Q. CITELLUS L.ATERALIS LATERALIS, Say
1823. Long's Exp. Rocky Mountains, 2, p. 46.
Arkansas River, below Canyon City, Pueblo County, Colorado.
130. CITELLUS LATERALIS WORTMANI, Allen
1895. Bull. Amer. Mus. Nat. Hist. VII, p. 335.
Kinney Ranch, Bitter Creek, Wyoming (Sweetwater County).
131. CITELLUS L.\TERALIS ARIZONENSIS, Bailey
1913. Proc. Biol. Soc. Washington, XXVI, p. 130.
Arizona, San Francisco Mountain.
132. CITELLUS LATERALIS CARYI, Howell
1917. Proc. Biol. Soc. Washington, XXX, p. 105.
Seven miles south of Fremont Peak, Wind River Mountains, Fremont,
Wyoming.
133. CITELLUS LATER,\LIS CINERASCENS, Merriam
1890. North .\mer. Fauna, No. 4, p. 20.
Helena, Lewis and Clarke County, Montana.
134. CITELLUS LATER.ALIS TESCORUM, Hollister
191 1. Smiths. Misc. Coll. LVI, no. 26, p. 2.
Head of Moose Pass Branch of Smoky River, Alberta, Canada.
135. CITELLUS LATER.ALIS CASTAXURUS, Merriam
1890. North Amer. Fauna, No. 4, p. 19.
Park City, Wasatch Mountains, Summit County, Utah.
136. CITELLUS LATER,ALIS CHRYSODEIRUS, Merriam
1890. North Amer. Fauna, Xo. 4, p. 91.
Fort Klamath, Klamath County, Oregon.
137. CITELLUS LATER.\LIS CONNECTENS, Howell
'93I- Joum. Mamm. Baltimore, 12, p. 161.
Homestead, Oregon.
138. CITELLUS L.\TER.\LIS TREPIDUS, Taylor
19:0. Univ. Calif. Pub. Zool. 5, p. 283.
Head of Big Creek, Pine Forest Mountains, Humboldt County, Xevada.
Synonym: perpallidiis, Grinnel], 1918, Univ. Calif. Pub. Zool. 17, p. 429.
Big Prospector Meadow, White Mountains, Mono
Counrv', California.
454 CITELLUS— MARMOTA
Uy. CITELLUS LATERALIS CERTUS, Goldman
1921. Joum. Mamni. Baltimore, 2, p. 232.
Charleston Peak, Clark Count>', Nevada.
140. CITELLUS LATER.'iLIS BERNARDINUS, Merriam
1898. Science, n.s., 8, p. 782.
San Bernardino Count>', California; .San Bernardino Peak.
Synonym: brciicniidiis, Merriam, 1893, Proc. Biol. Soc. Washington
V'lII, p. 134. Not of Brandt, 1844.
141- CITELLUS LATER-^LIS AIITRATUS, Howell
1931. Joum. Mamni. Baltimore, 12, p. 161.
South Yolla Bolly Mountain, California.
142. CITELLUS LATERALIS TRINITATIS, Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. 126.
Trinitv Mountains, Humboldt Countv, California (east of Hoopa
Valley).
143. CITELLUS SATURATUS, Rhoads
1895. Proc. Acad. Nat. Sci. Philadelphia, p. 43.
Lake Keechelus, Kittitas County, Washington.
144. CITELLUS MADRENSIS, Merriam
igoi. Proc. Washington Acad. Sci. Ill, p. 363.
Sierra Madre, Chihuahua, Mexico, near Guadalupe y Calvo.
Genus 30. MARMOTA, Blumenbach
1779. M.\RMOT.A, Blumenbach, Handb. Naturgesch, i, p. 79.
1780. Arctomvs, Schreber, Saugthiere, pis. CCVII-CCXI, ibid, text, IV, 72 1-743, 1782.
1922. Marmotops, Pocock, Proc. Zool. Soc. London, p. 1200. {M. mona.\, Linnaeus).
Type Species. — Mas inarmota, Linnaeus.
R.\NGE. — Holarctic : Alps (France, Switzerland, North Italy) and Car-
pathians; Poland, North Rumania (Bukowina); European Russia
(steppes of Rivers Don, Donez, Middle and Lower Volga, Mid LVal), northern
Kazakstan; Fergana, Pamir, Semirechyia, Altai, Tomsk; Transbaikalia;
Verhoyansk district, Anadyr region, and Kamtchatka (Russian localities quoted
by Vinogradov); Tibet, Chinese Turkestan, Kansu, northern Mongolia,
Manchuria; Afghanistan, Kashmir; Nepal, Sikkim, to Yunnan.
The greater part of Canada and the United States, from Alaska to Labrador
and the Atlantic coast of U.S.A.; south to California, New Mexico, and northern
Oklahoma, and Alabama. (Distribution maps of Nearctic species are published
by Howell, and in Anthony, Field Book North American Mammals, 1928.)
Number of Forms. — About fifty.
Ch.'\racters. — Skull much more powerfully ridged for muscle attachment
than in Citellus, and size becoming very large, largest of
family, head and body up to 620 mm. Postorbital process very thick and
heavy; little sign of interorbital constriction; parietal ridges usually join to form
MARMOTA 455
a sharp sagittal crest near hinder part of postorbital process. Frontals de-
pressed. Occiput strong, prominent. Infraorbital foramen wider below, but
not well open; masseter knob appears less produced than is usual in Cilellus;
upper border of zygomatic plate well ridged, relatively narrow. Jugal, as usual
in the family, long and extending to lachrymal. Palate not narrowed posteriorly.
Mandible as a rule less angular than in Citelhis or Cynomys. P. 3 very little
reduced, functional. Upper cheekteeth like those of Citellus, but rather less
constricted internally; strongly hypsodont, particularly on the inner side.
Cusps and ridges high, depressions deep, as in Citellus; often the third depres-
sion, at the back of the main teeth, wears out, like Citellus in this feature, but
unlike the few Cynomys examined. I\1.3 the largest tooth, with its posterior
elements more or less obliterated as a rule. Lower teeth like those of Citellus,
the posterointernal cusp not developed. Incisors, both upper and lower, usually
with traces of several faint grooves. Cheek-pouches (said to be) rudimentary or
absent. In the above notes ''Citellus'" refers to Citellus s.s.
Form thickset; tail less than or rarely exceeding a third head and body
length except in caudata group, in which it approaches half this measurement,
and perhaps in ciiligata group. Ear short. Hindfoot rather broad, with digits
arranged as in other terrestrial genera. Forefoot with D.3 the main digit; poUex
rudimentary; or absent in the type species. On this account Pocock restricted
the genus to the type species, and erected " Marmotops" for the others; but the
presence or absence of a minute and functionless digit of this type is of no
importance, and an examination of the skeleton of the manus of M. marmota
and M. caudata representing "Marmotops" presents very httle essential ditfer-
ence. Claws usually well developed, powerful. Mammae 10 in flaviventris,
caligata, marmota, bobak; 8 in monax.
It must be added that Marmotops is recognized as a subgenus by Howell,
'938-
The American species were revised by Howell in 191 5 (North Amer. Fauna,
no. 37). Three specific groups are recognized: the monax group ("Wood-
chucks": mammae 8, sagittal crest according to Howell weaker, less developed;
general appearance and coloration distinct at a glance from all others judging
by material examined); the flaviventris group (Yellow-bellied Marmots; of
western U.S.A.); and the caligata group (dark thick-furred types from Alaska,
extreme west Canada and adjacent parts of U.S.A. ; also from Xorth-east
Siberia; becoming rather larger than the above; apparently darkest of genus in
general coloration, and tail apparently tending to be rather longer). In both the
two latter groups, the mammae are 10.
In flaviventris group, the ear is stated to be smaller than in monax group,
and the tail is relatively longer. The monax group ranges right across Canada,
and in much of the eastern U.S..-\. Good distribution maps of the three groups
are published by Howell, and in Anthony, 1928, Field Book North American
Mammals.
The Palacarctic species are not yet revised. Other than the Siberian repre-
sentatives of the caligata group, I provisionally recognize three groups ; marmota
group (.\lps; tail approximatelv 27 per cent head and body length; general
Fig. 113. Marmota mabmota, Linnaeus.
B.M. No. 7.1. 1. 195 bis; •, i.
Fig. 1 14. Marmota marmota, Linnaeus.
B.M. No. 7.1. 1. 1 95 bis; ■ i.
MARMOTA
457
appearance as regards coloration at once distinguishable from all other species;
mammae lo) ; caiidata group (tail longest of genus, about 45 per cent head and
body length; rather large; thick-furred) includes caiidota (yellowish, with
conspicuous black mid-dorsal area noticeably
contrasting with sides), aurea, from which little-
dalei seems not more than racially distinct,
dichrous and stirlingi, differing from each other
in minor colour distinctions, but all clearly
separable from caudata.
M. bobak group: provisionally including all
other Palaearctic forms examined; tail normallv
strongly reduced, about a quarter length head
and body (or slightly more); coloration principally
light, often more or less unicolor; and typically
with short fur. Includes bobak, the related but
larger liinmlayaiia, sibirica (evidently differing in
colour), and centralis, a thicker-furred form, , .
1 • 1 • , , e L u ■ Marmota MARMOTA, Lmnaeus.
which IS now regarded as a race ol baibaana; cheekteeth- x 2
robusta is, I think, a race of himalayana.
Forms seen: aurea, baibacma, bobak, caligata, camtschatica, caudata, centralis,
cliftoiii, dichrous, Jlazinus, flaviventer, himalayana, littledalei, marmota, motiax,
okanagana, robusta, sibirica, stirlingi.
Fig. 115
List of Named Forms
monax Group
1. MARMOTA MONAX MOX.VC, Linnaeus
1758. Syst. Nat., loth Ed., vol. i, p. 60.
Maryland.
2. MARMOTA MONAX RIFESCEXS, Howell
1914. Proc. Biol. Soc. Washington, XXVII, p. 13.
Elk River, Minnesota, Sherburne County.
:?. MARMOTA MONAX PREBLORUM, Howell
1914. Proc. Biol. Soc. Washington, XXVII, p. 14.
Wilmington, Middlesex Countj', Massachusetts.
4. NURMOTA MONAX IGN.WA, Bangs
1899. Proc. New England Zool. Club, i, p. 13.
Black Bay, Strait of Belle Isle, Labrador,
.i. M.\RMOTA .MON.A.X CANADENSIS, Erxleben
1777. Syst. Regn. Anim. i, p. 363.
Quebec, Canada.
SynonjTii: empeira, Pallas, Nov. Sp. Quadr. Glir. Ord., p. 75, 1778.
6. MARMOTA MON.AX PETRENSIS, Howell
1915. North .^mer. Fauna, No. 37, p. 33.
Revelstoke, British Columbia, Canada.
7. MARMOTA MONAX OCHRACEA, Swarth
191 1. Univ. Calif. Publ. Zool. 7, p. 203.
Forty-mile Creek, .Alaska.
45S MARMOTA
S. MARMOTA MONAX BUNKERI, Black
■ 935- Journ. Mamm. Baltimore, i6, p. 319.
Lawrenci', Douglas County, Kansas.
flavivetitris Group
g. MARMOTA FLAVIVENTRIS FLAVIVENTRIS. Audubon & Bachman
1 84 1. Proc. Acad. Nat. Sci. Philadelphia, p. gg.
Mount Hood, Oregon.
10. NtARMOTA FLAVIVENTRIS A\-ARA, Bancs
1899. Proc. New England Zool. Club, i, p. 68.
Okanagan, British Columbia, Canada.
11. MARMOTA FLAVIVENTRI.S SIERRAE, Howull
1915. North Amcr. Fauna, No. 37, p. 43.
Head of Kern River, Mount Whitney, California, Tulare County.
12. MARMOTA FL-^WIVENTRIS FORTIROSTRIS, Grinnell
192 1. Univ. Calif. Publ. Zool. 21, p. 242.
McAfee Meadow, White Mountains, Mono County, California.
13. MARMOTA FL.WIVENTRIS PARVULA, Houtll
1915. Proc. Biol. Soc. Washington, XXVH, p. 14.
Jefferson, Nye County, Nevada.
14. MARMOTA FL.WIVENTRIS ENGELHARDTI, Allen
1905. Mus. Brooklyn Inst. Arts & .Sci. Science, Bull. 1, p. 120.
Briggs Meadows, Beaver Range Mountains, Beaver County, Utah.
15. MARMOTA FLAVIVENTRIS NOSOPHOR.-\, Howell
1914. Proc. Biol. Soc. Washington, XXVH, p. 15.
Willow Creek, Montana, Ravalli County, 7 miles east of Corvallis.
16. MARMOTA FL.A.VIVENTRIS DACOTA, .Merriam
1889. North .^mer. Fauna, No. 2, p. 8.
Custer, Black Hills, Custer County, South Dakota.
17. MARMOTA FLAVIVENTRIS LUTEOLA, Howell
1914. Proc. Biol. Soc. Washington, XXVH, p. 15.
Woods Post OHice, Medicine Bow Mountams, .Albany County,
Wyoming.
18. MARMOTA FL.A.VIVENTRIS CAMRIONI, Fiijnins
1915. Proc. Biol. Soc. Washington, XXVIII, p. 147.
Eight miles north of Higho, Jackson County, Colorado.
If). MARMOTA FLAVIVENTRIS WARRENI, Howell
1914. Proc. Biol. Soc. Washington, XXVH, p. 16.
Crested Butte, Gunnison County, Colorado.
20. MARM(.)TA FLAVIVENTRIS OBSCURA, Howell
1914. Proc. Biol. Soc. Washington, XXVH, p. 16.
Wheeler Peak, Taos County, New Me.xico; 5 miles south of Twining.
21. MARMOTA FL.AVIVENTRIS NOTIOROS, Warren
1934. Journ. Mamm. Baltimore, 15, p. 62.
Marion Lake, West Mountains, Custer County, Colorado.
MARMOTA 459
caligata Group
22. MARMOTA CALIGATA CALIGATA, Eschscholtz
1829. Zool. Atlas, pt. 2, p. I, p!. 6.
Near Bristol Bay, Alaska.
Synonym : (?) pruiiwsus, Gmelin, 1788, Syst. Nat. i , p. 144. Regarded as
unidentifiable by Howell, 191 5.
23. MARMOTA CALIGATA VIGILIS. Heller
1909. Univ. Calif. Publ. Zool., 5, p. 248.
West shore of Glacier Bay, Alaska.
24. MARMOTA CALIGATA SHICLDONI, Howell
1914. Proc. Biol. Soc. Washington, XXVII, p. 18.
Montague Island, Alaska.
25. MARMOTA CALIGATA OXYTONA, Hollister
1914. Science, n.s., vol. 39, p. 251.
Moose Pass Branch, Smoky River, Alberta, Canada.
Synonym: sibila, Hollister, 1912, Smiths. Misc. Coll. LVI, 35, p. i (not
of Wolf, 1808).
26. MARMOTA CALIGATA OKANAGANA, King
1836. Narr. Journ. Shores Arctic Ocean, vol. 2, p. 236.
Gold Range, British Columbia, Canada.
27. MARMOTA CALIGATA NIVARIA, Howell
1914. Proc. Biol. Soc. Washington, XXVII, p. 17.
Mountains near Upper St. Mary's Lake, Teton County, Montana.
28. M.\RMOTA C.'^LIGATA CASCADENSIS, Howell
1914. Proc. Biol. Soc. Washington, XXVII, p. 17.
Mount Rainier, Pierce County, Washington.
29. MARMOTA CALIGATA R.'^CEYI, Anderson
1932. Bull. Nat. Mus. Canada, 70, p. 112.
British Columbia; Itcha Mountains, Chilcotin Plateau.
30. MARMOTA CALIGATA BROWERI, Hall & Gilmore
1934. Canad. Field Nat., 48, p. 57.
North Alaska ; Point Lay, Arctic coast.
31. MARMOTA OLYMPUS, Merriam
1898. Proc. Acad. Nat. Sci. Philadelphia, p. 352.
Head of Soleduc River, Olympic Mountains, Clallam County, Wash-
ington.
32. MARMOTA VANCOUVERENSIS, Swarth
191 1. Univ. Calif. Publ. Zool. 7, p. 201.
Mount Douglas, Vancouver Island, British Columbia.
33. MARMOTA CAMTSCHATICA CAMTSCH.4TICA, Brandt
1843. Bull. Acad. St. Petersb., II, p. 364.
Kamtchatka.
34. MARMOTA CAMT.SCH.VriCA BUNGEI, Kasccnko
1901. .Ann. Mus. St. Petersb., VI, p. 615.
Omoloy, Vcrhoyansk Mountains, N. Siberia.
46o MARMOTA
35. MARMOTA CAMTSCHATICA DOPPELMAYRI. P.irula
1922. Ann. Mus. Zool. Acad. .Sci., XXII, 4, 80 pages.
The upper reaches of the River Nergili (east shore of Lake Baikal,
50 km. northwards from Sviatoi Nos).
36. MARMOTA CAMTSCHATICA CLIFTONI, Thomas
1002. .\nn. Mag. Nat. Hist. 7, IX, p. 444.
Versiansk Mountains, Yakutsk, N.-E. Siberia.
bohak Group
37. MARMOTA BOBAK, Mullcr
1776. Natursvst. Suppl. Regist. Band, p. 40.
Poland.
.Svnonvm: arctomvs, Pallas, 177S, Nov. Sp. Quadr. Glir. Ord., p. 75.
(Poland.)
38. M.ARMOTA HIMALAYANA HIMALAYANA, Hodgson
1 84 1. Journ. Asiat. Soc. Bengal, X, p. 777.
Nepal.
Synonym: talaricus, Jameson, 1847, L'Institut, XV, p. 384.
/lodgsoni, nianford, 1876, Yarkand Mission, Mamm., p. 35.
Nepal.
hemachalamis, Hodgson, 1843, Journ. .-\siat. Soc. Bengal,
XII, p. 410. Nepal.
39. MARMOTA HIMAL.\YAXA ROBCST.A, Milni-Edwards
1870. Nouv. Arch. Mus., VII, Bull., p. 92.
Moupin, Szechuan.
40. MARMOTA .SIBIRICA, Radde
1862. Reise .Sud. Ost. Sibir., p. 159.
Transbaikalia.
41. MARMOTA BAIB.\CINA BAIBACINA, Brandt
1843. Bull. Acad. Sci. St. Petersb., II, p. 364.
Altai.
42. MARM(JTA BA!B.\CINA CENTRALIS, Thomas
1909. Ann. Mag. Nat. Hist. 8, III, p. 260.
Aksai Plateau, 120 miles north of Kashgar. Turkestan.
caiulata Group
43. MARMOTA CAUDATA, Jacquemont
1844. Voy. dans I'lnde, IV, Zool., p. 66.
Kashmir.
44. MARMOT.^ AL'REA AUREA. Blanford
1S75. Journ. Asiat. Soc. Bengal, XLIV, pp. 106, 123.
Mountains west of Yarkand (E. Turkestan).
45. MARMOTA AUREA LITTLEDALEI, Thomas
1909. .Ann. Mag. Nat. Hist. 8, III, p. 259.
Alai Mountains, Pamir.
4(.. MARMOTA AUREA I'LAVINUS, Thomas
1909. Ann. Mag. Nat. Hist. 8, III, p. 259.
Hissar Mountains, 100 miles east of Samarkand.
MARMOTA— CYNOMYS 461
47. MARMOTA STIRMNGI, Thomas
1916. Journ. Bombay Nat. Hist. Soc. XXIV, p. 341.
Head of Chitral Nullah, Chitral (N.-W. Kashmir).
48. MARMOTA UICHROUS, Anderson
1875. Ann. Nat. Hist., XVI, p. 283.
Hills north of Kabul, Afghanistan.
marmota Group
49. MARMOTA MARMOTA, Linnaeus
1758. Syst. Nat., loth Ed., vol. i, p. 60.
Alps.
Synonym: alpina, Blumenbach, 1779, Handb. der Naturg, i, p. 80.
tigriiia, Bcchstein, 1801, Gemeinn Naturg. 1, 2nd ed.,
p. io2g.
alba, Bechstein, same reference, p. 1030.
nigra, Bechstein, same reference, p. 1030.
Not seen, and not allocated to group
50. MARMOTA MENZBIERl, Kashkarov
1925. Trans. Sci. Soc. Turkestan, 2, p. 47.
Tian-Shan, Central Asia.
Genus 31. CYNOMYS, Rafinesque
1817. Cynomys, Rafinesque, Amer. Monthly Mag., II, no. i, p. 45.
1916. Leucocrossuro.mys, Hollister, North Amer. Fauna, No. 40, p. 23. Spermophilus
gunnisoni, Baird. Valid as a subgenus.
Type Species. — Cynomys socialis, Rafinesque = Arctomys ludoviciana, Ord.
Range. — Western United States: forms named from Upper Missouri River,
Arizona, Wyoming, Utah, Colorado, New Mexico; and Coahuila,
northern Mexico. Good distribution maps of this genus are published by
Hollister, and in Anthony, Field Book North American Mammals, 1928,
pp. 219, 221.
Number of Forms. — Seven. The genus is revised by Hollister, 191 6,
North Amer. Fauna, no. 40.
Ch.xracters. — Dentition extremely hea\'y. Cheekteeth with the general plan
of CiteUiis, the inner border of each main tooth strongly
constricted. P. 3 very little reduced, nearly as large as P. 4. Upper cheekteeth
extremely hypsodont on inner side, and slanting outwards. Three depressions
on each tooth evidently remaining for a long time; these separate the four main
ridges; also M.3, which in related genera shows signs of simplification normally,
does not do so in this genus, the elements being as in the other molars (i.e. two
main ridges), and tending to persist. Lower cheekteeth about as in Citellus;
the posteroexternal cusp rather low, and in worn teeth separated from the cusp
in front of it by a small but well-marked re-entrant fold; M.3 with a narrow
depression running down the centre of the tooth.
Skull massive and angular. Toothrows markedly converging posteriorly.
Occipital region prominent. A well-marked sagittal crest developed in all
462 CYNOMYS
skulls seen. Infraorbital foramen triangular, its outer border much thickened,
and with a prominent masseter knob present. Zygomatic plate with upper
border well ridged, the infraorbital foramen apparently situated farther forward
with relation to this ridge than in related genera. Zygomatic width relatively
great. Mandible angular, powerfully ridged, the angular portion strongly pulled
inward, probably more so than in any other genus of squirrel.
Mammae 8 to 12 (Hollister). External form heavy, with tail much reduced,
probably not exceeding a quarter of head and body length. Digits of forefoot,
including the pollex, all with strong claws; D.3 the main digit, D.2 and D.4
subequal. Hindfoot with digits arranged as usual in terrestrial types.
In the typical subgenus, the jugal bone is described as strong, its outer
surface at angle of ascending branch very broad, triangular. C. mexicamis is
stated to have bullae larger than is usual in the genus.
In the subgenus Leiicocrossiiromvs, the jugal is "weak, thin and flat, the outer
surface at angle of ascending branch only very slightly thickened, the margin
rounded, not triangular. . . . Teeth smaller than in subgenus Cynomys, not so
much expanded laterally." The tail is tipped with white, instead of black (as
in the typical subgenus).
Remarks. — A very distinct genus. Not well represented in the British
Museum.
Forms seen: ludoviciaiius, gunnisoni.
List of Named Forms
Subgenus Cynomys, Rafinesque
1. CYNOMYS LUDOVICl.-^NUS LUDOVICI.'\NUS, Ord
1815. Guthrie's Geography, 2nd Amer. ed., vol. 2, p. 292.
Upper Missouri River.
.Synonym: socialis, Rafinesque, 1817, Amer. Monthly Mag., II, p. 45.
pyrrhotrichus, Elliot, 1905, Proc. Biol. .Soc. Washington,
XVIlI.p. 139. Oklahoma.
?>iissour!crisis. Warden, 1819, Stat. Pol. Hist. Ace. U.S. i,
p. 226.
latram, Harlan, 1825, Faun. Amer., p. 306.
2. CYNOMYS LUDOVICIANUS ARIZONENSIS, Mcarns
1890. Bull. Amer. Mas. Nat. Hist. II, p. 305.
Point of Mountain, near Wilcox, Cochise County, .Arizona.
3. CYNOMYS .MEXIC.\NUS. Mirriam
1S92. Proc. Biol. Soc. W.ashington. VII, p. 157.
La Ventura, Coahuila, Mexico.
Subgenus Leucocrossuromys, Hollister
4. CYNOMYS LEUCURUS, iMerriam
1890. North .Amer. Fauna, No. 40, p. 34.
Fort Bridger, Wyoming, Uinta County.
Synonym: le'disii, Allen. Bull. Amer. Mus. Nat. Hist. X, p. 456, iSgS.
Not of .Audubon S: Bachman, a Marmota from shores of
of Columbia River (see Hollister, North Amer. Fauna,
No. 40, p. 26, 1916).
CYNOMYS 463
5. CYNOMYS PARVIDENS, Allen
1905. Mus. Brookhii Inst. Arts & Sci., Science Hull, i, p. 119.
Buckskin Valley, Iron Count>', Utah.
6. CYNOMYS GUNNISONI GUNMSONI, Baird
1855. Proc. Acad. Nat. Sci. Philadelphia, VII, p. 334.
Cochetopa Pass, Saguache Count)', Colorado.
Synonym: co/»m6M«(«, True, Proc. U.S. Nat. Mus., VII, p. 593,1885.
7. CYNOMYS GUNNISONI ZUNIKNSIS, Hollister
1916. North Amer. Fauna, No. 40, p. 32.
Wingate, McKinley County, New Mexico.
The family Sciuridae is known fossil from the Oligocene.
FAMILY SCIURIDAE:
GENERAL WORKS OF REFERENCE
Forsyth Major, 1893, Proc. Zool Soc. London, 1893, p. 179. On some Miocene
Squirrels, with remarks on the dentition and classification of the Sciuridae.
Thom.\s, 1908, Journ. Bombay Nat. Hist. Soc, XVIII, p. 244. On the generic
position of the groups of Squirrels typified by Sciurus berdmorei and pernyi respect-
ively. (Rearrangement of genera from the Indo-Malayan region, other than Flying-
squirrels.)
Thomas, 1908, Ann. Mag. Nat. Hist. 8, I, p. i. The genera and subgenera of the
Sciuropterus group. (Rearrangement of genera of smaller Flying-squirrels.)
Thom.as, 1909. The generic arrangement of the African Squirrels: Ann. Mag. Nat.
Hist. 8, III, p. 467.
Allen, 1915, Review of the South American Sciuridae. Bull. Amer. Mus. Nat. Hist.,
XXXIV, p. 147.
Thomas, 1915, Ann. Mag. Nat. Hist. 8, XV, p. 383. The penis bone or baculum as a
guide to classification of certain Squirrels.
Miller, 1912, Catalogue of Mammals of Western Europe, pp. 897-946: Sciuridae and
Petauristidae : Sciurus, Citellus, Alaniiota, Sciuropterus ( ==Pleromys).
PocoCK, 1923, Classification of Sciuridae on the baculum. Proc. Zool. Soc. London,
p. 209, 1923.
PococK, 1922, Proc. Zool. Soc. London, p. 1171. On the external characters of the
Beaver and some Squirrels.
Howell, 1938, North Amer. Fauna, No. 56, p. i. Revision of North American Citellus
and rearrangement of genera and subgenera of North American Squirrels.
Allen, 1877, Monograph of North American Rodents, p. 637. Sciuridae.
Tullberg, 1899, Nova Acta Reg. Soc. Sci. Upsaliensis, XVIII, 3, 1.
Wroughton, 191 1, Journ. Bombay Nat. Hist. Soc, XX, p. 1012. Key to the Indo-
Malayan species of Petaurista.
Wroughton, 1919, Journ. Bombay Nat. Hist. Soc, XVI, p. 352. Indian Mammal
Sun-cy: Sciuridae.
Vinogradov, 1933, Tab. .^nal. Faune de I'URSS, Inst. Zool. Acad. Sciences, 10, p. i.
Key to Rodents of the U.S.S.R.: Sciuridae: Sciurus, Eutamias, Spermophilopsis,
Citellus, Marmola, Ptcromys.
Howell, 19 18, North .Amer. Fauna, No. 44. Revision of genus Glaucomys.
Thomas, 1888, Journ. Asiat. Soc. Bengal, LVII, p. 258. Eupetaurus.
Allen, 1914, Bull. Amer. Mus. Nat. Hist. XXXIII, p. 145. Review o{ Microsciurus.
Nelson, 1899, Proc Washington Acad. Sci., I, p. 38. Revision of Mexican and Central
.American Squirrels (Sciurus).
Allen, 1898, Bull. .Amer. Mus. Nat. Hist. X, p. 249. Revision of Tamiasciurus.
464 CASTORIDAE
Robinson &■ Kloss, igiS, Rec. Indian Miis., XV. pt. IV, pp. 171-250. Nominal List
of Oriental Sciuridac. (Arrangement of species and subspecies of Petaurista,
Eupetaurm, loniys, Beloiiiys, Pteroniyscus, PetauriUus, Hylopi-tes, Petinoinys, Acromys^
EoghtiiCfimys, Ratiifn, Ciillnsciurus, " Tomciilcs." Miiietis. Larisctis, Dreiiiomys,
Rliinosciiiriis, RhritJirosciiirus, Clyphotcs, " Taniinps," Fiiiuimhiiliis, Nannosciurus.)
Ingoldbv, 1927, Proc. Zool. Soc. London, p. 471. Revision of Hcliosciunis.
Howell, 1929, North Amer. Fauna, No. 52, p. i. Revision of Chipmunks, Taiiiias
"Etitamias."
Obolensky, 1927, C.R. Acad. Sci. Leningrad, p. 188. Revision of Palaearctic Ground-
squirrels : Citelhis, Sptriiwpln'lopsis.
Howell, 191 5. Revision of American species of Marntota, North. Ainer. Fauna, no. 37.
Hollister, 1 916, Revision of genus Cyiwinys. North Amer. Fauna, no. 40.
Hollister, East African Mammals: Sciuridac. Smiths. Inst. \J. S. Nat. Mus. Bull. 99,
1919, p. I.
Superfamily CASTOROIDAE
As here understood this contains one living family.
1896
1899
1918
1924
192S
Family CASTORIDAE
Thomas: Scuro.morph.^, part: Family Castoridae.
Tullberg: ScifROMORPH.^, part: Costoroidei, Family Castoridae.
Miller Sr Gidley: Superfamily Sciuroid.^e. Family Castoridae.
Winge : Family Sciuridac, part: Castorini.
Weber: C.\storoide.^ : Familv Castoridae.
GEOGR.'iPHlCAL DiSTRiBiTTiON. — Palaearctic and Nearctic. In North
America forms have been described from
Hudson Bay, Newfoundland, Alaska, Vancouver Island, Carolina, Michigan,
North Dakota, Texas, California, New Mexico, and Sonora (near Mexican
boundary line), Mexico. The genus formerly probably extended over the
greater part of the continent, and Anthony in Field Book of North American
]\Iammals, 192S, gives as the range for Castor canadensis, "most of North
America from Alaska and Labrador to the Rio Grande." In Europe, formerly
extending across most of the Continent, and including England; but now
restricted to Norway, and proiiably some of the main rivers of Central Europe,
as the Rhone, Elbe, Danube and Pripet (Flower); occurs in parts of the U.S.S.R.
(quoted by Vinogradov from basins of Rivers Vistula and Dnieper, former
Minsk, Smolensk, Chernigov, Poltava govts., former Voronej govt., basin of
River Sosva (north Ural mountains), and Mongolian Altai).
Number of Genera. — One.
ChaR-^cters. — Skull and external form heavy; zygomasseteric structure as
in typical specialized Sciuridae; the infraorbital foramen
forming a canal. Dental formula i. i, c. y, p. {, m. ;{ = 20. Cheekteeth excessively
hypsodont, but not evergrowing, the pattern changing little during life, and
characterized by narrow inner and outer enamel folds, as in certain Hystricoid
genera; bullae with neck protruding upwards and outwards; a pit-like depression
in basi-occipital; jugal in contact with the lachrymal, and immensely thickened
Fig. 1 1 6. Cynomys ludovicianus ludovicianvs, Ord.
B.M. No. 19.7.7.2841; X I.
Fig. 117. Cynomy'S ludovicianvs, Ord.
Cheekteeth; X 3.
30 — I,ivin|j; Rodents — I
Fig. 118. Castor fiber, Linnaeus.
Cheekteeth; X lA,
Fig. iiQ. Castor fiber, Linnaeus.
(From Miller's Cataloj^ue of the Mammals of Western Europe)
X i.
CASTORIDAE: CASTOR 465
anteriorly; externally highly modified for aquatic life, the hindfeet much en-
larged, the digits webbed; tail broad, flat, scaly and naked, the caudal vertebrae
flattened; anal and urinogcnita! orifices open within a common cloaca; tibia and
fibula united at base, but not fully fused (i.e. not comparing in this specialization
with Muridae, Dipodidae, etc.).
Remarks. — The diff'erences between Castor and the Sciuridae appear to
relate entirely to the internal characters (other than the cheek-
teeth), so far as superfamily separation goes. TuUberg mentions that it may
be that Castor has more in common with the Muridae than the Sciuridae, and
that it is possible that instead of forming a supergroup containing Sciuridae,
Castoridae and Geomyidae as he did, and one containing Aluridae, Anomalur-
idae, Ctenodactylidae, it might be correct to unite Castoridae, Geomyidae and
Muridae in one group, and possibly Ctenodactylidae, Anomaluridae and
Sciuridae in another. Bearing this in mind, and also that Weber regards the
group as a superfamily, it is retained as such here, though some doubt is felt
as regards the desirability of this classification; in most essential cranial charac-
ters the family stands close to the Sciuroidae.
It may be noted that, according to Miller & Gidley, many Castoroid genera
of Rodents are known fossil from the Holarctic and have been distinct from the
Sciuridae since the Oligocene; at least in their classification both groups are
quoted from that period.
There is one living Castoroid genus.
Genus I. CASTOR, Linnaeus
1758. Castor, Linnaeus, Syst. Nat., loth Ed., vol. i, p. 58.
Type Species. — Castor fiber, Linnaeus.
Range. — As in the family Castoridae.
Number of Forms. — Twenty.
Characters. — Skull with narrowed frontals, narrow braincase, rostrum
thick and relatively short. Temporal ridges forming a sharp
sagittal ridge in the adult, which divides anteriorly, the ridges extending forward
to about half-way along orbits, and forming short postorbital notches. Lambdoid
crest thick, occipital region prominent and angular. Paroccipital process of
medium length. Jugal greatly broadened anteriorly, a process directed upwards
on the anterior half of the upper border; jugal in contact with lachrymal. Bullae
of moderate size, the neck protruding sharply upwards and backwards, appearing
in superior view of skull as a conspicuous upwardly projecting tube, on either
side of the squamosals. Basioccipital w ith a pit-like depression situated near base
of foramen magnum. Palate slightly wider posteriorly than anteriorly, ending
in a short spinous process, at level just behind M.3. Incisive foramina rela-
tively small, narrow and situated considerably in front of the toothrows, as in
the Sciuridae. Mandible with coronoid process high, considerably higher than
the condylar, and angular portion flattened and rounded; the mandibular
466 CASTOR
symphysis extending back to P. 4 in adult. Upper part of zygomatic plate
deeply ridged. Nasals noticeably longer in Palacarctic forms than in American
species, so far as seen.
Incisors thick. Cheekteeth extremely hvpsodont, decreasing in size from
P.4 to M.3. Upper series with three long narrow enamel folds externally and
one internally, which tends to meet the first outer fold. Lower teeth with this
pattern reversed, the outer fold curving backwards between the second and
third inner folds.
Size very large, second only to Hxdrocliocnis in the Order at extreme develop-
ment. Form thickset; legs short; ears very small; hindfeet large, broad, with
five well-developed digits, the digits webbed, the claws powerful. D.2 with a
horny supplement rising beneath the claw, this probably used for dressing the
fur. Forefoot much smaller than hindfoot, with five well-developed digits, and
long curved claws, that of the poUex quite well developed, but noticeably
weaker than the others. D.3 the main digit. Fur very thick, consisting of a
dense soft underportion, and abundant growth of coarser longer hair. Tail
much broadened, almost without a vestige of hair, though slightly furred at
extreme base; of mediimi length (unique in the Order as regards structure).
The habits of these animals are too well known to need more than passing
mention, though that an animal of this size can cut down trees is remarkable
to say the least.
It is to be hoped that the excellent work in the preservation of these animals
by the late "Grey Owl" will lie carried on and will preserve them from
extinction.
More or less closely allied forms are known fossil from the Ilolarctic, as
already mentioned; the Castoroididae, containing a genus (Ctisturoii/cs) with
evergrowing laminate cheekteeth and slightly modified zygomasseteric structure
(Pleistocene), and the Chalicomyidae {Chalicomys, European Miocene and
Pliocene, Trogontherium, European Pliocene to Pleistocene, Palaeocastor and
others, North American Oligocene and Pliocene (Miller & Gidley)), seem
worthy of mention.
Forms examined: fiber, canaileiisis.
CASTORIDAE:
SPECIAL WORKS OF REFERENCE
J. Allen, Monograph North American Rodentia, 1877, p. 431.
TuLLBEBO, Nova Acta Reg. See. Sci. Upsalicnsis, XVIII, ser. 3, no. i, iSgg.
PococK, On the external characters of the Beaver and some Squirrels, Proc. Zool. Soc.
London, p. 1171, 1922.
Miller, Catalogue of Mammals of Western Europe, 1912, p. 947: Castoridae.
List of Named Forms
(The references and type localities are the work of Mr. R. W. Hayman.)
As already noted, so far as seen the Palaearctic forms differ from the North
American ones in the length of the nasals.
CASTOR 467
canadensis Group
1. CASTOR CANADKNSIS CANADENSIS, Kuhl
1820. Heitrage z. Zoologie, p. 64.
Hudson Bay.
Synonym: americanus, Richardson, Faun. Bor. Amer., 1829, p. 105.
2. CASTOR CANADENSIS BELUGAE, Taylor
1916. Univ. Calif. Publ. Zool. 12, p. 429.
Beluga River, Cook Inlet region, Alaska.
3. CASTOR CANADENSIS PHAEUS, Heller
1909. Univ. California Publ. Zool. 5, p. 250.
Admiralty Island, Alaska.
4. CASTOR CANADENSIS SAGITT.ATUS, Benson
I933- Journ. Mamm. Baltimore, 14, p. 320.
Indianpoint Creek, north-east of Barkerville, British Columbia.
5. CASTOR CANADENSIS LEUCODONTA, Gray
1869. Ann. Mag. Nat. Hist. 4, IV, p. 293.
Vancouver Island, British Columbia.
Synonym: pacificus, Rhoads, Trans. .Amer. Philos. Soc, n.s., vol. 19,
p. 422, 1898. Lake Keechelus, Cascade Mountains,
Washington.
6. CASTOR CANADENSIS MICHIGANENSIS, Bailey
1913. Proc. Biol. Soc. Washington, XXVI, p. 192.
Tahquamenaw River, Luce County, Michigan.
7. CASTOR CAN.ADENSIS MISSOURIENSIS, Bailey
1919. Journ. Mamm. Baltimore, i, p. 32.
Apple Creek, 7 miles east of Bismarck, Burleigh County, North Dakota.
8. CASTOR CANADENSIS CAROLINENSIS, Rhoads
1898. Trans. .Amer. Philos. Soc, n.s., 19, p. 420.
Dan Ri\er, Stokes County, North Carolina.
9. CASTOR CANADENSIS REPENTINUS, Goldman
1932. Journ. Mamm. Baltimore, 13, p. 266.
Bright .Angel Creek, Grand Canyon, .Arizona River, Colorado.
10. CASTOR CANADENSIS BAILEYI, Nelson
1927. Proc. Biol. Soc. Washington, XL, p. 125.
Humboldt River, near Winnemucca, Nevada.
11. CASTOR CANADENSIS TEXENSIS, BaUey
1905. North .Amer. Fauna, No. 25, p. 122.
Cummings Creek, Colorado County, Texas.
12. CASTOR CA.N.ADENSIS MEXICANUS, Bailey
1913. Proc. Biol. Soc. Washington, XXVI, p. 191.
Ruidoso Creek, 6 miles below Ruidoso, Lincoln County, New Mexico.
13. CASTOR CANADENSIS FROND.ATOR, Mcams
1897. Prelim, diag. new Mamm. Mexican border of U.S.A., p. 2 (Reprinted Proc. U.S.
Nat. Mus. XX, p. 502, 1898).
San Pedro River, Sonora, Mexico, near monument No. 98 of the
Mexican boundary line.
46S CASTOR
14. CASTOR CAECATOR, Bangs
1913. Bull. Mus. Comp. Zool. 54, p. 513.
Near Bay St. George, Newfoundland.
15. CASTOR SUBAURATUS SUBAURATUS, Taylor
1912. Univ. Calif. Publ. Zool. 10, p. 167.
Grayson, San Joaquin River, Stanislaus County, California.
16. CASTOR SUBAURATUS SHASTENSIS, Taylor
1916. Univ. Calif. Publ. Zool. 12, p. 433.
Casscl, Pitt River, Shasta County, California.
fiber Group
17. C.\STOR FIBER FIBER, Linnaeus
1758. Syst. Nat., loth Ed., vol. i, p. 58.
Sweden. {Range: Western European range of the genus.)
Synonym: albicus, Matschie, 1907, Sitz. Ber. Ges. Nat. Fr. Berlin,
p. 216. Anhalt, Gerniany.
balticus, Matschie, 1907. Sitz. Ber. Ges. Nat. n't. Berlin,
p. 217. Pomerania.
Miller, Catalogue Mammals of Western Europe, also quotes as
synonyms :
alhiis, Kerr, 1792, .Anim. Kingd., p. 222.
solitarius, Kerr, same reference, p. 224.
ia(-/fgn;i(s,Bechstein,Gemeinn. Naturgesch, Deutschlands i,
2nd ed., p. 913, 1801.
fiilvus, Bechstcin, same reference.
galliae, GeofiFroy, 1803, Cat. Mamm. Mus. Nat. Hist. Paris,
p. 168. {Rhone, France.) 1803.
m'ger, Desmarest, 1822, Mammalogie, pt. II, p. 27S.
varhis, Desmarest, same reference.
flatus, Desmarest, same reference.
gatlicus, Fischer, Synops. Mamm., p. 2S7, 1829.
proprius, Billberg, Linn. Samf., p. 34, 1833.
18. CASTOR FIBER VISTULANUS, Matschie
1907. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 219.
West Poland.
{Listed as a valid race by Vinogradov, Rodents occurring in
U.S.S.R., 1933)
iq. CASTOR FIBER POHLEI, Serebrennikov
1929. C.R. Acad. Leningrad, p. 275.
River Leplja, tributary of the N. Sosva, east slope of N. LTrals. Russia.
20. CASTOR FIBER BIRULAI, Serebrennikov
1929. C.R. Acad. Leningrad, p. 276.
River Bulungun, south of Kobdo, Mongolian Altai.
Superfamily GEOMYOIDAE
This group contains as here understood two families, the Geomyidae and
the Heteromyidae.
1896. Thomas: MYOMuRPH.^, part: Family Heteromyidae {with subfamilies Hetero-
myinae (Heleromys, Pcrognalhiis), and Dipodomyinae (Dipodomys, Microdipodops)).
Family Geomyidae.
GEOMYOIDAE 469
iSgg. Tullberg: Scu'romorpha, part: Geomyoidei. One family only recognized, the
Geomyidae, with subfamilies Dipodomyini and Geomyini.
1918. Miller & Gidley: Superfamily Sciuroidae: Family Geomyidae (living genera
referred to subfamily Geomyinae) ; and Family Heteromyidae.
1924. VVinge: Family "Saccomyidae" (=Heteromyidae) : Saccomyini and Geomyini.
J928. Weber: Geomyoidea: p'amily Heteromyidae, and Family Geomyidae.
Geographical Distribution. — America : the greater part of the United
States; South-western Canada ; Mexico and
Central America south to Venezuela, Colombia and Ecuador.
Characters. — Zygomassetcric structure essentially as in Sciuridae or
Castoridae, but rather more modified than in either group;
infraorbital foramen excessively reduced ; zygomatic plate tilted strongly
upwards.
Cheekteeth frequently evergrowing, and frequently becoming completely
simplified in pattern. Dental formula i. \, c. g, p. \, m. f = 20, the premolars not
or rarely showing any sign of reduction.
Large externally opening cheek-pouches present.
Fibula reduced and fully fused with the tibia high on the leg.
Zygoma abnormal; either extremely reduced and slender (Heteromyidae),
or comparatively robust, but with strongly shortened jugal which becomes
progressively reduced until the zygomatic arch is in specialized forms complete
without it (Geomyidae).
External form primitively Murine (Perognathus, Liomys, Heteromys) ; or
modified for bipedal saltatorial life {Microdipodops, Dipodomys), in which genera
the mastoids and auditory bullae become abnormally inflated, and the digits
of the hindfoot may be reduced to four; or in all Geomyidae much specialized
for underground life.
Remarks. — No doubt is felt in retaining this group as a superfamily. Most
zoologists who have classified the Order and given proper
consideration to zygomasseteric structure have placed this group in the neigh-
bourhood of the Sciuromorph series of Rodents, where they appear to belong.
In some ways the Geomyidae appear to me at their highest development
to be among the most highly specialized of all living Rodents.
Key to the Families of Geomyoidae
Skull and external form modified for subfossorial life; cheekteeth always
evergrowing in living members of the group, their structure always
near complete simplification. Incisors thick and powerful. Zygoma
comparatively robust, the jugal becoming progressively reduced
until the zygomatic arch is complete without it. Bullae and
mastoids never abnormally inflated. Palate much narrowed.
Frontals relatively narrow to extremely so. Family Geomyidae
Skull and external form never modified for subfossorial life; cheekteeth
rarely evergrowing (Dipodomys only), and their structure rarely
near complete simplification (adult Dipodomys only). Incisors thin
470 HETEROMVIDAE
and compressed. Zygoma much narrowed and reduced, thread-
like. A marked tendency towards great inHation of mastoids and
bullae. Palate not narrowed. Frontals relatively broad to ex-
tremely so. Family Heteromyidae
Family HETEROMYIDAE
1S96. Thomas: Myomorpha, part: Heteromyidae, with subfamilies Heteromyinae
{Heteromys, Perogiiathus), and Dipodoniyinae (Microdipoiiops, DipoiJoiiiys).
1S99. TiiUberg: Scuirc^morpha, part; Gcontyoidei ; Family Geomyidac, part, subfamily
Dipodomyini.
191S. Miller & Gidley: Supcrfamily Siii'RomAF. part: Family Heteromyidae.
1924. Winge: "Saccomyidae" (=Heteromyidae), part: " Saccomyini."
1925. Weber: GEOMYomKA, part: Family Heteromyidae.
Geographical Distribution. — Western North America, from British
Columbia southwards; west in U.S.A. to
the Dakotas and Texas ; through Mexico and Central America south to Ecuador,
Colombia and Venezuela.
Number of Genera. — Five.
Characters. — As indicated in the above key. "Orifice of infraorbital
foramen protected from muscle pressure by countersinking
in a vacuity which extends transversely through rostrum" (Miller & Gidley).
(Compare Geomyidae, p. 505.) Hindfoot long and narrow; in specialized
forms of bipedal saltatorial type. Cheekteeth hypsodont, but not evergrowing
except in Dipodoinvs; molars in adult usually with a two-lobed pattern, in
Dipodoinxs more or less simple.
The family is the subject of a recent and most extensive monograph by
Wood (Ann. Carnegie Mus, XXIV, p. 75, 1935); in this paper all known fossil
and recent forms have been very fully dealt with.
Wood divides the family into three subfamilies: Heteromyinae, Perogna-
thinae, and Dipodomyinae. The Heteromyinae contain, of living genera,
Heteromys and Lioinvs only. The Perognathinae contain Perogiiathiis and
Microdipodops. The Dipodomyinae Dipodoinxs alone. He remarks, discussing
the Dipodomyinae, "this subfamily is definitely related to the Perognathinae,
to which it shows much closer relationships than does either to the Hetero-
myinae; it may be that a more correct idea of relationships within the family
woidd be attained by consolidating these first two subfamilies."
For the purposes of the present work I propose to do so, recognizing only
two subfamilies, the Dipodomyinae to include Peiogiuitliiis and Microdipodops
and to be divided into two generic groups. The classification here adopted is
based on Wood's monograph, therefore, w-ith the above modification; this being
the most up-to-date work on the entire family, which in common with many
other North American groups is not well represented at the British Museum.
At first sight the family seems composed of two types of animal, the
"murine" Heteromys (from which, though unquestionably closely related to it,
Lioinvs was separated by Merriam in 1902 and is currently accepted as a full
HETEROMYIDAE: HETEROMYINAE 471
genus by American authors), and Perognatlms; and the "Dipodide" saltatorial
type, Microdipodops and Dipodomys.
It is tlicrt-forc of great interest that, according to Wood, Microdipodops is
more closely allied to Perogriathus (see notes under genus Microdipodops), and
not to Dipodomys as has been previously held ; and that Ileteromys (with Liomys)
stands rather apart from the rest of the family. From the teeth of the one skull
of Microdipodops examined it would appear that the above assumption as to
its relationship with Perognathiis rather than Dipodomys is correct, and that the
external saltatorial characters and the abnormal inflation of mastoids and bullae
has in each genus been developed independently.
Key to the Subi-amilies of Heteromyidae
"Lophs of lower premolars uniting first at buccal side, next at lingual;
those of upper premolars uniting first at lingual, next at buccal;
those of upper molars always and lower molars usually uniting at
the two ends surrounding a central basin; pattern of cheekteeth
preserved for a long time" (Wood). I^ullae showing no signs of
excessive inflation, and never reaching the level of grinding
surfaces of molars. (Incisors as far as seen not grooved.)
Subfamily Heteromyinae
{Heteromys, Liomys)
"Lophs of lower premolars uniting at centre of tooth; those of upper
premolars uniting first at or near centre of tooth; those of upper
molars uniting progressively from lingual to buccal margins;
those of lower molars uniting primitively at buccal margin, pro-
gressively at centre of tooth; the pattern being lost early in life"
(Wood). Bullae always well inflated; at highest development
e.xtremely so. Subfamily Dipodomyinae
Cheekteeth not evergrowing, in adult the pattern not completely
simplified; anterior zygomatic root not greatlv enlarged.
Bullae moderate (Perognathiis), or abnormally inflated
{Microdipodops). Perognathiis Group (Perognathi)
( Perognathus, Microdipodops)
Cheekteeth evergrowing, in adiflt and usually early in life the
pattern near complete simplification; anterior zygomatic
root greatly enlarged; bullae much inflated.
Dipodomys Group (Dipodomyes)
(Dipodomys)
Subfamily HETEROMYINAE
Geographical Distribution. — Southern Texas and Mexico southwards to
Panama, Colombia, Venezuela, Ecuador.
Characters. — As indicated in the above key. Skull with bullae little
inflated, the mastoids not or scarcely showing in superior
aspect; nasals thick and projecting far forwards over incisors, which are narrow
472 HETEROMYINAE: HETEROMYS
and opisthodont. Frontals normally scarcely constricted at all, and usually
with relatively well-developed supraorbital ridges which extend backwards over
the sides of the braincase. Palate moderately broad; two pairs of pits present
for the pterygoid muscles; hamulars not joining the bullae. Infraorbital fora-
men, as is usual in the family, minute, and situated far forwards, on side of
rostrum. Incisive foramina very small, and situated far in front of toothrow
Upper incisors (of those seen) plain. Lower jaw small and weak. External form
Murine, not specialized for saltatorial life. Fur usually bristly.
Containing two closely allied genera.
Key to the Genera of Heteromyinae
Adult pattern of cheekteeth less complicated, the enamel islands wearing
out; posterior loop of P. 4 with no deep re-entrant anterior fold.
LlOMYS
Adult pattern of cheekteeth not simplified, the enamel islands per-
sisting; posterior loop in crown of P. 4 with a deep re-entrant
anterior fold. IIeteromys
Genus I. HETEROMYS, Desmarest
1817. HETEROMYS, Desmarest, Nouv. Diet. Hist. Nat., vol. 14, p. 181.
1S23. Saccomys, Cuvier, Dents des Mamm., p. 186. (Sacconiys anthopihis; this name is
usually considered synonymous with Heteromys; the type species of Saccomys is
presumably unidentifiable.)
1902. Xylomys, Merriam, Proc. Biol. Soc. Washington, XV, p. 43. Heteromys
nelsoni, Merriam. Valid as a subgenus.
Type Species. — Mits anomalus, Thompson.
Range. — Southern Mexico (Vera Cruz, Oaxaca, Yucatan), through Guate-
mala, Salvador, Honduras, Nicaragua, Costa Rica and Panama to
Ecuador, Colombia and Venezuela.
Number of Forms. — Twenty-two.
Characters. — Skull as described aliove; upper molars of adult in two loops
with a median enamel island which is long, frequently open
exteriorly, and persistent. P. 4 with a fold extending across and dividing the
tooth completely into two lobes, and with a well-marked fold entering anteriorly
into the inner side of the hinder lobe. Lower molars with the same elements of
the upper teeth, but the folds when open do so interiorly. P. 4 also with a small
anterior outer fold, which may wear out. M.3 is not reduced in size. Tail
longer than head and body as a rule, poorly haired, with scales visible. Fur
normally bristly or spinv. Forefoot with medium claws; D.5 short, three centre
digits longer; pollex rudimentary. Hindfoot narrow, long though not extremely
Fig. I20. Heteromys anomalus anomalus, Thompson.
B.M. No. 97.4.7.2, ?; X 2j.
Fig. 121. Heteromys anomalvs anomalus, Thompson.
B.M. No. 97.4.7.2, 9; X zi-
474
HETEROMYS
so; with the three central digits much elongated, D.5 shorter, the hallux short.
This arrangement of digits is constant in all non-saltatorial members of the
family I have seen.
H. nelsoni, not represented at the British Museum, is separated subgeneri-
callv as Xxlomxs. with the following characters:
"Pelage soft, without stiff bristles . . . skull light, braincase high and rounded,
supraorbital beads small and faint ; upper surface of maxillary root of zygomata
large, heavy and rectangular; frontals much elongated, pushing nasals and pre-
maxillae far forward; underjaw broad, without trace ot tubercle over root of
incisor and with angle very slightly everted. Dentition heavy. Posterior prism
of last upper molar more or less completelv double, the crown of the tooth
Fig. 122. Heteromys anomalus axomalus, Thompson.
Cheekteeth: B.M. No. 97.4.7.2, $; x 9.
presenting two complete transverse loops and a more or less perfect posterior
loop."
The species of Heteromys are fully revised by Goldman (North Amer.
Fauna, No. 34, p. 14, 191 1). A key to these will be seen on consulting the above-
mentioned work. With the exception of H. gaiimeri, all seem ver)' closely allied.
\r\ gaiimeri, the sole is hairy from near posterior tubercle to the heel; in all other
species it is naked.
Forms seen : (uioiiicilns, mistidlis, bicolor, desinarestianus, gaiimeri, goldmani,
longicaiidatiis, " mchmohiicas" rcpeiis.
List of Named Forms
(The references and type localities in all genera of Ileteromvidae are the
vork of Mr. G. W. C. Holt.)
HETEROMYS 475
Subgenus Heteromys, Desmarest
1. HKTEROMYS ANOMALUS ANOMALUS, Thompson
1815. Trans. Linn. Soc. XI, p. 161.
St. .^nn's Barracks, island of Trinidad.
Synonym: melanoleucas. Gray, 1868, Proc. Zool. Soc. London, p. 204.
Venezuela (see Alston, Biol. Cent. Amer. Mamm.,
p. 167, Ann. Mag. Nat. Hist. 5, VI, 1880).
2. HKTICROMVS A.\OMAI-US BRACHIALIS, Osgood
1912. Field Mus. Zool. Pub. 10, p. 54.
El Panorama, Rio Aurare, Venezuela.
,. HKTKROMYS JK.SUPI, Alkn
1899. Bull. Amer. Mus. Nat. Hist. XII, p. 201.
Colombia.
4. HETEROMY.S BICOLOR, Gray
1868. Proc. Zool. Soc. London, p. 202.
Salle, Honduras.
5. HETEROMY.S LOMITENSIS, Allen
1912. Bull. Amer. Mus. Nat. Hist. XXXI, p. 77.
Las Lomitas, Cauca, Columbia.
6. HETEROMYS AUSTRALI.S AUSTRALI.S, Thomas
1901. Arm. Mag. Nat. Hist., ser. 7, VII, p. 194.
St. Javier, Lower Cachabi River, N. Ecuador.
7. HETEROMYS AUSTRALIS CONSCIUS, Goldman
1913. Smiths. Misc. Coll. LX, no. 22, p. 8.
Cana, mountains of E. Panama.
8. HETEROMYS DESMARESTIANUS DESMARESTIANUS, Gray
1868. Proc. Zool. Soc. London, p. 204.
Coban, Guatemala.
0. HETEROMYS DESMARESTIANUS GRISEUS, Mcrriam
1902. Proc. Biol. Soc. Washington, XV, p. 42.
Mountains near Tonala, Chiapas, Mexico.
10. HETEROMYS DESMARESTIANUS PSAKASTUS, Dickey
1928. Proc. Biol. Soc. Washington, XLI, p. 10.
Los Essemiles, Dept. Chalatenango, EI Salvador.
11. HETEROMYS ZON.ALIS, Goldman
1912. Smiths. Misc. Coll. LVI, no. 36, p. 9.
Rio Indio, near Gatun, Canal zone, Panama.
12. HETEROMYS LONGICAUDATUS, Gray
1868. Proc. Zool. Soc. London, p. 204.
Mexico.
13. HETEROMYS GOI.DMANI, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 41.
Chicharras, Chiapas, Mexico.
14. HETEROMYS r>:PTURUS, Mcrriam
1902. Proc. Biol. Soc. Washington, XV, p. 42.
Mountains near Santo Domingo, Oa.\aca, Mexico.
47*' HETEROMYS— LIOMYS
15- HETEROMYS TEMPORALIS, Goldman
191 1. North Amer. Fauna, no. 34, p. 26.
Motzorongo, Vera Cruz, Mexico.
lb. HETEROMYS REPENS, liaiiss
1902. Bull. Mus. Comp. Zool. XXXIX, p. 45.
Boquete, southern slope of Volcan de Chiriqui, Panama.
■ 7. HETEROMYS ORi:STERUS, Harns
1932. Occ. Pap. Mus. Zool. Univ. Michigan, no. 248, p. 4.
El Copey de Dota, Cordillerade Talamanca, Costa Rica.
iS. HETEROMYS PANAMENSIS, Uoldmaii
1912. Smiths. Misc. Coll. LVI, no. 36, p. 9.
Cerro .'\zul. near headwaters of Chagres River, Panama.
19. HETEROMYS CR.ASSIRO.STRIS, Goldman
1912. Smiths. Misc. Coll. LX, no. 2, p. 10.
Near head of Rio Limon, Mount Pirri, E. Panama.
20. HETEROMYS FUSCATUS, Allen
1908. Bull. Amer. Mus. Nat. Hist. XXIV, p. 652.
Tunia, Nicaragua.
21. HETEROMYS GAUMERI, Allen & Chapman
1897. Bull. Amer. Mus. Nat. Hist. IX, p. 9.
Chichenitza, Yucatan, Mexico.
Subgenus Xylomys, Merriam
22. HETEROMYS NEl,SONI, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 43.
Pinabete, Chiapas, Mexico.
Genus 2. LIOMYS, Merriam
1902. LlOMYS, Merriam, Proc. Biol. Soc. Washington, XV, p. 44,
Type Species. — Heteromvs alleni, Coues.
Range. — Southern Texas, Mexico, Guatemala, Nicaragua, Honduras, Costa
Rica and Panama.
Number of Forms. — Twenty-nine.
Char-^cters. — Like Heteromys, but angle of mandible more strongly everted,
and teeth becoming more simplified, the enamel islands not
persistent, but wearing out with age; P. 4 with the posterior loop slightly
notched but with no deep re-entrant angle.
This genus is revised by Goldman (North. Amer. Fauna, p. 32, no. 34, 191 1).
He recognizes three specific groups:
the irroratus group, with light grevish coloration and five-tuberculate soles
on hindfeet;
the pictiis group, characterized by rich orange rufous lateral lines, and six-
tuberculate hindfeet; and
LIOMYS 477
the crispus group, including small species with short tails, plain coloration,
and peculiar dental characters.
(The dental characters do not seem to be constant, according to Wood.)
Forms seen: adspersus, alleni, bulleri, "hispidus," irroratus, nigrescens,
obscurus, picttis, salvini.
List of Named Forms
picttis Group
1. LIOMYS PICTUS PICTUS, Thomas
1893. Ann. Mag. Nat. Hist. 6, XII, p. 233.
Mineral San Sebastian, Jalisco, Mexico.
Synonym: hispidus, Allen, 1897, Bull. Amer. Mus. Nat. Hist. IX,
p. 56. Compostela, Nayarit, Mexico.
2. LIOMYS PICTUS ESCUIN.^PAE, Allen
1906. Bull. Amer. Mus. Nat. Hist. XXII, p. 211.
Escuinapa, Sinaloa, Mexico.
3. LIOMYS PICTUS SONORANUS, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 47.
Alamos, Sonora, Mexico.
4. LIOMYS PICTUS PLANTINARENSIS, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 46.
Plantinar, Jalisco, Mexico.
5. LIOMYS PICTUS PARVICEPS, Goldman
1904. Proc. Biol. Soc. Washington, XVII, p. 82.
La Salada, 40 miles south of Uruapan, Michoacan, Mexico.
6. LIO.MYS PICTUS ROSTRATUS, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 46.
Near Ometepec, Guerrero, Mexico.
7. LIO.MYS PICTUS PHAEURUS, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 48.
Pinotepa, Oaxaca, Mexico.
8. LIOMYS PICTUS ISTHMIUS, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 46.
Tehuantepec, Oaxaca, Mexico.
9. LIOMYS PICTUS VER.\ECRUCIS, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 47.
San Andres Tuxtla, Vera Cruz, Mexico.
Synon>Tn: orbitalis, Merriam, 1902, Proc. Biol. Soc. Washington, XV,
p. 48. Catemaco, Vera Cruz, Mexico.
10. LIOMYS PICTUS OBSCURUS, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 48.
Carrizal, Vera Cruz, Mexico.
Synonym: paralitis, Elliot, 1903, Field Columb. Mus. Publ. 80, zool.
ser., vol. 3, p. 233. San Carlos, Vera Cruz.
11. LIOM\"S .ANNECTENS, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 43.
Pluma, Oaxaca, Mexico.
478 LIOMYS
crispus Group
12. LIOMYS CRISI'IS CRISI'LS. Mernam
IQ02. Proc. Biol. Soc. Washinf»ton, XV, p. 49.
Tonala, Chiapas, Mexico.
1.1. LIOMYS CRISPUS SKTOSUS, Mernam
igo2. Proc. BioL Soc. Washington, XV, p. 49.
Hufhuetan, Chiapas, Mexico.
14. LIOMYS VULC.ANI. MVn
igoS. Bull. Amer. Mus. Nat. Hist. XXIV, p. 652.
Volcan de Chinandega, Nicaragua.
15. LIOMYS HETEROTHRIX, Mernam
1902. Proc. Biol. Soc. Washington, XV, p. 50.
San Pedro Sula, Honduras.
16. LIO.MYS S.ALVINI S.-\LVINI. Thomas
1893. .Ann. Mag. Nat. Hist. 6, XI, p. 331.
Duenas, Guatemala.
17. LIOMYS S.YLVINI NIGRESCENS, Thomas
1S93. -Ann. Mag. Nat. Hist. 6, XII, p. 234.
Costa Rica.
18. LIOMYS ANTHONY I, Goodwin
1932. Amer. Mus. Nov. no. 528, p. 2.
Sacapulas, Central Guatemala.
iq. LIOMYS ADSPERSL'S, Peters
1874. Monatsb. k. preuss. Akad. Wiss. Berlin, p. 357.
Panama.
irroratus Group
20. LIOMYS IRRORATLS IRRORATUS, Gray
1868. Proc. Zool. Soc. London, p. 205.
State of Oaxaca, Mexico.
Synonym: (7/6o/;mAo/»s, Gray, Proc. Zool. Soc. London, p. 205, 1S6S.
La Parada, Oaxaca, Mexico.
21. LIOMYS IRRORATUS TORRIDUS, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 45.
Cuicatlan, Oaxaca, Mexico.
Synonym: exiguus, Elliot, Field Columb. Mus. Publ. 71, zool. ser.,
vol, 3, p. 146, 1903. Puente de Ixtla, Morelos, Mexico.
22. LIOMYS IRROR.ATUS MINOR, Merriam
igo2. Proc. Biol. Soc. Washington, XV, p. 45.
Huajuapam, Oaxaca, Mexico.
23. LIOMYS IRRORATUS ALLENI, Coues
1881. Bull. Mus. Comp. Zool. Harvard Coll. VIII, p. 187.
Rio Verde, San Luis Potosi, Mexico.
24. LIOMYS IRRORATUS PRETIOSL'S, Goldman
191 1. North Amer. Fauna, no. 34, p. 58.
Metlaltoyuca, Puebla, Mexico.
DIPODOMYINAE 479
25. LIOMYS IRRORATUS TEXENSIS, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 44.
Brownsville, Cameron County, Texas.
26. LIOMV.S IRRORATLS CANUS. Merriam
1902. Proc. Biol. Soc. Washinuton, XV, p. 44.
Near Parral, Chihuahua, Mexico.
27. LIOMYS IRROR.\TLS JALISCF.NSIS, Allen
1906. Bull. .Vmer. Mus. Nat. Hist., XXII, p. 251.
Las Canoas, about 20 miles west of Zapotlan, Jalisco, Mexico.
28. LIOMYS BULLERL Thomas
1893. Ann. Mag. Nat. Hist. 6, XL p. 330.
Laguna, Sierra de Juanactlan, Jalisco, Mexico.
29. LIOMYS GUERRERENSIS, Goldman
191 1. North Amer. Fauna, no. 34, p. 62.
Omilteme, Guerrero, Mexico.
Subfamily DIPODOMYINAE
Geographical Distribution. — The northern part of the range of the
family, south to Central Mexico.
Number of Genera.- — As here understood three, divided into two generic
groups.
Characters. — As indicated in the key, p. 471 ; external form Murine in
Perognat/tus, otherwise specialized for bipedal saltatorial life.
Skull characterized by wide braincase and moderately or abnormally inflated
bullae, the skull becoming gradually narrower towards the anterior zygomatic
root; frontals not or scarcely constricted, as in Heteromyinae; rostrum slender;
nasals projecting far forwards over incisors, which are narrowed, opisthodont,
and grooved in living genera. Incisive foramina very small; palate relatively
broad. Cheekteeth hypsodont, at extreme development evergrowing; M. f
tending to become reduced in size.
A key to the two generic groups is given on p. 471.
The Perognathtts Group
Cheekteeth not evergrowing, not becoming simplified in pattern; anterior
zygomatic root not greatly enlarged on joining the lachrymal.
Key to the Genera of the Perognathus Grout
Mastoids and bullae not abnormally inflated; hindfoot shorter; not
specialized for saltatorial life. Perognathus
Mastoids and bullae abnormally inflated, at maximum for familv and
perhaps for the whole Order; hindfoot longer; specialized for
saltatorial life. Microdipodops
31 — Living Rodents — I
4!So PEROGNATHUS
Genus i. PEROGNATHUS. Wied.
iS3ij. Perognathis, Wied, Xo\a Acta Phys. Med. Acad. Caes. Leop. Carol, XIX,
pt. I, p. 368.
1848. Cricetodipus, Peale. Mamni. and Ornith. Wilkes Exped., VIII, 2nd ed., p. 52.
(Cricetodipus parvus, Peale.)
1889. Ch.\etodipi's, Merriani, North Amer. Fauna, no. i, p. 5. Perognathus spinnliis,
IVIerriam. Valid as a subgenus.
Type Species. — Perognathus fasciatus, Wied.
R.\NGE. — Western North America from British Columbia to Central Mexico
(in U.S.A. known from Washington, Oregon, Idaho, Wyoming,
North and South Dakota, Nebraska, California (and Lower California), Nevada,
Utah, Arizona, Colorado, New Mexico, Kansas, Oklahoma, Texas).
Number of Forms. — About one hundred and twentv-six.
Ch.\r.\cti:rs. — Frontals scarcely constricted, with supraorbital ridges feebly
marked or absent; bullae inflated, the mastoids in the typical
subgenus appearing in superior aspect of the skull; the bullae nearly meeting
anteriorly. Incisive foramina minute; palate broad, extending to M.3.
Cheekteeth originally showing marked signs of three longitudinal rows of
cusps (to be seen for instance in No. 2.3.6.27, P. hispidiis, Texas, at British
Museum); but soon wearing down; general adult plan not unlike that of
Heteromxs; a long fold more or less separating the molars into two lobes;
premolar with narrow anterior lobe and much wider posterior one; M. 5 very
small. P. 4 lower with four cusps, one at each corner, apparently more or less
persistent; this tooth reduced, smaller than M.i; and with three folds, one
external, one internal, one anterior; lower molars much like the upper series
in general arrangement; in very young teeth, there is a pattern apparently
strongly reminiscent of Murinae, i.e. three rows of cusps, the outer row very
much reduced.
Size usually small to very small; tail about subequal to head and body,
or mav be longer, well haired; hindfoot moderately long, general arrangement
of the digits as in Heteromxs; forefoot not abnormal. Spines present on the
rump of some species (subgenus Clmetodipus).
Two well-marked subgenera are admitted : the typical, and Chaetodipiis,
Merriam.
In Chaetodipiis, the soles are naked, the pelage harsh, often with spiny
bristles on rump.
In Perognathus s.s. the pelage is normal, without spines; the soles are usually
hairy {except formosus).
In Chaetodipiis, the bullae are less inflated than in Perognathus, the mastoids
relatively small, not projecting beyond plane of occiput; the "mastoid side of
parietal is equal to or shorter than the other sides" (Osgood), the audital bullae
are separated by full width of basisphenoid, and the ascending branches ot the
supraoccipital are heavy and laminate, instead of slender and threadlike in
Fig. 123. Perocnathus hispidus hispidls, Baird.
B.M. No. 2.3.6.47, 5; X 3J.
Fig. 124. Perocn.^thls hispidus hispidus, Baird.
B.M. No. z. 3.6.47, f, X 3 J.
482
PEROGNATHUS
Perogmithtis pnipt-r. As indicated above, the characters of the bullae of Perog-
nathus s.s. do not agree with one of these characters.
The genus has been fully revised by Osgood (North Amer. Fauna, no. 18,
1900).
^-SiO^fc
Fig. 125. Perognathus hispidus hispidus, Baird.
Cheekteeth: B.M. No. 2.3.6.47, $; X 17.
From Osgood's key, it would appear that the essential characters of the ten
specific groups currently recognized are as follow:
Subgenus Perognathus
1. Antitragus not lobed; hindfoot 20 or less.
fasciatus group: tail about equal to head and body, or slightly shorter;
lower premolar smaller than or subequal to M.3.
longiiiiembiis group: tail longer than head and body except pacificus;
lower premolar larger than M.3.
2. Antitragus lobed; hindfoot more than 20.
parvus group: sole normal for subgenus; tail moderate.
formosiis group: sole naked; tail long and heavily crested.
Subgenus Chaetodipus
I. Rump with spines or bristles.
caUfornicus group: lateral line present; bristles moderate, usually
confined to rump; ears elongated.
PEKOGNATIIUS 483
intermedins group: similar to californiciis, but cars not elongated.
(Rump not spiny in artus.)
spinutus group: no lateral line, or this very faint; pelage very hispid,
with bristles extending to sides.
2. Rump without bristles.
hispidiis group: tail not crested, shorter than head and body; skull with
supraorbital bead in adult.
baileyi group: tail crested; no supraorbital bead in adult; tail much
longer than head and body; interparietal width about equal to
interorbital width.
penicillatus group: tail slightly longer than head and body, crested;
interparietal width exceeds interorbital width.
Forms seen: flams, formosus, infraluteus, lordi, parvus, panamintinus;
angustirostris, femoralis, Idspidus, inornatus, pemnsulae, pertiix, pricei.
List of Named Forms
Subgenus Perognathus, Wied
fasciatus Group
1. PEROGNATHUS, FASCIATUS FASCIATUS, Wied
1839. Nova .^cta Phys. Med. Acad. Caes. Leop. Carol., XIX, pt. i, p. 369. Upper
Missouri River, North-western N. Dakota.
2. PEROGNATHUS FASCIATUS INFRALUTEUS, Thomas
1893. Ann. Mag. Nat. Hist. 6, II, p. 406.
Loveland, Larimer County, Colorado.
3. PEROGNATHUS FASCI.\TUS LITUS, Gary
1911. Proc. Biol. Soc. Washington, XXIV, p. 61.
Sun, Sweetwater Valley, Fremont County, Wyoming.
4. PEROGN.ATHUS FLAVESCENS FL.WESCENS, Merriam
i88g. North Amer. Fauna, no. i, p. 11.
Kennedy, Cherry County', Nebraska.
5. PEROGNATHUS FL.^VESCENS COPEI, Rhoads
1893. Proc. Acad. Nat. Sci. Philadelphia, p. 404.
Near Mobeetie, Wheeler County, Texas.
6. PEROGN.-\THUS FL.'WESCENS PERNIGER, Osgood
1904. Proc. Biol. Soc. Washington, XVII, p. 127.
Vermilion, Clay County, South Dakota.
7. PEROGNWTHUS MERRIAMI MHRRIAMI, .-Mlm
1892. Bull. Amer. Mus. Nat. Hist., IV, p. 45.
Brownsville, Cameron County, Texas.
Synonym: mearnsi, .Allen, 1896, Bull. Amer. Mus. Nat. Hist., VIII,
p. 237. Watson's Ranch, Bexar County, Texas.
4S4 PEROGNATHUS
8. PEROGNATHUS MERRIAMI GII.VUS, Osgood
I goo. North Amer. Fauna, no. 18, p. 22.
Eddy, near Carlsbad, Eddy County, New Mexico.
t). PHROCJNATHIS FLAVL'S H..AVL'S, Baird
1855. Proc. .■\cad. Nat. Sci. Philadelphia, VII, p. 332.
El Paso, El Paso County, Texas.
10. PEROGN.VrHU.S FLAVUS PIPERl, Goldman
1917. Proc. Biol. Soc. Washinijton, XXX, p. 148.
Twenty-three miles south-west of Newcastle, Weston County,
Wyoming.
11. PEROGN.\THLS FLAVUS BIMACUL.-\TUS, Merriam
1889. North Amer. Fauna, no. i, p. 12.
Fort Whipple, Yavapai County, Arizona.
12. PERt)GNATHUS FL.AVUS FULIGINOSUS, iXIerriam
iSgo. North .Amer. Fauna, no. 3, p. 74.
Cedar belt north-east of San Francisco Mountain, Coconino County,
Arizona.
13. PEROGNATHUS FL.AVL'S MEXIC.^NUS, Merriam
1S94. Proc. Acad. Nat. Sci. Philadelphia, p. 265.
Tlalpam, Federal district, Mexico.
14. PEROGN.ATHUS FLAVUS HOPIENSIS, Goldman
1932. Proc. Biol. Soc. Washington, XLV, p. 89.
Oraibi, Hopi Indian Reservation, Navajo County, Arizona.
15. PEROGNATHUS FLAVUS SONORIENSIS, Nflson & Goldman
1934. Joum. Washington Acad. Sci. XXIV, p. 267.
Costa Rica Ranch, Sonora, Mexico.
16. PEROGNATHUS APACHE APACHE, Merriam
1889. North Amer. Fauna, no. i, p. 14.
Keam Canyon, Apache County, Arizona.
17. PEROGN.VPHUS APACHE CLEOMOPHILA, Goldman
1918. Proc. Biol. Soc. Washington, XXXI, p. 23.
Wmona, Coconino County, Arizona.
iS. PEROGN.a.THUS APACHE CARYI, Goldman
igiS. Proc. Biol. Soc. Washington, XXXI, p. 24.
Eight miles west of Rifle, Garfield County, Colorado.
IQ. PEROGNATHUS APACHE MEL.ANOTIS, Osgood
1900. North Amer. Fauna, no. 18, p. 27.
Casas Grandes, Chihuahua, Mexico.
20. PEROGNATHUS GVPSI, Dice
1929. Occ. Pap. Mus. Zool. Univ. Mich., 203, p. i.
New Mexico: White Sands, 12 miles south-west of Alamogordo, Otero
County.
21. PEROGN.VPHl'S CAI.LISTUS, Osgood
igoo. North Amer. Fauna, no. 18, p. 28.
Kinney Ranch, Sweetwater County, Wyoming.
I
PEROGNATHUS 485
longimembris Group
23. PEROGNATHUS KLIHATUS, Klliot
1903. Field Columb, Mus. Publ. 87, 7.00I. ser., vol. 3, p. 252.
Lockwood Valley, near Mount Finos, Ventura County, California.
Regarded by Osgood as identical with longimembris longimembris
(Proc. Biol. Soc. Washington, XXXI, p. g6, 1918).
21. PF.ROGNATHUS LONGIMEMBRI.S LONGIMEMBRIS, Coues
1875. Proc. Acad. Nat. Sci. Philadelphia, p. 305.
Old Fort Tejon, Tehachapi Mountains, Kern County, California.
24- PEROGNATHUS LONGIMEMBRIS PANAMINTINUS, Merriam
1894. Proc. Acad. Nat. Sci. Philadelphia, p. 265.
Perognathus Flat, Panamint Mountains, Inyo Count>', California.
25. I'EROGN.-\THUS LONGIMEMBRIS ARENICOLA, Stephens
1900. Proc. Biol. Soc. Washington, XIII, p. 153.
San Felipe Mountains, San Diego County, California.
26. PEROGNATHUS LONGIMEMBRIS BANGSI, Mearns
1898. Bull. Amer. Mus. Nat. Hist. X, p. 300.
Palm Springs, Colorado Desert, Riverside County, California.
27. PEROGNATHUS LONGIMEMBRIS BREVINASUS, Osgood
igoo. North Amer. Fauna, no. 18, p. 30.
San Bernardino, San Bernardino Count>', California.
2.S. PEROGNATHUS LONGIMEMBRIS AESTIVUS, Huey
1928. Trans. S. Diego Soc. Nat. Hist. 5, p. 87.
Sangre de Cristo, Lower California, Mexico.
2Q. PEROGNATHUS LONGIMEMBRIS VENUSTUS, Huey
1930. Trans. S. Diego Soc. Nat. Hist. 6, p. 233.
Lower California, Mexico, San Agustin.
30. PEROGNATHUS LONGIMEMBRIS ARIZONENSIS, Goldman
193 1. Proc. Biol. Soc. Washington, XL IV, p. 134.
North side of Marble Canyon of Colorado River, Arizona.
31. PEROGN.ATHUS LONGIMEMBRIS CANTWELLI, Blocker
1932. Proc. Biol. Soc. Washington, XLV, p. 128.
Hyperion, Los Angeles County, California.
32. PEROGN.-\THUS LONGIMEMBRIS KINOENSIS, Huey
1935. Trans. S. Diego Soc. Nat. Hist. 8, p. 73.
Bahia Kino, Sonora, Mexico.
33. PEROGNATHUS LONGIMEMBRIS ARCUS, Benson
1935. Univ. Cal. Pub. Zool. XL, p. 451.
Utah: Rainbow Bridge, San Juan County.
34. PI-ROGN.VrHUS PERICALLES, Elliot
1903. Field Columb. Mus. publ. 87, zool. ser. vol. 3, p. 252.
Keeler, Owens Lake, Inyo County, California.
35. PEROGNATHUS BOMBYCINUS, Osgood
1907. Proc. Biol. Soc. Washington, XX, p. 19.
Yuma, Yuma County, .\rizona.
486 PEROGNATHUS
36. PEROGNATHUS NEVADENSIS, Merriam
1S94. Proc. Acad. Nat. Sci. Philadelphia, p. 264.
Halleck, E. Humboldt Valley, Elko County, Nevada.
37. PEROGNATHl'S PACIFICU.S, Mcanis
1898. Bull. Anier. Mus. Nat. Hist. X, p. 299.
Mexican boundary monument no. 258, shore of Pacific Ocean, San
Diego County, California.
38. PEROGN-VnU'S AMPLUS AMPLUS, Osgood
igoo. North Amer. Fauna, no. 18, p. 32.
Fort Verde, Yavapai County, Arizona.
39. PEROGN.-\THUS AMPLUS TAYLORI, Goldman
1932. Journ. Washington Acad. Sci. XXII. p. 488.
Santa Rita Range Reserve, Pima County, Arizona.
40. Pi:ROf;\.ATHUS AMPLUS ROTUNDUS, Goldman
1932. Journ. Washington Acad. Sci. XXII, p. 387.
WcUton, Yuma County, Arizona.
41. Pl';ROC;N.VrHUS AMPLUS PERGRACILIS, Goldman
1932. Journ. Washington Acad. Sci. XXII, p. 387.
Hackberry, Mohave County, Arizona.
42. PEROGN.ATHUS AMPLUS JACKSONI, Goldman
1933. Journ. Washington Acad. Sci. XXIII, p. 465.
Yavapai County, Arizona.
43. PEROGNATHUS AMPLUS CINERIS, Benson
1933. Proc. Biol. Soc. Washington, XLVI, p. loq.
Wupatki Ruins, Coconino County, Arizona.
44. PEROGN.ATHUS AMPLUS AMMODYTES, Benson
1933. Proc. Biol. Soc. Washington, XLVI, p. no.
Two miles south of Cameron, Coconino County, .Arizona.
45. PEROGNATHUS INORN.ATUS INORN.ATUS, Merriam
1889. North Anier. Fauna, no. i, p. 15.
Fresno, Fresno County, California.
4h. PERDGN.ATHUS INORNATUS NEGLECTUS, Tax lor
1912. Univ. Calif. Publ. Zool. X, p. 155.
McKittrick, Kern County, California.
parvus Group
47. PEROGN.\THUS PARVUS PARVUS, Peak-
1S48. U.S. Explor. Exped. vol. 8, mamm. S: ornith, p. 53.
Oregon; probably neighbourhood of the Dalles, Wasco County.
Synonym: monticola, Baird, 1857, Mamm. N. Amer. p. 422. Montana.
48. PEROGN.ATHUS PARVUS IDAHOENSIS, Goldman
1922. Proc. Biol. Soc. Washington, XXXV, p. 105.
Echo Crater, 20 miles south of Arco, Blaine Count>-, Idaho.
49. PEROGNATHUS PARVUS MOLLIPILOSUS, Coucs
1875. Proc. Acad. Nat. Sci. Philadelphia, p. 296.
Fort Crook, Shasta County, California.
PEROGNATHUS 487
50. PEROGNATHUS PAKVLS OLIVACEUS, Merriam
1889. North Amer. Fauna, no. i, p. 15.
Kclton, Boxelder County, Utah.
Synonym: olivaceus amoemis, Merriam, North Amer. Fauna, no. i,
p. 16. Nephi, Juab County, Utah, 1889.
51. PEROGNATHUS PARVUS CUARUS, Goldman
igiy. Proc. Biol. Soc. Washington, XXX, p. 147.
Cumberland, Lincoln County, Wyoming.
52. PEROGNATHUS PARVUS MAGRUDERENSIS, Osgood
1900. North .\mer. Fauna, no. 18, p. 38.
Mt. Magruder, Nevada, near boundary between Inyo County, Cali-
fornia, and Esmeralda County, Nevada.
53. PEROGNATHUS XANTHONOTUS, Grinnell
1912. Proc. Biol. Soc. Washington, XXV, p. 128.
Freeman Canyon, Kern County, California.
54. PEROGNATHUS ALTICOLA ALTICOLA, Rhoads
1893. Proc. Acad. Nat. Sci. Philadelphia, p. 412.
Squirrel Inn, San Bernardino Mountains, San Bernardino County,
California.
55. PEROGNATHUS ALTICOLA INEXPECT.'iTUS, Huey
1926. Proc. Biol. Soc. Washington, XXXIX, p. 121.
Fourteen miles west of Lebec, Kern County', California.
56. PEROGN.ATHUS LAINGI, .Anderson
1932. Bull. Nat. Mus. Canada, 70, p. 100.
British Columbia, .Anarchist Mountain, near Osoyoos-Bridesville
summit, about 8 miles east of Osoyoos Lake.
57. PEROGN.ATHUS LORDI LCmDI, Gray
1868. Proc. Zool. Soc. London, p. 202.
Southern British Columbia, Canada.
58. PEROG.NATHUS LORDI COLUMBIANUS, Merriam
1894. Proc. Acad. Nat. Sci. Philadelphia, p. 263.
Pasco, Franklin Countj', Washington.
formosus Group
59. PEROGNATHUS FORMOSUS FORMOSUS, Merriam
1889. North Amer. Fauna, no. i, p. 17.
St. George, Washington Countj', Utah.
60. PEROGN.ATH17S FORMOSUS CINERASCENS, Nelson & Goldman
1929. Proc. Biol. Soc. Washington, XLII, p. 105.
Lower California : San Felipe.
61. PEROGNATHUS MESEMBRINUS, Elliot
1903. Field Columb. Mus. publ. 87, zool. ser. vol. 3, p. 251.
Palm Springs, Riverside Count\', California.
Subgenus Chaetoilipiis, Merriam
baileyi Group
62. PEROGN.ATHUS B.MLEYl BAILEYI, .Merriam
1894. Proc. Acad. Nat. Sci. Philadelphia, p. 262.
Magdalena, Sonora, Mexico.
488 I'EROGNATHUS
63. PEROGNATHUS BAll.KYl RL'DINORIS, Elliot
1903. Field Columb. Mus. publ. 74, zool. ser. vol. 3, p. 167.
San Quintin, Lower California, Mexico.
(14. PEROGNATHUS BAII.EYl INSULARIS, Tounsend
1912. Bull. Amer. Mus. Nat. Hist. XXXI, p. 122.
Tihuron Island, Gulf of California, Sonora, Mexico.
65. PEROGNATHUS BAILEYI DOMENSIS, Goldman
192S. Proc. Biol. Soc. Washington, XLI, p. 204.
Arizona ; Castle Dome, at base of Castle Dome Peak.
66. PEROGNATHUS BAILEYI HUEYI, Nelson & Goldman
192S. Proc. Biol. Soc. Washington, XLII, p. 106.
Lower California : San Felipe.
67. PEROGNATHUS BAILEYI FORNICATUS, Burt
1932. Trans. S. Diego Soc. Nat. Hist. 7, p. 164.
Montserrat Island, Gulf of California.
68. PEROGN.ATHUS KNEKUS, Elliot
1903. Field Columb. Mus. publ. 74, zool. ser. vol. 3, p. 169.
Rosarito, San Pedro Martir Mountains, Lower California.
hispidus Group
6(,. PEROGN.-\THUS HISPIDUS HISPIDUS, Baird
1857. Mamm. N. Amer. p. 421.
Charco Escondido, Tamaulipas, Mexico.
Synonym : paradoxus spilotiis, Merriam, 18S9, North .-Xmer. Fauna, no. i,
p. 25. Gainesville, Cook County, Texas.
70. PEROGNATHUS HISPIDUS PAR.-\DOXUS, Merriam
1889. North .ALmer. Fauna, no. i, p. 24.
Banner, Trego County, Kansas.
Synonym : latirostiis, Rhoads, .Amer. Nat. XXVIII, p. 185, 1S94. Rocky
Mountains.
conditi, Allen, 1894, Bull. Amer. Mus. Nat. Hist. VI,
p. 318. San Bernardino Ranch, Cochisa County,
Arizona.
71. PEROGNATHUS HISPIDUS MAXIMUS, Elliot
1903. Field Columb. Mus. publ. 87, zool. ser. vol. 3, p. 253.
Noble, Cleveland County, Oklahoma.
72. PERCJGNATHUS HISPIDUS ZACATECAE, Osgood
1900. North Amer. Fauna, no. 18, p. 45.
Valparaiso, Zacatecas, Mexico.
penicillatus Group
73. PEROGNATHUS PENTCILL.'^TUS PENICILLATUS, Woodhouse
1S52. Proc. Acad. Nat. Sci. Philadelphia, VI, p. 200.
■San Francisco Mountain, Coconino County, .Arizona.
74. PEROGNATHUS PENICILLATUS ALBULUS, Nelson & Goldman
1923. Proc. Biol. Soc. Washington, XXXVI, p. 159.
Magdalena Island, Lower California, Mexico.
I
PEROGNATHUS 489
75. PEROGNATHUS PENICILLATUS ANGUSTIROSTRIS, Osgood
1900. North Amor. Fauna, no. 18, p. 47.
Carriso Creek, Colorado Desert, Imperial Counts', California.
76. PER(K;N.VrHU.S PENICILL-VrUS PRICEI, Allen
1894. Bull. Amer. Mus. Nat. Hist. VI, p. 318.
Oposura, Sonera, Me.\ico.
77. PEROGNATHUS PENICILLATUS EREMICUS, Mearns
1898. Bull. Amer. Mus. Nat. Hist. X, p. 300.
Fort Hancock, El Paso County, Texas.
78. PEROGNATHUS PENICILLATUS AMMOPHILUS, Osgood
1907. Free. Biol. Soc. Washington, XX, p. 20.
Margarita Island, Lower California, Mexico.
79. PER0C;N,^THUS PENICILL.^TUS siccus, Osgood
1907. Proc. Biol. Soc. Washington, XX, p. 20.
Ceralbo Island, Gulf of California, Mexico.
80. PEROGNATHUS PENICILL.ATUS SERI, Nelson
191 2. Proc. Biol. Soc. Washington, XXV, p. 116.
Tiburon Island, Gulf of California, Sonora, Mexico.
Synonym: goldmani, Tovvnsend, 1912, Bull. Amer. Mus. Nat. Hist.
XXXI, p. 122. Same locality.
81. PEROGNATHUS PENICILLATUS MINIMUS, Burt
1932. Trans. S. Diego Soc. Nat. Hist. 7, p. 164.
Turner Island, Gulf of California.
82. PEROGNATHUS HELLERI, Elliot
1903. Field Columb. Mus. publ. 74, zool. ser. vol. 3, p. 166.
San Quintin, Lower California, Mexico.
83. PEROGNATHUS STEPHENSI, Merriam
1894. Proc. Acad. Nat. Sci. Philadelphia, p. 267.
Mesquite Valley, Inyo County, California.
84. PEROGNATHUS ARENARIUS ARENARIUS, Merriam
1894. Proc. Calif. Acad. Sci. ser. 2, vol. 4, p. 461.
San Jorge, near Comondu, Lower California, Mexico.
85. PEROGN.i^THUS .■\REN.'\RIUS ALBESCENS, Huey
1926. Proc. Biol. Soc. Washington, XXXIX, p. 67.
Lower California: San Felipe.
86. PEROGNATHUS ARENARIUS AMBIGUUS, Nelson & Goldman
1929. Proc. Biol. Soc. Washington, XLII, p. 108.
Lower California: Yubay, 30 miles south-east of Calamahue.
87. PEROGNATHUS ARENARIUS SUBI.UCIDUS, Nelson & Goldman
1929. Proc. Biol. Soc. Washington, XLII, p. 109.
Lower California : La Paz.
88. PEROGNATHUS PERNIX PERNIX, .Allen
1898. Bull. Amer. Mus. Nat. Hist. X, p. 149.
Rosario, Sinaloa, Mexico.
Sg. I'ER(K;N.ATHUS PERNIX ROSTRATUS, Osgood
1900. North Amer. Fauna, no. 18, p. 51.
Camoa, Rio Mayo, Sonora, Mexico.
490 PEROGNATHUS
intermedins Group
.;o. PKROGNATHLS INTERMEDIUS INTERMEDIUS, Merriam
1889. North Amer. Fauna, no. i, p. 18.
Mud Spring, Mohave County, Arizona.
Synonym; obsciinis, Merriam, i88g. North Amer. Fauna, no. 1, p. 20.
Camp Apache, Grant County, New Mexico.
<ii. PEROGN..\THUS INTERMEDIUS PHASMA, Goldman
1918. Proc. Biol. Soc. Washington, XXXI, p. 22.
Tinajas Atlas, Gila Mountains, Yuma County, Arizona.
92. PEROGNATHUS INTERMEDIUS ATER, Dice
1929. Occ. Pap. Mus. Zool. Univ. Mich. no. 203, p. 2.
New Mexico: Malpais Spring, Otero County, 15 miles west of Three
Rivers.
93. PEROGN.^THUS INTERMEDIUS RUPESTRIS, Benson
1932. Univ. Cal. Pub. Zool. XXXVIII, p. 337.
New Mexico ; Lava beds nearest to Kenzin, Dona Ana County.
94. PEROGNATHUS INTERMEDIUS NTGRIMONTIS, Blossom
1933. Occ. Pap. Mus. Zool. Univ. Mich. 265, p. i.
Arizona; Black Mountain, 10 miles south of Tucson, Pima County.
95. PEROGNATHUS INTERMEDRS CRINTTUS, Ben.wn
1934. Proc. Biol. Soc. Washington, XLVII, p. 199.
Arizona: 2 miles west of Wupatki Ruins, Coconino County.
96. PEROGNATHUS INTERMEDIUS UMBROSUS, Benson
1934. Proc. Biol. Soc. Washington, XLVII, p. 200.
Camp Verde, Yavapai County, Arizona.
97. PEROGN.^THUS INTERMEDIUS PINTCATE, Blossom
1933. Occ. Pap. Mus. Zool. Univ. Mich. 273, p. 4.
Sonora, Mexico ; Papago Tanks, Pmacate Mountains.
oS. PEROGNATHUS NELSONI NELSONI, Merriam
1894. Proc. Acad. Nat. Sci. Philadelphia, p. 266.
Hacienda la Parada, San Luis Potosi, Mexico.
<)9. PEROGNATHUS NELSONI CANESCENS, Murriam
1894. Proc. Acad. Nat. Sci. Philadelphia, p. 267.
Jaral, Coahuila, Mexico.
100. PEROGNATHUS GOLDMANI, Osgood
1900. North Amer. Fauna, no. 18, p. 54.
Sinaloa, State of Sinaloa, Mexico.
loi. PEROGNATHUS ARTUS, Osgood
1900. North Amer. Fauna, no. 18, p. 55.
Batopilas, Chihuahua, Mexico.
102. PEROGNATHUS FALLAX FALL.'^X, Merriam
1889. North Amer. Fauna, no. i, p. 19.
Reche Canyon, San Bernardino County, California.
103. PEROGN.Vl'HUS FALLAX PALLIDUS, Mearns
1901. Proc. Biol. Soc. Washington, XIV, p. 135.
Mountain Spring, halfway up east slope of Coast Range Mountains,
Imperial County. California.
PEROGNATHUS 491
104. PEROGNATHUS FALLAX INOPINUS, Nelson & Goldman
1929. Proc. Diol. Soc. Washington, XLII, p. no.
Lower California ; Turtle Bay.
105. PEROGNATHUS ANTHONYI, Osgood
1900. North Amer. Fauna, no. 18, p. 56.
South Bay, Cerros Island, Lower California, Mexico.
californicus Group
106. PEROGNATHUS FEMORALIS FEMORALIS, Allen
1891. Bull. Amer. Mus. Nat. Hist. I H, p. 281.
Dulzura, San Diego Count>', California.
107. PEROGNATHUS FEMORALIS MESOPOLIUS, Elliot
1903. Field Columb. Mus. publ. 74, zool. ser. vol. 3, p. 168.
Pinon, San Pedro Martir Mountains, Lower California, Mexico.
108. PEROGNATHUS CALIFORNICUS CALIFORNICUS, Merriam
1889. North Amer. Fauna, no. i, p. 26.
Berkeley, .Alameda County, California.
Synonym: armatus, Merriam, 1889, North Amer. Fauna, no. i, p. 27.
Mt. Diablo, Contra Costa County, California.
109. PEROGNATHUS CALIFORNICUS DISPAR, Osgood
1900. North Amer. Fauna, no. 18, p. 58.
Carpenteria, Santa Barbara County, California.
no. PEROGNATHUS CALIFORNICUS OCHRUS, Osgood
1904. Proc. Biol. Soc. Washington, XVII, p. 128.
Santiago Springs, 16 miles south-west of McKittrick, Kern County,
California.
111. PEROGN.ATHUS CALIFORNICUS BERNARDINUS, Benson
1930. Univ. Calif. Publ. Zool. XXXII, p. 449.
San Bernardino County, California.
spinatus Group
112. PEROGN.ATHUS SPIN.ATLS SPIN.ATUS, Merriam
1889. North Amer. Fauna, no. i, p. 21.
Colorado River, San Bernardino County, California (twenty-five miles
below the Needles).
113. PEROGNATHUS SPINATUS PEXINSUL.AE, Merriam
1894. Proc. Calif. Acad. Sci. ser. 2, vol. 4, p. 460.
San Jose del Cabo, Lower California, Mexico.
114. PEROGN.ATHUS SPIN.ATUS MAGDALEN.AE, Osgood
1907. Proc. Biol. Soc. Washington, XX, p. 21.
Magdalena Island, Lower California, Mexico.
115. P1;R0GNATHUS SPIN.ATUS OCCULTUS, Nelson
1912. Proc. Biol. Soc. Washington, XXV, p. n6.
Carmen Island, Lower California, Mexico.
Synonym: spinatus iielsnm, Townsend, 1912, Bull. .Amer. Mus. Nat.
Hist. XXXI, p. 122.
492 PEROGXATHUS— MICRODIPODOPS
1 16. PEROGNATHL^S SPINATL"S LAAIBI, Benson
1930. Univ. Calif. Publ. Zool. XXXII. p. 452.
Espintu Santo Island, Lower California, Mexico.
117. PKR(JGN.-\THL-S SPLN.ATUS RUFESCEN.S, Hucy
1930. Trans. S. Diego Soc. Nat. Hist. 6, p. 231.
Mouth of Palm Canyon, Borego Valley, San Diego County-. California.
U.S. PEROGN.ATHUS SPINATL'S PRIET.AE. Huey
1930. Trans. S. Diego Soc. Nat. Hist. 6, p. 232.
Lower California : 25 miles north of Punta Prieta.
iiq. PEROGN.->iTHUS SPIN,-\TUS MARCOSENSIS, Burt
1932. Trans. S. Diego Soc. Nat. Hist. 7, p. 166.
S. Marcos Island, Gulf of California.
120. PKR()C;N.\THUS SPINATUS GUARDIAE, Burt
1932. Trans. S. Diego Soc. Nat. Hist. 7, p. 165.
Angel de la Guardia Island, Gulf of California.
121. PER0GN.-\THUS SPIN.-^TUS PULLUS, Burt
1932. Trans. S. Diego Soc. Nat. Hist. 7, p. 166.
Coronados Island, Gulf of California.
122. PEROGXATHUS SPIX.\TUS SEORSC.S. Burt
1932. Trans. S. Diego Soc. Nat. Hist. 7, p. 167.
Daurzante Island, Gulf of California.
123. PEROGX.-\THUS SPINATUS L.^TIJUGUI.ARIS, Burt
1932. Trans. S. Diego Soc. Nat. Hist. 7. p. 168.
San Francisco Island, Gulf of California.
124. PEROGXATHUS BRYAXTI, Mernam
1894. Proc. Calif. Acad. Sci. ser. 2, vol. 4, p. 45S.
San Jose Island, Lower California, Mexico.
125. PEROGXATHUS MARGARITAE, Merriam
1894. Proc. Calif. ,\cad. Sci. ser. 2, vol. 4, p. 459.
Margarita Island, Lower California, Mexico.
126. PEROGXATHUS EVERMAXXI, Xelson & Goldman
1929. Proc. Biol. Soc. Washington, XLII, p. iii.
Lower California: Mejia Island, near north end of Angel de la Cluardia
Island.
Genus 2. MICRODIPODOPS, Merriam
1891. MICRODIPODOPS, Merriam, North .Amer. Fauna, no. 5, p. 115.
Type Species. — Microdipodops megacepliaJus, Merriam.
Range. — Western U.S.A.: known from California, Oregon and Nevada.
Number of Forms. — Seven.
Char.\cters. — As indicated above, this genus has usually been regarded as
most closely allied to Dipodomys, but Wood states that "many
of the resemblances to Dipodomys are obviously connected with its ricochetal
MICRODIPODOPS 493
habits and are not necessarily significant of close relationships. The foot
structure seems indicative of relationship with Perognathns." And further,
"Of the characters allying Microdipodops with Dipodomys, all but two, the trans-
verse processes of the caudal vertebrae and the process of the pubis at the anterior
end of the obturator foramen, are either obviously habitus characters or else are
shared with Perognathus too."
This genus is represented by one skull and skin only at the British ^Museum.
Essential skull characters near Dipodomys, but anterior zygomatic root not
abnormally inflated, and bullae and mastoids even larger, extending relatively
further forward in the skull, almost in contact superiorly, and nearly half as long
as the skull at greatest length. (This excessive inflation of bullae is reminiscent
of that present in some of the Old World genera as Salpingotus and Cardio-
cranius (Dipodidae), and to a lesser degree Pachvuromys (Gerbillinae).)
" Upper molars form enamel lakes by surrounding median valley as Liomys;
P.4 as in Perognathus; M. ij much reduced; cheekteeth extremely highcrowned,
but apparently not evergrowing" (Wood).
The dental pattern seems much nearer Perognathus than Dipodomys in the
one skull seen.
Tail not tufted, well haired; hindfoot greatly lengthened, sole densely hairy;
five toes present.
The bullae curve forward at the sides, and overlap the posterior portion of
the zygoma; the abnormal inflation found in this genus is carried to a further
degree than in any other Rodent genus I have examined.
Forms seen: megacephalus.
List of Named Forms
1. MICRODIPODOPS MEG.ACEPH.JiLUS MEGACEPHALUS, Merriam
1891. North .\mer. Fauna, no. 5, p. 116.
Halleck, East Humboldt Valley, Elko County, Nevada.
2. MICRODIPODOPS MEG.^CEPH.\LUS OREGONUS, Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. 127.
Lake Alvord, Alvord Desert, Harney County, Oregon.
3. MICRODIPODOPS MEG.ACEPH.-SiLUS LUCIDUS, Goldman
1926. Proc. Biol. Soc. Washington, XXXIX, p. 127.
Clayton Valley, Blair, Nevada.
4. MICRODIPODOPS MEGACEPHALUS DICKEYI, Goldman
1927. Proc. Biol. Soc. Washington, XL, p. 115.
Three miles south-east of Oasis, Mono County, California.
5. MICRODIPODOPS CALIFORNICUS, Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. 128.
Sierra Valley, near Vinton, Plumas County, California.
6. .MICRODIPODOPS P.M.LIDUS. Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. 127.
Ten miles east of Stillwater, Churchill Count}', Nevada.
494 MICRODIPODOPS— DIPODOMYS
7. MICRODIPODOPS I'OLIOXOTLS, Grinnell
1914. Univ. Calif. Publ. Zool. XII, p. 302.
McKecvcr's Ranch, 2 miles south of Benton Station, Mono County,
California.
The Dipodomys Group
Cheekteeth evergrowing, early in life completely simplified in pattern.
Anterior zygomatic root greatly enlarged on joining lachrymal. External form
bipedal saltatorial; hallux present or absent; a tendency present towards fusion
of some of the cervical vertebrae (parallel — Dipodinae).
Containing one genus only.
Genus 3. DIPODOMYS, Gray
1841. Dipodomys, Gray, .^nn. Mag. Nat. Hist. VII, p. 521.
1S67. Perodipvs, Fitzinger, Sitzber. math.-nat. CI. k. Akad. Wiss. Wien. LVI,
abth. I, p. 126. {Dipodomys agilis, Gambel.)
1890. DiPODOPS, Merriam, North Anier. Fauna, No. 3, p. 71. (Dipodomys agilis,
Gambel.)
Type Species. — Dipodomys phillipsii. Gray.
R.\NGE. — Western United States, and Mexico (in U.S.A. known from
Oregon, Wyoming, California, Utah, Arizona, Colorado, New
Mexico, Oklahoma, Texas, also Low^er California; and in Mexico south to
Vera Cruz).
Number of Forms. — Eighty-two.
Rem.^rks. — Formerly the genus " Perodipiis" was recognized to contain the
forms with the minute hallux present; but this division is not
now maintained, and it seems that the hallux may be even absent or present in
different specimens of the same species, w'hich demonstrates clearly the in-
advisability of retaining generic names for forms in which a minute and func-
tionless digit usually present may have become suppressed; as for instance such
forms as " Scartiirtis" or " Marmotops."
If two species have gone 99 per cent of the way towards suppressing a digit
and one of them goes the other i per cent and loses the digit altogether, it is
surely at very most a specific, perhaps even a racial distinction; certainly not
a generic one.
Char.\cters. — Apices of bullae in contact for a short distance behind the
posterior portion of the palate. Mastoids enormously in-
flated, taking up most of the posterior part of superior border of skull, and
projecting considerably beyond the occipital plane. Skull progressively broader
from back of lachrymals to mastoids; at all points very broad. Superior border
of zygomatic plate usually heavily ridged, but degree of spreading and ridging
of this portion of the skull variable in the different species. Skull differing
markedly from Microdipodops in the large expansion of the upper part of the
anterior zygomatic root. Nasals projecting conspicuously forward beyond the
Fig. 126. DipODO>ris merriami xielanurus, Merriam.
B.M. No. 98.3. 1. 158, cJ; X zj.
Fig. 127. DiPODOMYS merri.vmi mel^nurls, Merriam.
B.M. No. 9S.3.1.158 cJ : X 2j.
32 — Living Rodents — 1
496
DIPODOMVS
incisors; the top of the nose in the Uving animal can be seen on close inspection
to be curiously projecting forwards, no doubt caused by this bone formation.
Mandible with angular process somewhat pulled inwards, and coronoid process
small. Occipital region much reduced, between the mastoids. In D. deserti
the mastoid inflation reaches its greatest degree.
Lower incisors said to be frequently grooved in D. spcctabilis, as well as the
upper ones.
Cheekteeth evergrowing, "with tendency for thinning and breaking of
enamel on buccal and lingual margins of teeth leaving only an anterior and
Fig. 1 28. DiPODOMYS merriami melanurus, Merriam.
Cheekteeth: B.M. No. 98.3.1.158, o^; .■ 13.
posterior blade in more progressive species." Upper cheekteeth wider than
long, simplifying to a ring pattern in adult, P. 4 slightly the largest, and M.3
slightly the smallest tooth. Lower teeth like the upper series except that P.4
appears to retain one inner and one outer shallow indentation normally.
When cut, the teeth present a more complicated pattern, which is said to
vary somewhat in the different species. Calcaneum articulating w^ith the
navicular. Some of the cervical vertebrae tend to fuse.
Size largest of family. Form Dipodide. Hindlimbs elongated, foot very
long and narrow, soles hairy. Ilindfoot with three main digits; D.5 moderately
developed; hallux vestigial or absent, when present placed high on the leg, as in
Allactaga; but hindfoot differing from this genus in the length and position
of D.5. Tail considerably longer than head and body as a rule, well haired,
tufted terminally. Ear large. Forefoot with minute pollex; the two centre
digits (3 and 4) subequal and tending to be slightly longer than D.2 and D.5 so
far as seen. Claws long, sharp.
Forms seen: agilis, aitihiouus, coliiiiibiamis, deserti, exilis, leripes, iinlamiriis,
merriami, iiitratoides, ordii, richardsoni, simiolus, spectabilis, " streatori."
DIPODOMYS 497
Nine specific groups are currently recognized. As no revision of the genus
has been pubhshed, no description or key to these groups is at present avail-
able. Six of them are characterized by Grinnell, A Geographical Study of the
Kangaroo-Rats of California, Univ. Calif. Publ. Zool., XXIV, p. i, 1922.
Wood keys certain species which have been examined by him on dental
characters. His results were as follows:
"Crowns of teeth persist an appreciable time after all teeth are erupted.
E!namcl complete through life. conipactus
Enamel interrupted slightly after much wear. nilratoides
Enamel interrupted slightly after little wear. mcrriami
Crowns of teeth destroyed bv or shortly after the time the last tooth
has been erupted.
Enamel breaks small, developing late.
Unworn teeth with oval ends. ordii
Unworn teeth with square ends. agilis
Enamel breaks small to medium, with an appreciable period before
they show on grinding surface.
Unworn teeth with square ends. heermantii
Unworn teeth with oval ends. spectabilis
Enamel breaks very large, developing very early. deserti."
Further characters which 1 have compiled from Grinnell's diagnosis of his
groups are:
Maxillary arches
In heermanni group, very broad, widely spreading and as a rule with
posteroexternal angles prominent and sharp.
In ordii group, broad, spreading fairly widely as a rule, with angles more
or less well developed.
In mcrriami group, broad, widely spreading, sharply angled.
In agilis group, rather narrow, narrowly spreading, and as a rule weakly
angled.
In tnicrops group, very narrow, spreading narrowly, and very weakly
angled.
In deserti group, extremely narrow, with very narrow posteroexternal
angles, indicated but faintly.
The spectabilis group, which appears to consist of the largest forms of the
genus as a rule (other than deserti, which is nearly as large), agrees as far as I have
seen with the broad "maxillary- arch type," as heermanni.
Normally four toes are present in the hindtoot in the groups typified bv
heermanni, spectabilis, phillipsii, mcrriami, deserti; and normally five toes are
present in the groups typified by agilis, ordii and microps.
In the heermanni grouYi the metatarsal of the first toe is developed, according
to (jrinnell.
In both cranial and dental characters D. deserti appears to be very distinct
from other forms of the genus.
49S DIPODOMYS
For the characters of /). phillipsii see Merriam, Proc. Biol. Soc. Washington,
VIII, p. 83, 1893, "Rediscovery of the Mexican Kangaroo-rat Dipodomvs
phillipsii. Gray." The mastoids are described as "both actually and relatively
smaller than in any other species."
The remaining characters given by Grinnell appear for the most part
not to divide the groups ver\' clearly, nor are the measurements given by
Anthony in Field Book of North American Mammals 192S for the forms occur-
ring north of Mexico indicative of any clear size distinction between the various
groups, except that, as indicated above, the spcctabilis group and the deserti
group approach the maximum size, and that the smallest forms belong to the
merriami and the compact us groups; but even here the measurements of total
length overlap those of the smaller members of the ordii group.
List of X.amhd Forms
heermanni Group
1. DIPODOMVS HHERM.^N'NI HEERM.iWNI. Le Conte
1853. Proc. Acad. Nat. Sci. Philadelphia, VI, p. 224.
Sierra Nevada, California.
Synonym: slrecitori, Merriam, 1894, Proc. Biol. Soc. Washington,
IX, p. 113. Carbondale, Amador County, California.
2. DIPODOMYS HEERM.ANNI C.^LIFORNICUS, Merriam
1S90. North Amer. Fauna, no. 4, p. 49.
L'kiah, Mendocino County, California.
Synonym: calijormciis tritntatis, Kellogg, 1916, Uni\'. Calif. Publ. Zool.
XII, p. 366. Hellena, Trinity County, California.
californicKs pallididiis, Bangs, iSgg, Proc. New Engl. Zool.
Club. I, p. 65. Sites, Colusa County. California.
3. DIPODOMYS HEERMANNI EXIMIUS, Grinnell
1919. Proc. Biol. Soc. Washington, XXXII, p. 205.
Marysville Buttes, 3 miles north-\\'est of Sutter, Sutter County, Cali-
fornia.
4. DIPODOMYS HEERM.^NNI TULARENSIS, Merriam
1904. Proc. Biol. Soc. Washington, XVII, p. 143.
Alila, now Earlimart, Tulare County, California.
5. DIPODOMYS HEERMANNI DIXONI, Grinnel!
1919. Univ. Calif. Publ. Zool. XXI, p. 45.
Delhi, near Merced Ri\'er, Alerced County, California.
6. DIPODOMYS HEERMANNI BERKELEYENSIS, Grinnell
1919. Proc. Biol. Soc. Washington, XXXII, p. 204.
Berkeley (Head of Dwight Bay), Alameda County, California.
7. DIPODOMYS HEI'RMANNI GOLDMANI, Merriam
1904. Proc. Biol. Soc. Washington, XVII, p. 143.
Salinas, Monterey County, California.
8. DIPODOMYS HEERMANNI JOLONENSIS. Grinnell
1919. Proc. Biol. Soc. Washington, XXXII, p. 203.
One mile south-west of Jolon, Monterey County, California.
k
DIPODOMYS 499
9. DIPODOMYS HKKR.MANM SWAKTHI, Grinndl
1919. Univ. Publ. Calif. Zool. XXI, p. 44.
Seven miles south-east of Simmler, Carrizo Plain, San Luis Obispo
County, California.
10. DIPODOMV.S HKERMAN.M SAX.ATILI.S, Grinnell & Linsdale
1929. Univ. Calif. Publ. Zool. XXX, p. 453.
Mesa, near Dales, Tehema County, California.
11. DIPODOMYS HEERM.^NNI GABRIELSONI, Goldman
1925. Proc. niol. Soc. Washington, XXXVIII, p. 33.
Brownsboro, Jackson Count>', Oregon.
12. DIPODOMYS MORROENSIS, Merriam
1907. Proc. Biol. Soc. Washington, XX, p. 78.
Morro, San Luis Obispo County, California.
13. DIPODOMYS MOH.\VENSIS, Grinnell
1918. Univ. Cal. Publ. Zool. XVII, p. 428.
Warren, Kem Countj', California.
14. DIPODOMYS LELXOGENYS, Grinnell
1919. Univ. Calif. Publ. Zool. XXI, p. 46.
Pellisier Ranch, 5 miles north of Benton Station, Mono County,
California.
15. DIPODO.MYS P.-\NAMINTIN'US, Merriam
1894. Proc. Biol. Soc. Washington, IX, p. 114.
Willow Creek, Panamint Mountains, Inyo County', California.
16. DIPODOMYS STEPHENSI, Merriam
1907. Proc. Biol. Soc. Washington, XX, p. 78.
San Jacinto Valley, Riverside Count\-, California.
17. DIPODOMYS INGENS, Merriam
1904. Proc. Biol. Soc. Washington, XVII, p. 141.
Painted Rock, 20 miles south-east of Simmler, Carrizo Plain, San Luis
Obispo County-, California.
18. DIPODOMYS GRAVIPES, Huey
1925. Proc. Biol. Soc. Washington, XXXVIII. p. 83.
Santo Domingo Mission, Lower California, Mexico.
spectabilis Group
19. DIPODOMYS SPECTABILIS SPECTABILIS, Merriam
1890. North .\mer. Fauna, no. 4, p. 46.
Dos Cabezos, Cochise County, .\rizona.
zo. DIPODOMYS SPECTABILIS BAILEYI, Goldman
1923. Proc. Biol. Soc. Washington. XXXVI, p. 140.
Forty miles west of Roswell, Chaves County, Xew Mexico.
21. DIPODOMYS SPECT.\BILIS CRATODON, .Merriam
1907. Proc. Biol, Soc. Washington, XX, p. 75.
Chicalote, .^guas Calientes, \lexico.
22. DIPODOMYS SPECT.-VBILIS ZYGO.M.VriCUS, Goldman
1923. Proc. Biol. Soc. Washington, XXXVI, p. 140.
Parral, Southern Chihuahua, Mexico.
500 DIPODOMYS
2j. DIPODOMYS SPFXTABILIS PKRRI.ANDUS, Goldman
■ 933- Joum. Washington Acad. Sci. XXIII, p. 466.
Calabassus, Santa Cruz County, Arizona.
24. DII>ODOMYS SPECTABILIS CLARENCEI, Goldman
I933' Journ. Washington Acad. Sci. XXIII, p. 467.
Blanco, San Juan County, New Mexico.
25. DIPODOMYS NELSONI, Merriam
1907. Proc. Biol. Soc. Washington, XX, p. 75.
La Ventura, Coahuila, Mexico.
pliiUipsii Group
2h. DIPODOMYS PHILLIPSII, Gray
1841. Ann. Mag. Nat. Hist. VII, p. 522.
Valley of Mexico, Mexico.
Synonym: hoJticus, Wagner, 1846, Arch. Naturg. p. 176.
27. DIPODOMYS ELATOR, Mernam
1894. Proc. Biol. Soc. Washington, IX, p. log.
Henrietta, Clay County, Texas.
28. DIPODOMYS PEROTENSIS. Merriam
1894. Proc. Biol. Soc. Washington, IX, p. iii.
Perote, Vera Cniz, Mexico.
29. DIPODOMYS ORNATUS, Merriam
1894. Proc. Biol. Soc. Washington, IX, p. no.
Berriozabel, Zacatecas, Mexico.
merriaini Group
30. DIPODOMYS MERRIAMI MERRIAMI, Mearns
i8go. Bull. Amer. Mus. Nat. Hist. II, p. 290.
New Ri\er, Maricopa County, Arizona.
Synonym: tnerriawi nevadensis, Merriani, 1894, Proc. Biol. Soc. Wash-
ington, IX, p. III. Pyramid Lake, Washoe County,
Nevada.
merriami nitratus, Merriam, 1S94, Proc. Biol. Soc. Wash-
ington, IX, p. 112. Keeler, Inyo County, California.
merriami morlivallis, Elliot, 1903, Field Columb. Mus. Publ.
87, zool. ser. vol. 3, p. 250. Furnace Creek, Inyo
County, California.
merriami kerncnsis, Merriam, 1907, Proc. Biol. Soc. Wash-
ington, XX, p. 77. Onyx, Kern County, California.
ji. DIPODOMYS MERRIAMI AMBIGUUS, Merriam
1890. North .•\mer. Fauna, no. 4, p. 42.
El Paso, El Paso County, Texas.
32. DIPODOMYS MERRIAMI ATRONASUS, Mernam
1894. Proc. Biol. Soc. Washington, IX, p. 113.
Hacienda la Parada, San Luis Potosi, Mexico,
33. DIPODOMYS MERRIAMI PARVUS, Rhoads
1894. Amer. Nat. XXVIII, p. 69.
Reche Canyon, San Bernardino County, California.
DIPODOMYS 501
34. DIPODOMYS MERRIAMI SIMIOLUS, Rhoads
1893. Proc. Acad. Nat. Sci. Philadelphia, p. 410.
Agua Calicnte, near I'alni Springs, Riverside County, California.
Synonym: similis, Rhoads, 1893, Proc. Acad. Nat. Sci. Philadelphia,
p. 411. Whitewater, Riverside County, California.
35. DIPODOMYS MERRIAMI ARKNIVAGUS, Elliot
1903. Field Columb. Mus. Publ. 87, zool. ser. vol. 3, p. 249.
San Felipe, Lower California, Mexico.
36. DIPODOMYS MERRIAMI MELANURUS, Mcrriam
1893. Proc. Calif. Acad. Sci. ser. 2, vol. 3, p. 345.
San Jose del Cabo, Lower California, Mexico.
37. DIPODOMYS MERRIAMI SEMIPALLIDUS, Huey
1927. Trans. S. Diego Soc. Nat. Hist. 5, p. 65.
Santa Catarina, Lower California, Mexico.
38. DIPODOMYS MERRIAMI M.^YENSIS, Goldman
1928. Proc. Biol. Soc. Washington, XLI, p. 141.
Alamos, Sonora, Mexico.
39. DIPODOMYS MERRIAMI VULCANI, Benson
1934. Proc. Biol. Soc. Washington, XLVII, p. 181.
Towoweap Valley, Moha\e County, Arizona.
40. DIPODOMYS MERRIAMI FRENATUS, Bole
1936. Sci. Publ. Cleveland Mus. 5, no. i, p. i.
Toquerville, Washington County, Utah.
41. DIPODOMYS NITR.\TOIDES NITR.ATOIDES, Merriam
1894. Proc. Biol. Soc. Washington, IX, p. 112.
Tipton, San Joaquin Valley, Tulare County, California.
42. DIPODOMYS NlTR.\TOIDES EXILIS, Merriam
1894. Proc. Biol. Soc. Washington, IX, p. 113.
Fresno, Fresno County, California.
43. DIPODOMYS XITRATOIDES BREVINASUS, Grinnell
1920. Journ. Mamm. Baltimore, i, p. 179.
Hayes Station, Fresno County, California (19 miles south-west of
Mendota).
44. DIPODOMYS PLATYCEPHALUS, Merriam
1907. Proc. Biol. Soc. W'ashington, XX, p. 76.
Calmalli, Lower California, Mexico.
45. DIPODOMYS MARGARITAE. Merriam
1907. Proc. Biol. .Soc. Washington, XX, p. 76.
Margarita Island, Lower California, Mexico.
46. DIPODOMYS INSULARIS, Merriam
1907. Proc. Biol. Soc. Washington, XX, p. 77.
San Jose Island, Lower California, Mexico.
47. DIPODOMYS MITCHEI.LI, Mcarns
1897. Proc. U.S. Nat. Mus. XIX, p. 719.
Tiburon Island, Gulf of California, Sonora, Mexico.
502 DIPODOMYS
crdii Group
48. DIPODOMYS ORDIl ORDII, Woodhoust-
1S53. Proc. Acad. Nat. Sci. Philadelphia, VI, p. 224.
El Paso, El Paso County, Texas.
4.). DIPODOMYS ORDII COLUMBIANUS, Merriam
1894. Proc. Biol. Soc. Washington. IX, p. 115.
Umatilla, Umatilla County, Oregon.
50. DIPODOMYS ORDII MONOENSIS, Grinnell
iQig. Univ. Calif. Publ. Zool. XXI, p. 46. . ^ r^ ,■
Pfllisier Ranch, 5 miles north of Benton Station, Mono County, Cali-
fornia.
51. DIPODOMYS ORDII UTAHKNSIS, Merriam
1904. Proc. Biol. Soc. Wash. XVII, p. 143.
Ogden, Weber County, Utah.
52. DIPODOMYS ORDII CHAPMANI. Mcarns
1890. Bull. Amer. Nat. Hist. II, p. 291.
Fort Verde, Yavapai County, .Arizona.
53. DIPODOMYS ORDII OBSCURCS, .^Uen
1903. Bull. Amer. Mus. Nat. Hist. XIX, p. 603.
Rio Sestin, Durango, Mexico.
54. DIPODOMYS ORDII MONTANUS, Baird
iSc;5. Proc. Acad. Nat. Sci. Philadelphia, VII, p. 334.
Fort Massachusetts (now Fort Garland), Costilla County, Colorado.
DIPODOMYS ORDII LONGIPKS, Merr
rnani
1S90. North Amer. Fauna, no. 3, p. 72. ^ , ■
Foot of Echo Cliflfs, Painted Desert, Coconino County, Arizona.
56. DIPODOMYS ORDII LUTI-:OI,US, Goldman
1917. Proc. Biol. Soc. Washington, XXX, p. 112.
Casper, Natrona County, Wyoming.
57. DIPODOMYS ORDII RICHARDSON, -Ail™
1891. Bull. Amer. Mus. Nat. Hist. Ill, p. 277.
One of the sources of Beaver River, Beaver County, Oklahoma.
58. DIPODOMYS ORDII PALMERl. Allen
1891. Bull. Amer. Mus. Nat. Hist. Ill, p. 276.
San Luis Potosi, State of San Luis Potosi, Mexico.
5.). DIPODOMYS ORDII CCPIDINEUS, Goldman
1924. Joum. Washington Acad. Sci. XIV, p. 372. ,,..,,.. „
Kanab Wash, Arizona (southern boundary of Kainab Indian Reserva-
tion).
1.0. DIPODOMYS ORDII KVEXLS, Goldman
1933. Journ. Washington Acad. Sci. XXIII, p. 468.
Salida, Chaffee County, Colorado.
bi. DIPODOMYS ORDII CLEOMOPHILA, Goldman
1933. Joum. Washington Acad. Sci. XXIII, p. 469.
Coconino Counr\^ (Winoma), Arizona.
DIPODOMYS 503
62. DIPODOMYS ORDII NEXIl.IS, Goldman
'933- Jiiurn. Washington Acad. Sci., XXIII, p. 470.
Naturila, Montrose County, Colorado.
compactus Group
63. DIPODOMYS COMPACTUS, True
1889. Proc. U.S. Nat. Mus., II, 1888, p. 160.
Padre Island, Cameron County, Te.\as.
64. DIPODOMYS SENNETTI, .-Mien
1891. Bull. .Amer. Mus. Nat. Hist., Ill, p. 226.
Santa Rosa, Cameron County, Texas.
agilis Group
65. DIPODOMYS AGILIS AGILIS, Gambel
1848. Proc. Acad. Nat. Sci. Philadelphia, IV, p. 77.
Los Angeles, Los Angeles County, California.
66. DIPODOMYS AGILIS SIMULANS, Merriam
1904. Proc. Biol. Soc. Washington, XVII, p. 144.
Dulzura, San Diego Count\', California.
67. DIPODOMYS AGILIS PENINSULARIS, Merriam
1907. Proc. Biol. Soc. Washington, XX, p. 79.
Santo Domingo, Lower California, Mexico.
68. DIPODOMYS AGILIS CABEZONAE, Merriam
1904. Proc. Biol. Soc. Washington, XVII, p. 144.
Cabezon, San Gorgonio Pass, Riverside Count\', California.
6q. DIPODOMYS .AGILIS PERPLEXUS, Merriam
1907. Proc. Biol. Soc. Washington, XX, p. 79.
Walker Basin, Kern County, California.
70. DIPODOMYS AGILIS MARTIRENSIS, Huey
1927. Trans. S. Diego Soc. Nat. Hist. 5, p. 7.
La Grulla, Sierra San Pedro Martir, Lower California, Mexico.
71. DIPODOMYS .AGILIS L.ATIM.AXILLARIS, Huey
1925. Proc. Biol. Soc. Washington, XXXVIII, p. 84.
Two miles west of Santo Domingo Mission, Lower California; Lat.
30° 45 ' N., Long. 115" 58' W.
72. DIPODOMYS VENUSTUS VENUSTUS, Merriam
1904. Proc. Biol. Soc. Washington, XVII, p. 142.
Santa Cruz, Santa Cruz County, California.
73. DIPODOMYS VENUSTLS SANCTILUCI.AE, Grinnell
1 919. Proc. Biol. Soc. Washington, XXXII, p. 204.
One mile south of Jolon, Monterey County, California.
74- DIPODOMYS ELEPHANTINUS, Grinnell
1919. Univ. Calif. Publ. Zool. XXI, p. 43.
One mile north of Cook P.O., Bear Valley, San Benito Valley, Cali-
fornia.
504 DIPODOMYS
microps Group
75. n]P()Df)MYS MICROPS MICROPS, Akrriam
1904. Proc. liiol. Soc. Washington, XVII, p. 145.
Lone Pine, Owens Valley, Inyo County, California.
76. DIPODOMYS MICROPS PREBLEI, Goldman
if)2i. Journ. Mamm. Baltimore, 2, p. 233.
Narrows, Malheur Lake, Harney County, Oregon.
77. DIPODOMYS MICROPS CELSUS, Goldman
1924. Journ. Washington .^cad. Sci. XIV, p. 372.
Six miles north of Wolf Hole, Arizona.
7S. DIPODOMYS MICROPS LEUCOTIS, Goldman
1031. Proc. Biol. Soc. Washington, XLIV, p. 135.
Houserock Valley, Marble Canyon, Colorado Ri\er, Arizona.
7Q. DIPODOMYS MICROPS AQUILOXIUS, Willett
I935- Journ. Mamm. Baltimore, 16, p. 63.
Three miles east of Eagleville, Modoc County, California.
So. DIPODOMYS LEVIPES, Merriam
1904. Proc. Biol. Soc. Washington, XVII, p. 145.
Perognathus Flat, Panamint Mountains, Inyo County, Californa.
desert i Group
81. DIPODOMYS DESERTI DESERTI, Stephens
1887. -Amer. Nat. XXI, p. 42.
Mohave River, San Bernardino County, California.
Synonym: deserti helleri, Elliot, 1903, Field. Columb. Mus. Publ. Zoo!.
ser. vol. 3, p. 249. Keeler, Owens Lake, Inyo County,
California.
82. DIPODOMYS DESERTI SONORIENSIS, Goldman
1923. Proc. Biol. Soc. Washington, XXXVI, p. 139.
La Libertad Ranch, 30 miles east of Sierra Seri, Sonora, Mexico.
The family Heteromyidae is known fossil from the Oligocene, from North
America only. Wood recognizes five extinct genera placed in the living sub-
families, as well as a number of Oligocene types.
GENERAL WORKS OF REFERENCE
Wood, Ann. Carnegie Mus. XXIV, p. 73, 1935. (Monographic review of living and
fossil Heteromyidae.)
GoLDM.^N, North Amer. Fauna, no. 34, 191 1. Revision of the genera Heteromys and
Liomys.
Osgood, North Amer. Fauna, no. 18, 1900. Revision of the genus Perognathus.
Grinnell, a Geographical Study of the Kangaroo-Rats of California, L'niv. Calif. Publ.
Zool. XXIV, p. I, 1922.
Howell, 1932, Proc. .Amer. .Acad. .Arts Sci. Boston, LXVII, p. 378. The Saltatorial
Rodent Dipodomys, Functional and comparative anatomy of its muscular and
osseous systems.
CouES, Monograph North American Rodentia, p. 487, 1877. "Saccomyidae."
GEOMYIDAE 505
Family GEOMYIDAE
1896. Thomas: Myomorpha, part: Family Geomyidae.
1899. TuUberg: Scilromorpha, part: Geomyoidei: Family Geomyidae, part, subfamily
Geomyini.
1918. Milier & Gidley: Superfamily Sciuhoidae, part: Family Geomyidae.
1924. Winge: Family "Saccomyidae" ( -^Heteromyidae), part, Geomyini.
1928. Weber: Geo.myoidea, part: Family Geomyidae.
Geographical Distribution. — North America, and Central America;
from British Columbia through Western
and Central United States, and also from Florida and Texas, south through
Mexico to Panama.
Number of Genera. — Nine are currently recognized.
Characters. — Cheekteeth |, evergrowing in living genera, simplified in
pattern. Skull much modified for subfossorial life. Fibula
reduced, and fused with tibia high on the leg (as in Muridae). Externally
specialized for underground life; digits of hindfoot five; claws of forefoot-
strongly lengthened. Incisors thick. Infraorbital foramen always forming
long canal, "its orifice protected from muscle pressure by countersinking in an
oblique sulcus" (Miller & Gidley). Mastoids never excessively inflated. Zy-
goma robust (at any rate as compared with the Heteromyidae); the jugal pro-
gressively shortened until the zygomatic arch is in extreme forms complete
without it.
The infraorbital foramen seems more reduced in this family, and in the
Heteromyidae, than in any living Rodents.
Skull characters. According to Merriam, it may be mentioned that there
are strong cranial differences between the sexes in this group.
The skull is flattened, the bullae moderately large, with neck directed for-
ward and outward; mastoids noticeable in back view of skull between the ex-
occipital and the upper border of the supraoccipital. Squamosals largely
developed. Palate long, very narrow, a deep pit each side between last molars,
"posterior to which the palatines usually unite with the pterygoids to form a
palatopterygoid plate on each side of the posterior nares." Incisive foramina
excessively small, jugal short, never approaching lachr)mal. Anterior border
of zygomatic plate usually prominently ridged. Occipital region powerfully
developed, though relatively low; squamosals usually with strong ridges present,
which frequently unite to form a sagittal crest. Lower incisor forming powerful
process between condylar and angular processes. Coronoid higher than con-
dyle. Incisors thick, the upper ones prominently grooved, except in Thomomys.
Premaxillae very large and hea\T, nasals usually narrow.
Cheekteeth rootless and simplified, the premolar, the largest tooth in the
series, being more or less eight-shaped, with an outer and an inner fold.
Other molars ring-shaped, except sometimes M.3, which may have a posterior
heel.
As figured by Merriam, the unworn teeth are less simplified when cut than
in the adult; evidently they simplify very soon in life.
5o6 GEOMYIDAE
External form as in other underground Rodents, thickset, with eyes and
cars small. Large cheekpouches present, which open externally. Tail usually
naked, moderately hairy in the more northern species, rather longer than the
hindtoot, the tip said to be supplied with tactile nerves, and to be used as a
guide when the animal runs backwards, which according to Merriam they do
as easily as thev run forwards.
Forefoot with five digits, bearing very large and powerful claws, D.3 the
longest, the pollex and D.5 the shortest.
According to Merriam the development of the claws varies greatly, and the
hairiness on the tail of northern species varies seasonally. Hindfeet with
general arrangement of digits the same as in forefoot, but claws less enlarged.
The Geomyidae, exclusive of Thomomys, were monographed very fully by
Merriam, North American Fauna, no. 8, 1895, pp. ii-2vS. He divided the
former genus Geomvs into eight genera, based mainly on the presence or absence
of enamel plate in the upper premolar and first two molars; the number of
grooves of the incisors, and certain bones in the interior part of the skull. Most
authors have retained these genera.
The familv as a whole is so inadequately represented at the British Museum
that I have mostly to give abridged versions of Merriam's original genus descrip-
tions.
I have seen only two skulls of Cratogeomys, one of Platygeomys, four of
Orthogeomys, one of Zygogeoiitys, and few, at any rate less than ten, of Macro-
geomys and Heterogeomys.
As regards the presence or absence of enamel plate, the following teeth have
a constant pattern throughout the family, excepting the genus Thomomys:
Lower molars: a single posterior enamel plate only.
Lower premolar: four enamel plates always present.
Third upper molar: three enamel plates, one inner, one outer, one anterior.
In Thomomys, there are present in:
Lower molars : two enamel plates, an anterior and a posterior.
Lower premolar : as in the rest of the family.
Third upper molar : two enamel plates only.
It is perhaps not out of place to remark that there is a very strong resem-
blance between all the genera included in the family as regards essential cranial
and dental characters, and that two famous zoologists at least have considered
that the seven extra genera of Merriam are of at most subgeneric value only.
Key to the Genera of Geomyidae
(modified from that of ]\Ierriam)
Frontals with no marked constriction between the orbits. Orthogeomys
Frontals strongly constricted between the orbits.
Third upper molar with two enamel plates; lower molars with an
anterior enamel plate; forefoot relatively more slender, and
GEOMYIDAE: THOMOMYS 507
claws lighter (Bailey). Incisors not grooved, or with a single
fine sulcus on inner side. Thomomys
Third upper molar with three enamel plates; lower molars without
anterior enamel plate; forefoot relatively heavier. Incisors
strongly grooved.
No enamel plate on posterior surface of upper premolar.
Posterior enamel plate present on M.i and M.2.
Upper incisor bisulcate. Geomys
Upper incisor unisulcate. Pappogeomys
Posterior enamel plate absent on M.i and M.2.
Breadth of cranium across squamosals much less than
zygomatic width ; lambdoid crest not sinuous ;
angle of mandible short. Cratogeomys
Breadth of cranium across squamosals greater than zygo-
matic width ; lambdoid crest strongly sinuous ;
angle of mandible very long. Platygeomys
Enamel present on posterior surface of upper premolar.
Posterior enamel present on inner side only of JNI.i and M.2;
incisors bisulcate (zygoma complete without jugal).
Zygogeomys
Posterior enamel complete on M.i and M.z. Incisors unisul-
cate.
Postorbital process absent; palatopterygoid long and
slender, the pterygoid part narrow. Heterogeomy^s
Postorbital process strongly marked; palatopterygoid short
and broad, the pterygoid part broad. Macrogeo.mys
The key is weakened by the fact that in Orthogeomys, posterior enamel plate
may be present or absent in the upper premolar. It is, according to INIerriam,
becoming suppressed, and when present (O. laiifrons), is restricted to the inner
fourth. In other species of Orthogeomvs it is absent.
In addition to the characters indicated above, the genus Thomomys appears
to differ from the other genera, as regards those examined, in that the cheekteeth
are less rounded in aspect, though whether this character is absolutely constant
I do not know.
Genus i. THOMOMYS, VVied
1839. Thomomys, Wied, Nova Acta Phys. Med. Acad. Caes. Leop. Carol. XIX, pt. i,
P- 377.
1903. Megascaphels, Elliot, Field Columb. Mus. Publ. 76, Zool. ser. vol. 3, p. 190.
(Diplostoma bulbivortim, Richardson.) Valid as a subgenus.
Type Species. — Thomomys rtifescens, Wied.
5oS
THOMOMYS
Range. — "From the \allev of Mexico and Mount Orizaba northward to
British Cohinibia and North Saskatchewan River; and from
Pacific coast eastward to the great Plains" (Merriani).
NfMBER OF Forms. — I have Hsted one hundred and ninety-two.
Ch.\r.^cters. — As already noted, the arrangement of the enamel of the lower
molars and M.3 differs from the other genera in that there
are two enamel plates on the lower teeth instead of one, and that there are only
Fig.
29. Thomomys PERP.\LLiDis PERP.\LLIDL s, Merriam.
B.M. No. 2CJ.1 1.7.43. I; ^5.
two enamel plates on M. 3 instead of the usual number of three. There are also
two enamel plates on M.i and M.2; in P. 4 there are four enamel plates.
Upper incisor plain, or with a narrow sulcus close to the inner side of the
tooth, the main groove characteristic of other genera of the family absent. The
sulcus when present may rarely, as in inonticola, be relatively large and deep.
Molars, so far as seen, less rounded than in Geomys and allies, the upper teeth
with a tendency to point outwards at the centre of each tooth, the lower teeth
with tendency to point inwards. Lower incisor root forming large process
Fig. 130. Thomomys perpallidus perpallidus, Merriam.
B.M. No. 29.1 1.7.43, ^; -■> 2i.
b
Fig. 131. Thomomys perp.\llidus perpallidus, Merriam.
Cheekteeth: B.M. No. 29.11.7.43, V; x 6.
5IO THOMOMYS
which turns angular portion ot mandihlc noticeably outwards. The basi-
occipital seems in those seen to tend to be relatively narrower than in other
genera. Sagittal crest rarely formed in skulls seen, and more often than not
undeveloped in the large series of skulls figured by Bailey in his revision of the
genus.
Externally differing from allied genera in the relatively smaller size of the
forefoot.
The species bidhirorus, the largest known form, is separated as a subgenus
Megascaplieus by American authors, differing in the following characters from
normal Thomomys :
"Central surface of exoccipital next condyle occupied by a deep groove run-
ning obliquely to axis of skull; bullae flatter, less inflated; pterv.goids broad
laterally, concave internally, with hamuli converging at tips."
This genus was fully revised by Bailey (North Amer. Fauna, no. 39,
1915).
He divides the genus into twelve specific groups, and he keys these groups as
follows :
"Rostrum deep and evenly sloping in front of upper molars.
Pterygoid concave on inner surface and convex on outer; mammae in
four pairs. bulbirorus group
Pterygoid flat and straight.
Mammae in three pairs (inguinal 2. pectoral i). uiiihrinus group
Mammae in four pairs (inguinal 2, pectoral 2).
Skull short and wide; colour mainly dark or light ochraceous.
bottae group
Skull not conspicuously short and wide.
Skull long and narrow; colour dark. alpinus group
Skull not conspicuously long and narrow; colour mainly
pale.
Colour pale buffy yellowish, or grey and black.
Colour buffy yellowish (except apache). perpuUidiis group
Colour grey and black. iownsendi group
Colour tawny. fulviis group
Rostrum slender, abruptly arched in front oi upper molars.
Mammae in six pairs or more. talpoides group
Mammae in four or five pairs.
Mammae in five pairs (inguinal 2, pectoral 3). fossor group
Mammae in four pairs (inguinal 2, pectoral 2).
Ears rather large and rounded at tips. douglasi group
Ears large or small and pointed at tips.
Ears relatively large and pointed. montkola group
Ears relatively small and pointed. fiisciis group."
It may be noted as a matter of interest that only two groups, talpoides and
fiiscus, appear to range as far north as Canada.
Forms examined: anitae, alticola, atronirliis, annularis, altivalhs, bottae,
hulhiToriis, douglasi, >?ioiitiiola, perpallidiis, talpoides, tiAtecus, iinibnniis.
THOMOMYS SI I
List of Named Forms
(The references and type localities for all members of the family Geomyidae
are the work of Mr. G. W. C. Holt. Air. Holt has also provided mc with notes
on the relationships of the distinct species recently described.)
Subgenus Thomo?nys, Wied
tozctisendi Group
1. THOMO.MYS TOWNSKNDI TOW.XSKNDI, Bachman
1839. Joum. Acad. Nat. .Sci. Philadelphia, VIII, p. 105.
Probably Southern Idaho, near Nampa, Canyon County.
Synonym: nevadensis atrogriseus, Bailey, 1914, Proc. Biol. Soc. Wash-
ington, XXVII, p. 118. Southern Idaho.
2. THtJ.MOMV.S TOWNSENDI NEV.ADENSIS, Merriam
1897. Proc. Biol. Soc. Washington, XI, p. 213.
Austin, Lamber County, Nevada.
3. THOMOMYS RELICTUS, Grinnell
1926. Univ. Cal. Publ. Zool. XXX, p. 2.
Susanville, Lassen County, California.
bottae Group
4. THOMOMYS BOTTAE BOTTAE, Eydou.x & Gervais
1836. Mag. de. Zool. VI, p. 23.
Coast of California.
5. THOMOMYS BOTTAE LATICEPS, Baird
1855. Proc. .Acad. Nat. Sci. Philadelphia, VII, p. 335.
Humboldt Bay, Humboldt County, California.
6. THOMOMYS BOTTAE LEUCODON, Merriam
1897. Proc. Biol. Soc. Washington, XI, p. 215.
Grant Pass, Rogue River Valley, Oregon.
7. THOMOMYS BOTTAE N.^WUS, Merriam
1901. Proc. Biol. Soc. W'asbington, XIV, p. 112.
Red Bluff, Tehama County, California.
S. THOMOMYS BOTTAE ME\VA. Merriam
1908. Proc. Biol. Soc. Washington, XXI, p. 146.
Raymond, Madera County, California.
9. THOMOMYS BOTTAE .MINOR, Baiky
1914. Proc. Biol. Soc. Washington, XXVII, p. 116.
Fort Bragg, Mendocino County, California.
10. THOMOMYS BOTT.AE OIABOLl, Grinnell
1914. Univ. Cal. Publ. Zool. XII, p. 313.
Sweeney's Ranch, Diablo Range, Merced County-, California.
11. THOMOMYS BOTTAE ANGUL.ARIS, Merriam
1897. Proc. Biol. .Soc. Washington, XI, p. 214.
Los Bancs, Merced Counrv', California.
33 — Living Kodents — I
512 THOMOMYS
12. 'rHUMOMYS liOTTAK PALI.ESCENS, Uhoads
1S05. Proc. Acad. Nat. Sci. Philadelphia, p. 36.
(Jrapelands, San Bernardino Valley, San Bernardino County, Cali-
fornia.
13. THOMOMYS BOTTAK INFRAl'ALLIDL'.S, Gnnnell
1914. Univ. Cal. Publ. Zool. XII, p. 314.
Seven miles south-east of Simmlcr, Carrizo Plain, San Luis Obispo
County, California.
14. 'niOMliMYS BOTTAE NIGRICANS, Rhoads
1895. Proc. .Acad. Nat. Sci. Philadelphia, p. 36.
Witch Creel;, 7 miles west of Julian, San Diego County, California.
Synonym: aphrastiis. Elliot, 1903, Field Columb. Mus. Publ. 79, zool.
ser. vol. 3, p. 219. San Tomas, Lower California.
15. THOMOMYS BOTTAf: PASCAI.IS, Mcrriam
1901. Proc. Biol. Soc. Washington, XIV, p. iii.
Fresno, San Joaquin Valley, Fresno County, California.
16. THOMCJMYS BOTTAE PUERTAE, Gnniiell
1914. Univ. Calif. Publ. Zool. XII, p. 315.
La Puerta, 5 miles west of Vallecitos, Eastern San Diego County,
California.
17. THOMO.MYS BOTTAE ANITAE, Allen
1898. Bull. Amer. Mus. Nat. Hist. X, p. 146.
Santa Anita, Lower California, Mexico.
iS. THOMOMYS BOTTAE .M/FICOLA, Allen
1899. Bull. Amer. Mus. Nat. Hist. XII, p. 13.
Sierra Laguna, Lower California, Mexico.
19. THO.MO.MYS BOTTAE RUSSEOLUS, Nelson & Goldm.in
1909. Proc. Biol. Soc. Washington, XXII, p. 25.
San .^ngel, 30 miles west of San Ignacio, Lower California, Mexico.
20. THOMOMYS BOTT.AE ABBOTTI, Huey
1928. Trans. S. Diego Soc. Nat. Hist. 5, p. 89.
El Rosario, Lower California, Mexico.
21. THOMOM'I'S BOTTAE DEPRESSLS, Hall
1932. Univ. Calif. Publ. XXXVIII, p. 326.
Churchill County, Nevada (Dixie Meadows, at south end of Humboldt
Salt Marsh).
22. THO-MOMYS BOTT.AE CINEREUS. Hall.
1932. Univ. Calif. Publ. Zool. XXXVIII, p, 327.
Smith's Valley, Lyon County, Nevada.
zy. THOMOMYS BOTTAE EACRYM.MJS, Hall
1932. Univ. Calif. Publ. Zool. XXXVIII, p. 32S.
Arlemont, Esmeralda County, Nevada.
24. THOM(JMYS BOTTAE CCRTATUS, Hall
1932. Univ. Calif. Publ. Zool. XXXVIII, p. 329.
.San Antonio, Nye County, Nevada.
25. THOMOMYS BOTT.AE Fl'MOSUS, Hall
1932. Univ. Calif. Publ. Zool. XXXVIII, p. 329.
Moore's Creek, Nye County, Nevada.
THOMOMYS S13
26. THOMOMYS BOTTAE NANUS, Hall
1932. Univ. Calif. Publ. Zool. XXXVIII, p. 331.
Whiterock Spring, Nye County, Nevada.
27. THOMOMYS BOTTAE BUKVIDKNS, Hall
1932. Univ. Calif. Publ. Zool. XXXVIII, p. 330.
Breen Creek, Nye County, Nevada.
28. THOMOMYS BOTTAE NASUTUS, Hall
1932. Proc. Biol. Soc. Washington, XLV, p. 96.
Black River, Apache County, .Arizona.
29. THOMOMYS BOTTAE RLTDOSAE, Hall
1932. Proc. Biol. Soc. Washington, XLV, p. 96.
Ruidoso, Lincoln County, New Mexico.
30. THOMO.MYS BOTTAE LUCIDUS, Hall
1932. Proc. Biol. Soc. Washington, XLV, p. 67.
Las Palmas Canyon, Lower California, Mexico.
31. THOMOMYS BOTT.AE CAT.AVINENSIS, Huey
1932. Trans. S. Diego Soc. Nat. Hist. 7, p. 45.
Catavina, Lower California, Mexico.
32. THOMOMYS BOTTAE INGENS, Grinnell
1933. Univ. Calif. Publ. Zool. XXXVIII, p. 405.
Millux, Kern Count>', California.
33. THOMOMYS BOTTAE DIVERGENS, Nelson & Goldman
1934. Joum. Mamni. Baltimore, 15, p. 122.
Huachinera, Sonora, Mexico.
34. THOMO.MYS BOTT.AE CONVERGENS, Nelson & Goldman
'934' Joum. Mamm. Baltimore, 15, p. 123.
Hermosilla, Sonora, Mexico.
35. THOMOM\-S BOTTAE S.\XATILIS, Grinnell
1934. Proc. Biol. Soc. Washington, XLVII, p. 193.
Susanville, Lassen County, California.
36. THOMOMYS BOTTAE TRUMBLLLENSIS, Hall & Davis
1934. Proc. Biol. Soc. Washington, XLVII, p. 51.
Nixon Spring, Mount Trumbull, Mohave County, .Arizona.
37. THO.MO.MYS BOTTAE VANROSSEMI, Huey
1934. Trans. S. Diego Soc. Nat. Hist. S, p. i.
Punta Penascosa, Sonora, Mexico.
38. THO.MOM^'S BOTTAE VESCUS, Hall & Davis
1935. Univ. Calif. Publ. Zool. XL, p. 389.
Mount Jefferson, Nye County, Nevada.
3g. THOMOM\-S BOTTAE CONCISOR. Hall & Davis
1935. Univ. Calif. Publ. Zool. XL, p. 390.
Monitor Valley, Nye County, Nevada.
40. THOMOMYS BOTTAE ABSTRUSUS, Hall & Davis
1935. Univ. Calif. Publ. Zool. XL. p. 391.
Tulle Peak, Nve Count\', Nevada.
514 THOMOMVS
41. THOMOMVS B( )'l"rAi; LATUS. Hall S: Davis
1935. Univ. Calif. Publ. Zool. XL, p. 393.
Chern,' Creek, White Pine County, Nevada.
42. THOMOMYS ROTTAE KXTHNL'ATLS, Cioklman
1935. Proc. Biol. Soc. WashinRton, XLVIII, p. 149.
Willcox, Cochise County, .-Vrizona.
43. THOMOMVS liOTTAK OPULEXTUS, Goldman
1935. Proc. Biol. .Soc. Washington, XLVIII, p. 150.
Las Palonias, .Sierra County, New Mexico.
44. THOMOMVS BOTT.AE CONKIN.'^LIS, Goldman
1936. Joum. Washington .Acad. Sci. XXVI, p. 119.
Thirty-five miles east of Rock Springs, Texas.
45. THOMOMVS BOTTAE CONNECTEXS, Hall
1936. Joum. Washington Acad. Sci. XXVI, p. 296.
Clawson Dairy, 5 miles north of .Albuquerque, Bernalillo County, New
Mexico.
46. THOMOMVS BOTT.AE DESITUS, C;oldman
1936. Journ. Washington Acad. Sci. XXVI, p. 113.
Big Sandy River Valley and desert region south-eastward to Wicken-
burg, Arizona.
47. THOM(JMVS BOTT.AE GUADALUPENSIS, Goldman
1936. Journ. Washington Acad. Sci. XXVI, p. 117.
McKittrick Canyon, Guadelupe Mountains, Texas.
4S. THOMOMVS BOTTAE HOWELLI, Goldman
1936. Journ. Washington .Acad. Sci. XXVI, p. 116.
Grand Junction, Mesa County, Colorado.
4<). THO.MOMVS BOTTAE INTERN.-ATUS, Goldman
1936. Journ. Washington .Acad. Sci. XXVI, p. 115.
Salida, Chaffee County, Colorado.
50. THfJMO.MVS BOTTAE HCALPAIENSIS, Goldman
1936. Journ. Washington .A.cad. Sci. XXVI, p. 113.
Hualpai Peak, Hualpai Mountains, Mohave County, Arizona.
51. THOMOMYS BOTTAE OPTABILIS. Goldman
1916. Joum. Washington Acad. Sci. XXVI, p. 116.
Coventry, Naturita Creek Valley, Montrose County, Colorado.
52. THO.MOMVS B(JTTAE DETUMIDUS, Grinnell
1936. Univ. Calif. Publ. Zool. XL, p. 405.
One and a half miles south of town of Pistol River, Curn- County,
Oregon.
53. THOMOMVS BOTTAE ACRIROSTR.ATLS, GnnncI!
1936. Univ. Calif. Publ. Zool. XL, p. 40S.
Valley of Mad River, 7 miles above Ruth, Trimty County, California.
54. THOMOMYS BOTTAE AGRIOOLARIS, GrinncU
1916. Univ. Calif. Publ. Zool. XL. p. 409.
Stralock Farm, 3 miles west of Davis, Yolo County, California.
55. THOMOMVS BOTT.\E SIEVIECGCS, Gnnntll
1936. Univ. Calif. Publ. Zool. XL, p. 406.
Near Coyote Peak. 3,000 ft. altitude, Humboldt County, California.
k
THOMOMYS SIS
56. THOMOMYS BOTTAK PIUTKNSIS, Grinncll & Hill
1936. Proc. Biol. Soc. Washinjjton, XLIX, p. 103.
Kern Countj-, California; Krc-nch Gulch, Piute Mountains, 2 J miles
north-west of Claraville.
57. THOMOMYS MURALIS, Goldman
1936. Journ. Washington Acad. Sci. XXVI, p. nz.
Lower end of Prospect Valley, Grand Canyon, Hualpai Indian Reserva-
tion, Arizona.
58. THOMOMYS MAGDALENAK, Nelson & Goldman
1909. Proc. Biol. Soc. Washington, XXII, p. 24.
Magdalena Island, Lower Cahfomia, Mexico.
5.;. THOMOMYS ALTIVALI.IS, Uhoads
1895. Proc. Acad. Nat. Sci. Philadelphia, p. 34.
San Bernardino Mountains, California.
alpinus Group
60. THOMOMYS ALPINUS ALFLNl\S, Merriam
1897. Proc. Biol. Soc. Washington, XI, p. 216.
Big Cottonwood Meadows, 8 miles south-east of Mount Whitney Peak,
Tulare Coimty, California.
61. THOMOMYS ALPINUS AWAHNEE, Merriam
1908. Proc. Biol. Soc. Washington, XXI, p. 146.
Yosemite Valley, Mariposa County, California.
6;. THO.MO.MYS NEGLECTUS, Bailey
1914. Proc. Biol. Soc. Washington, XXVII, p. 117.
Bear Flat Meadows, .San Antonio Peak, San Gabriel Mountains, Los
Angeles County, California.
6j. THOMOMYS JACINTEUS, Gnnnell & Swarth
1914. Proc. Calif. Acad. Sci. 4, IV, p. 154.
Round Valley, San Jacinto Mountains, Riverside County, California.
64. THOMOMYS MARTIRENSIS, Allen
1898. Bull. Amer. Mus. Nat. Hist. X, p. 147.
San Pedro Martir Mountains, Lower California.
perpallidiis Group
65. THOMOMYS PERPALLIDUS PERPALLIDUS. Merriam
1886. Science, VIII, p. 588.
Palm Springs, Riverside County, California.
66. THOMO.MYS PERP.^LLIDUS .i\LBATUS, Grinnell
191 2. Univ. Calif. Publ. Zool. X, p. 172.
West side of Colorado Ri\er, at Old Hanlon Ranch, Imperial County,
California.
67. THOMO.MYS PERPALLIDUS MOH.WENSIS, Grinnell
1918. Univ. Calif. Publ. Zool. XVII, p. 427.
Moha\e River bottom near Victonille, San Bernardino County-,
California.
5i6 THOMOMYS
fiS. THOMOMYS 1>1;R1'AI.I,1DLS LHUYSONOTUS, Grmncll
9J2. Univ. Calif. Publ. Zool. X, p. 174.
Ehrenberg, Yuma County. Arizona.
hi). 'IHOMOMYS 1'1:r1'ALI,1DLS P1:KPI-;s, Mtrriam
1901. Proc. Biol. Soc. Washington, XIV, p. iii.
Lone Pine, Owen's Valley, Inyo County-, California.
Synonym: scapterus, Elliot, IQ03, Field Columb. Mus. Publ. 87, zool.
ser. vol. 3, p. 248. Hannopec Canyon, Panamint
Mountains, Inyo County, California.
70. THOMOiXnS PI-RI'ALI.IDUS AMARGOSAE, Grinnell
1921. Univ. Calif. Publ. Zool. XXI, p. 239.
Shoshone, .Amarijosa River, Inyo County, California.
71. THOMOMYS PERPALLIDUS CANUS, Bailey
1910. Proc. Biol. Soc. Washington, XXIII, p. 79.
Deep Hole, .Smoke Creek Desert, Washoe County, Nevada.
72. THOMOMYS PERPALLIDUS AUREUS, Allen
1893. Bull. Amer. Mus. Nat. Hist. V, p. 49.
Bluff City, San Juan County, Utah.
73. THO.MO.MYS PERPALLIDUS APACHE, Hailey
1910. Proc. Biol. Soc. Washington, XXIII, p. 79.
Lake la Jara, Jicarilla Apache Indian Reservation, Xew Mexico.
74. THOM(_)M"i'S PI;rP.\LL1DUS ALBICAUD.-^TUS, Hall
1930. Univ. Calif. Publ. Zool. XXXII, p. 444.
Provo, Utah County, Utah.
75. THOMOMYS PERPALLIDUS AUREIVENTRIS, Hall
1930. Univ. Calif. Publ. Zool. XXXII, p. 444.
Kelton, Bo.\ Elder County, Utah.
76. THCJMOMYS PERPALLIDUS CENTRALIS, Hall
1930. Univ. Calif. Publ. Zool. XXXII, p. 445.
Baker, White Pine County, Nevada.
77. THOMOMYS PERPALLIDUS PLANIROSTRIS, Burt
1 93 1. Proc. Biol. Soc. Washington, XLIV, p. 38.
Zion National Peak, Washington County, Utah.
7S. THO.MO.MYS PERPALLIDUS OSGOODl, Goldman
1931. Joum. Washington Acad. Sci. XXI, p. 424.
Hanksville, Wayne County, Utah.
7y. THOMt)MVS PERPALLIDUS DISSIMII.IS, Goklman
1931. Joum. Washington Acad. Sci. XXI, p. 425.
Mount Ellen, Garfield County, Utah.
So. THO-Mli.MYS PERPALLIDUS ABSONUS, Goldman
1931. Journ. Washington Acad. Sci. XXI, p. 425.
Houseruck Valley, Coconino County, Arizona.
Si. THOMOMYS I'ERPALLIDL S DEPAUPI';R,ATUS, Cirinntll & Hill
1936. Journ. Mamm. Baltimore, 17, p. 4.
ICast base Tinajas Altas Mountains, 7 miles south of Raven Butte,
^'unia County, .Arizona.
THOMOMYS S17
S2. 'I-HOMOiMVS I'KRPALLIDUS RIPARIUS, Grinnill & Hill
1936. Journ. Mamm. Baltimore, 17, p. 4.
Blythe, Riverside County, California.
83. THOMOMYS I'ROVinKNTIALIS, Griniicll
1932. Univ. Calif. Pub). Zool. XXXVIII, p. i.
Providence Range, San Bernardino Counh,-, California.
84. THOMOMYS OREOECUS, Burt
1932. Trans. S. Diego Soc. Nat. Hist. 7, p. 154.
Greenwater, Black Mountains, Inyo County', California.
8.^. THOMOMYS ARGUSKNSIS, Huey
1932. Trans. S. Diego Soc. Nat. Hist. 7, p. 43.
Argus Mountains, Inyo County, California.
86. THO.MO.MYS fHHLLEOECUS. Burt
'933- Journ. Mamm. Baltimore, 14, p. 56.
Sheep Mountains, Clark County, Nevada.
87. THOMOMYS SOLITARIUS, Grinncll
1926. Univ. Calif. Publ. Zool. XXX, p. 177.
Stewart Valley, Alineral County, Nevada.
88. THOMOMYS ALEXANDRAE, Goldman
1933- Journ. Washington Acad. Sci. XXIII, p. 464.
Rainbow Lodge, Coconino County, .Arizona.
89. THOMOMYS MELANOTIS, Grinnell
1918. Univ. Calif, Publ. Zool. XVII, p. 425.
Big Prospector Meadow, White Mountains, Mono County, California.
00. THOMOMYS CABEZONAE, Mcrriam
igoi. Proc. Biol. Soc. Washington, XIV, p. 1 10.
Cabezon, San Gorgonio Pass, California (Riverside County).
91. THOMOMYS OPER.A.RIUS, Merriam
1897. Proc. Biol. Soc. Washington, XI, p. 215.
Keeler, east side Owen's Lake, Inyo County, California.
<12. THO.MOMYS I-ATIROSTRIS, Merriam
1901. Proc. Biol. Soc. Washington, XI\', p. 107.
Little Colorado River, Painted Desert, Coconino County, Arizona.
93. THOMOMYS CERVINUS, .Allen
1895. Bull. .Amer. .Mus. Nat. Hist. VII, p. 203.
Phoenix, Maricopa County, .Arizona.
94. THOMO^n•S H.\RQUAHALAE, Grinnell & Hill
1936. Journ. Mamm. Baltimore, 17, p. 7.
Ranegras Plain, 10 miles west of Hope, Yuma County, .Arizona.
95. THOMOMYS SINALOAE, Merriam
1901. Proc. Biol. .Soc. Washington, XIV, p. loS.
.Mtata, .Sinaloa, Mexico.
fulvus Group
96. THOMOMY^S FULVUS FUI.VUS. Woodhouse
1852. Proc. .Acad. Nat. Sci. Philadelphia, VI, p. 201.
San Francisco Mountain, Coconino County, Arizona.
5i8 THOMOiMYS
Q7. THOMOMVS I'l l.VLS PKRVAGUS. Mi-rriam
1901. Proc. Biol. Soc. Washington, XIV, p. no.
Espanola, Santa Fe County, New Mexico.
.)S. THt)MO>ns i TLVUS OKSKRTORUM. AU-rnum
igoi. Proc. Biol. Soc. Washington, XIV, p. 114.
Mud Spring, Detrital Valley, Mohave County, .Arizona.
.).). THO.MOMYS Fl'I.VUS INTl- RMP:DIL"S, Mcarns
1897. Proc. U.S. Nat. Mus. XIX, p. 719.
Summit of Huachuca Mountains, Southern .Arizona.
100.
THOMnMYS FULVUS TEXKNSIS, Bailey
1902. Proc. Biol. Soc. Washington, XV, p. 119.
Head of Limpia Creek, David Mountains, Jeflf Davis County, Texas.
101. THOMOMYS FULVLS TOLTECUS, Allen
1893. Bull. Amer. Mus. Nat. Hist. V, p. 52.
Juarez, Chihuahua, Mexico.
102. THOMOMYS FULVUS SUBOIT.S, Goldman
1928. Proc. Biol. Soc. Washington, XLI, p. 203.
Old Searchlight Ferr>', Colorado River, north-west of Kingman, .Arizona.
io,v THOMOMYS FULVUS FLAVIDUS, Goldman
193T. Journ. Washington Acad. Sci. XXI, p. 417.
Parker, Yuma County, .Arizona.
104. THOMOMYS FULVUS MODICUS, Goldman
1931. Journ. Washington Acad. Sci. XXI, p. 418.
La Osa, Pima County, Arizona.
105. THOAIOMYS FULVUS CATALINAE. Goldman
1931. Journ. Washington Acad. Sci. XXI, p. 419.
Summerhaven, Pima County, Arizona.
106. THOMOMYS FULVUS GR.^HAMENSIS, Goldman
1 93 1. Journ. Washington Acad. Sci. XXI, p. 420.
Graham Mountains, Graham County, Arizona.
107. THOMOMYS FULVUS COl.LINUS, Goldman
1931. Journ. Washington Acad. Sci. XXI, p. 421.
Fly Park, Cochise County, Arizona.
loS. THOMOMYS FLil>VUS PUSILLUS. Goldman
1 93 1. Journ. Washington Acad. Sci. XXI, p. 422.
Coyote Mountains, Pima County, Arizona.
109. THOMOMYS FULVUS PERAMPI.US, (ioldman
1931. Journ. W'ashington Acad. Sci. XXI, p. 423.
Wheatfield Creek, Tunicha Mountains, .Apache County, Arizona.
no. THOMOMYS FULVUS PHASMA, Goldman
1933. Proc. Biol. Soc. Washington, XLVI, p. 72.
Two miles south of Tulc Tank, Tule Desert, .Arizona (Yuma County).
III. THOMO-MVS FULVUS SUBSIMILIS, G.ildman
1933. Proc. Biol. Soc. Washington, XLVI, p. 74.
Harquahala Mountains, A'uma County, .Arizona.
THOMOMYS 519
112. THOMOMYS FULVUS MIITABII.IS, Goldman
1933. Proc. Biol. Soc. Washington, XLVI, p. 75.
Camp Verde, Yavapai County, Arizona.
113. THOMOMY.S KITLVUS ICMOTUS, Goldman
'933- Proc. Biol. Soc. WashinRton, XLVI, p. 76.
Animas Park, Animas Mountains, Hidalgo County, New Mexico.
114. THOMOMYS MKARNSl, Bailey
1914. Proc. Biol. Soc. Washington, XXVII. p. 117.
Gray's Ranch, .Animas Valley, south-west corner of Grant County,
New Mexico.
115. THOMOMYS HAIl.KVI liAII.KYI, Mcrriam
1901. Proc. Biol. Soc. Washington. XIV, p. 109.
Sierra Blanca, El Paso County, Texas.
116. THOMOMYS BAILEYl TULAKOSAE, Hall
1933- Univ. Calif. Publ. Zool. XXX.VIII, p. 411.
Tularosa, Otero County, New Mexico.
117. THOMOMYS L.ACHUGUILLA LACHUGUILLA, Bailey
1902. Proc. Biol. Soc. Washington, XV, p. 120.
Near El Paso, El Paso County, Texas.
118. THO.MOMYS LACHUGl'ILLA LIMITARIS, Goldman
1936. Journ, Washington Acad. Sci. XXVI, p. 118.
Four miles west of Boquillas. Brewster County, Texas.
iiy. THOMOMYS PECTORALIS, Goldman
1936. Journ. Washington ."Vcad. Sci. XXVI, p. 120.
Vicinity of Carlsbad Cave, Carlsbad Cave National Monument, Eddy
County, New Mexico.
120. THOMOMYS BURTI BURTI, Huey
1932. Trans. S. Diego Soc. Nat. Hist. 7, p. 158.
Madera Canyon, Santa Rita Mountains, Arizona (Santa Cruz County).
121. THOMOMYS BURTI QL'ERCINUS. Burt &: Campbell
1934- Journ. Mamm. Baltimore, 15, p. 150.
Pena Blanca Spring, Pajarito Mountains, Arizona (near Mexican
boundary).
122. THOMOMYS BURTI PROXIMUS, Burt & Campbell
1934. Joum. Mamm. Baltimore, 15, p. 151.
Santa Rita \Iountains, Pima Countj', Arizona.
iimbrimis Group
123. THO.MOMYS UMBRINUS UMBRINUS, Rich.irdson
1829. Fauna Borcali-.Americana, vol. i, p. 202.
.Southern Mexico ; probably the vicinity of Boca del Monte, Vera Cruz.
.24. THOMOMYS UMBRINUS ORIZ.\BAE, Mernam
1893. Proc. Biol. Soc. Washington, VIII, p. 145.
Mount Orizaba, Puebla, Mexico.
125. THOMOMYS UMBRINUS PEREGRINUS, Merriam
1893. Proc. Biol. Soc. Washington, VIII, p. 146.
Salazar, State of .Mexico, Mexico.
520 THOMOMVS
126. THOMOMYS tMBRlNUS ALBIGLLARIS, Nelson i: Goldman
1934. Journ. Manim. Baltimore, 15, p. 106.
El Chico, Sierra de Pachuca, Hidalgo, Mexico.
127. THOMOMYS UMBRINUS MARTINENSIS, Nelson &• (ioldman
1934. Journ. Mamm. Baltimore, 15, p. 108.
San Martin Te.\nielcuan, Puebla, Mexico.
128. THOMOMYS UMBRINUS TOI.UCAE, Nelson & Goldman
IQ34. Journ. Mamm. Baltimore, 15, p. log.
Volcano of Toluca, Mexico.
120. THOMO.MYS UMBRINl'S VUI.CANIUS, Nelson & Goldman
1934. Journ. Mamm. Baltimore, 15, p. 109.
Popocatepetl, Mexico.
130. THOMOMYS UMBRINUS SUPERNUS, Nelson & Goldman
1934. Journ. Mamm. Baltimore, 15, p. no.
Santa Rosa, Guanajuato, Mexico.
131. THO.MOMYS UMBRINUS POTOSINUS, Nelson & Goldman
1034. Journ. Mamm. Baltimore, 15, p. iii.
La Tinaja, San Luis Potosi, Mexico.
132. THOMOMYS UMBRINUS ATR<~iDORSALlS, Nelson &: Goldman
1934. Journ. Mamm. Baltimore, 15, p. iii.
-Alvarez, San Luis Potosi, Mexico.
133. THOMOMYS I'MBRINUS ZACATECAE, Nelson & Goldman
1934. Journ. Mamm. Baltimore, 15, p. 112.
Berriozabel, Zacatecas, Mexico.
134. THOMOMYS UMBRINITS ENIXUS, Nelson & Goldman
1934. Journ. Mamm. Baltimore, 15, p. 112.
Sierra Moroni, Zacatecas, Mexico.
135. THOMOMYS UMBRINUS CRASSIDENS, Nelson & Goldman
1934. Journ. Mamm. Baltimore, 15, p. 113.
Sierra de Valparaiso, Zacatecas, Mexico.
136. THOMOMYS UMBRINUS CHIHUAHUAE, Nelson & Goldman
1934. Journ. Mamm. Baltimore, 15, p. 114.
■Sierra Madre, Chihuahua, Mexico.
137. THOMOMYS UMBRINUS DURANGI, Nelson & Goldman
1934. Journ. Mamm. Baltimore, 15, p. 114.
Durango, Durango, Mexico.
13S. THOMOMYS UMBRINUS EVEXUS, Nelson & Goldman
1934. Journ. Mamm. Baltimore, 15, p. 115.
Mount San Gabriel, Durango, Mexico.
13Q. THOMOMYS UMBRINUS M.\DRENSIS, Nelson & Goldman
1934. Journ. Mamm. Baltimore, 15, p. 115.
Pilares Canyon, Colonia Garcia, Mexico.
140. THOMOMYS UMBRINUS CALIGINCXSUS, Nelson & Goldman
1934. Journ. Mamm. Baltimore, 15, p. 116.
Altamirano, Sierra Madre, Chihuahua. Mexico.
THOMOMYS 521
.41. THOMOMYS UMBKINUS CHIRICAHUAE, Nelson & Goldman
1934. Joum. Mamm. Baltimore, 15, p. 117.
Chiricahua Mountains, Arizona.
142. THOMOMYS UMBRINL'S SONORIKNSIS, Nelson & CJoldman
1934. Joum. Mamm. Baltimore, 15, p. 118.
Chinapa, Sonora River Valley, Mexico.
143. THOMOMYS UMBRINUS EXTIMUS, Nelson & Goldman
1934. Joum. Mamm. Baltimore, 15, p. iig.
Colomo, Nayarit, Me.\ico.
144. THOMOMYS UMBRINUS MUSCULUS, Nelson & Goldman
1934. Joum. Mamm. Baltimore, 15, p. iig.
Sierra de Teponahuaxtla, Nayarit, Mexico.
145. THOMOMYS UMBRINUS EXIMIUS, Nelson & Goldman
1934. Joum. Mamm. Baltimore, 15, p. 118.
Choix, Sinaloa, Mexico.
146. THOMOMYS NELSON I, Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. 109.
Parral, Chihuahua, Mexico.
147. THOMOMYS SHELDONI, Bailey
1915. North Amer. Fauna, no. 39, p. 93.
Santa Teresa, Nayarit, Mexico.
148. THOMOMYS GOLDMAN!, Merriam
1901. Proc. Biol. Soc. Washington, XIV. p. 108.
Mapimi, Durango, Mexico.
i4y. THOMOMYS PERDITUS, Merriam
1 901. Proc. Biol. Soc. Washington. XIV, p. 108.
Lanipazos, Nuevo Leon, Mexico.
150. THOMOMYS .ATROVARIUS, Allen
1S98. Bull. Amer. Mus. Nat. Hist. X, p. 148.
Tatemeles, Sinaloa, Mexico.
151. THOMOMYS SIMULUS SIMULUS, Nelson & Goldman
1934. Joum. Mamm. Baltimore, 15, p. 120.
Alamos, .Southern Sonora, Mexico.
152. THOMOMYS SIMULUS PARVICEPS, Nelson & Goldman
1934. Journ. Mamm. Baltimore, 15, p. I2i.
Chacala, Western Durango, Mexico,
talpoides Group
153. THOMOMYS TALPOIDES TALPOIDES, Richardson
1828. Zool. Joum. vol. 3, p. 518.
Near Fort Carlton, Saskatchewan, Canada.
Synonym: borealis, Richardson, 6th .\nn. Kept. Brit. .'\ssn. for 1836,
V. pp. 150, 157, 1837 (fide Bailey).
154- THOMOMYS TALPOIDES RUFESCENS. Wied
1839. Nova Acta. Phys. Med. .Acad. Caes. Leop. Carol. XIX, pt. i, p. 378.
Minnetaree Village, now Old Fort Clark, about 6 miles south of
Stanton, Mercer Count\', North Dakota.
522 THOMOMYS
155. THOMO.MVS TAI.l'OIUl-S LI.USIUS, Coiies
1875. Proc. Acad. Nat. Sci. Philadelphia, p. 138.
BridKcr Pass, iS niiles south-west of Rawlins, Carbon County, Wyo-
minji.
.56. THOMOMYS TAI.POIDKS BLLLATUS, Ba.ley
1014. Proc. Biol. Soc. Washington, XXVII, p. 115.
PowdervUle, Custer County, Montana.
157. TH(nK.)MYS TALPOIDES NEBULOSCS, Bailey
1914. Proc. Biol. Soc. Washington, XXVII, p. 116.
Jack Boyden's Ranch, Sand Creek Canyon. Crook County, Wyoming.
158. THOMOMYS T.ALPOIDES CARYI. Bailey
1914. Proc. Biol. Soc. Washington, XXVII, p. 115.
Head of 'I'rappcr Creek, Bighorn Mountains, Bighorn County, Wyo-
ming.
159. THOMOMYS TALPOIDES PRYORI, Bailey
1914. Proc. Biol. Soc. Washington, XXVII, p. 116.
Sage Creek, Pryor Mountains, Montana.
1(10. THOMOMYS T.ALPOIDES AGRESTIS, Merriam
190S. Proc. Biol. Soc. Washington, XXI, p. 144.
Medano Ranch, San Luis Valley, Colorado.
ihi. THOMOMYS TALPOIDES MACROTIS, F. Miller
1920. Proc. Colorado Mus. 0, p. 41.
D'.Arcy Ranch, Parker, Douglas County, Colorado.
162. THOMOMYS COLUMBIANUS, Bailey
1914. Proc. Biol. Soc. Washington, XXVII, p. 117.
Touchet, Walla Walla County, Washington.
153. THOMOMYS OCirS, Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. 114.
Six miles south-west of Old Fort Bridger, Uinta County, Wyoming.
164. THOMOMYS IDAHOENSIS, Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. 114.
Birch Creek, Fremont County, Idaho.
165. THOMOM^'S PYCiMAEL'S, Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. 115.
Montpelier Creek, Bear County, Idaho.
fossor Group
If. I.. THOMOMYS F'OSSOR, Allen
1893. Bull. Amer. Mus. Nat. Hist. V, p. 51.
Florida, La Plata County, Colorado.
ib7. THOMOMYS HRIDGERI, Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. 113.
Six miles south-west of Old Fort Bridger, Uinta County, Wyoming.
16S. THOMOMYS ITNTA, Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. 112.
Black's I'^ork, north base of Crilbert's Peak, Uinta Mountains, Summit
County, Utah.
THOMOMYS 523
i6(). THOMOMYS QUADRATUS QUADRATUS, Merriam
1897. Proc. Biol. Soc. Washington, XI, p. 214.
The Dalles, Wasco County, Oregon.
170. THOMOMYS QUADIl.-\TL"S FISHKRI, Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. iii.
Beckwith, Sierra Valley, Plumas County, California.
171. THOMOMYS QUADR.A1US WALLOWA, Hall & Orr
1933. Proc. Biol. Soc. Washington. XLVI, p. 41.
Catherine Peak, Telocaset, Oregon.
172. THOMO.MYS QUADRATUS MONOENSIS, Huey
1934. Trans. S. Diego Soc. Nat. Hist. 7, p. 373.
Dexter Creek Meadow, Mono County, California.
173. THOMOMYS KALCIFER, Grinnell
1926. Univ. Cal. Publ, Zool. XXX, p. 180.
Bell's Ranch, Nye County, Nevada.
douglasii group
174. THOMOMYS DOUGLASII DOUGLASII, Richardson
1829. Fauna Boreali-.\mericana, vol. i, p. 200.
Near mouth of Columbia River, Oregon.
175. THOMOMYS DOUGLASII YELMKNSIS, .Merriam
1899. Proc. Biol. Soc. Washington, XIII, p. 21.
Penino, Yelm Prairie, Thurston County, Washington.
176. THOMOMYS DOUGLASII OREGONUS, Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. 115.
Ely, near Oregon City, Clackamas County, Oregon.
177. THOMOMYS DOUGLASII TACOMENSIS, Taylor
1919. Proc. Biol. Soc. Washington, XXXII, p. 169.
Si.\ miles south of Tacoma, Pierce County, Washington.
178. THOMOMYS DOUGLASII MELANOPS, Merriam
1899. Proc. Biol. Soc. Washington, XIII, p. 21.
Timberline at head of Soleduc River, Olympic Mountains, Clallam
Counrv', Washington.
179. THOMOMYS DOUGLASII SHAWI, Taylor
1921. Proc. Biol. Soc. Washington, XXXIV, p. 121.
Owyhigh Lake, Mount Rainier, Pierce County, Washington.
180. THO.MOMYS DOUGL-^^SII LIMOSUS, Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. 116.
White Salmon, Gorge of the Columbia, Klickitat County, Washington.
181. THOMOMYS NIGER, Merriam
1901. Proc. Biol. Soc. Washington, XIV, p. 117.
Seaton, Umpqua River, Douglas County, Oregon.
monticola Group
182. THOMO.MYS MONTICOLA MONTICOLA, Allen
1893. Bull. Amer. Mus. Nat. Hist. V, p. 48.
Mount Tallac, El Dorado Counr\'. California.
524 THOMOMYS— GEOMYS
183. THOMOMYS MONTICOLA MAZAMA, Mernam
1S97. Proc. Biol. Soc. Washington, XI, p. 214.
Anna Creek, near Crater Lake, Mt. Mazama, Klamath County, Oregon.
1)54. THOMOMYS MONTICOLA PINETORUM, Mcrruim
iiigg. North Amer. Fauna, no. 16, p. 97.
Sisson, Siskiyou County, California.
Synonym: monticola premaxillaris, Grinnell, 1914, Univ. Calif. Publ.
Zool. XH, p. 312. Two miles south of S. Yolla Bolly
Mountain, Tehama County, California.
185. THOMOMYS MONTICOLA NASICUS, Mernam
iSgy. Proc. Biol. Soc. Washington, XI, p. 216.
Farewell Bend, Deschutes River, Crook County, Oregon.
iSh. THOMOMYS MONTICOLA HELLERI, Elliot
1903. Field Columb. Mus. Publ. Zool. Ser. 74, vol. 3, p. 165.
Goldbeach, Rogue River, Curry County, Oregon.
ftiscus group
1S7. THOMOMYS FUSCCS FUSCUS, Merriam
1 89 1. North Amer. Fauna, no. 5, p. 70.
Mountains at head of Big Lost River, Custer Coimty, Idaho.
iSS. THOMOMYS FUSCUS SATURATUS, Bailey
1914. Proc. Biol. Soc. Washington, XXVII, p. 117.
Silver, near Saltese, Coeur D'Alene Mountains, Missoula County,
Montana.
i8q. THOMOMYS FUSCUS LORINGI, Bailey
1914. Proc. Biol. Soc. Washington, XXVII, p. 118.
South Edmonton, .A,lberta. Canada.
lyo. THOMOMYS FUSCUS MYOPS. Mernam
1 90 1. Proc. Biol. Soc. W^ashington, XIV, p. 112.
Conconulh', east base of Cascade Range, Okanogan Count\', Washing-
ton.
191. THOMOMYS HESPERUS, Mcvriam
1901. Proc. Biol. Soc. Washington, XIV, p. 116.
Tillamook, Tillamook County, Oregon.
Subgenus Megascapheus, Elliot
192. THOMOMYS BULBIVORUS, Richardson
1829. Fauna Boreali-.4mericana, vol. i, p. 206.
Columbia River, probably near Portland, Oregon.
Genus 2. GEOMYS, Rafinesque
1817. Geomys, Rafinesc|ue, .-Xmer. Monthly Mag. II, p. 45.
Type Species. — Geomxs pinctis, IXifmeiquc—Mus iuza. Barton.
Range. — "Middle U.S.A. from Red River Valley in North-west Minnesota
and north-eastern North Dakota to Mexican boundary along Rio
Grande; also southern half of Alabama and Georgia, and northern half of
b
GEOMYS 525
Florida." Evidently now known to extend across the border into North-eastern
Mexico (Tamaulipas).
Number of Forms. — Nineteen.
Characters. — Upper premolar with posterior enamel plate absent. M.i
and M.2 with two enamel plates each, the posterior one in-
complete. M.3 with no well-marked heel. Upper incisors with two grooves,
the main one placed centrally. " Orbitosphenoid small and narrow, not reaching
alisphenoid . . . alisphenoid short posteriorly . . . pterygoids large, always
forming more than half of palatopterygoid extensions."
(Further characters appertaining to the detail cranial characters of this genus
will be found in p. 109 of Merriam's monograph (reference on p. 506).)
IMcrriam gives a most interesting account of the activities of a live Pocket-
Gopher in which the animal's method of using the cheekpouches is fullv ex-
plained. He states: "A live Geomys from Vernon, Texas, has been carefully
observed for the purpose of ascertaining how the reser\-e food is placed in the
cheekpouches. The animal soon became sufficiently tame to eat freely from
hand, and was commonly fed bits of potato of which he was particularly fond.
The manner of eating was peculiar and interesting, and showed an ability to use
the huge forefeet and claws in a way previously unsuspected. After satisfying
the immediate demands of hunger it was his practice to fill one or both cheek-
pouches. . . . The usual course is as follows: a piece of potato, root or other food
is seized between the incisor teeth and is immediately transferred to the fore-
paws which are held in a horizontal position, the tips of the claws curving
toward one another. If the food required reduction in size, the trimming is
done while held in this position. The piece is passed rapidly across the side of
the face with a sort of w iping motion which forces it into the open mouth of the
pouch. Sometimes a single rapid stroke with one hand is sufficient, at other
times both hands are used ... in such cases the long claws of one hand are used
to draw down the lower side of the opening, while the food is poked in with the
other. It is obviously impossible for the animal to pass food from the mouth
to the pouches without the aid of the foreclaws. The most remarkable thing
connected with the pouches is the way they are emptied. The forefeet are
brought back simultaneously along the sides of the head until they reach a
point opposite the hinder end of the pouches. They are then pressed firmly
against the head and carried rapidly forward. In this way the contents of the
pouches are promptly dumped in front of the animal."
In connection with the last paragraph, it may be of interest to note that I
have seen very much the same way of emptying pouches practised by the Golden
Hamster, Mcsocricetiis auratus; though in this case of coiu-se the cheekpouch
does not open externally.
The tail is already noted in these animals as being used apparently for
feeling purposes ; the tails of those examined appear, though sometimes naked,
to be quite devoid of any scales.
The species of Geomys were revised by IMerriam, who recognized three
specific groups : the tuza group, in which the tail is more naked than in the others ;
526 GEOMYS
the hiirstiriiis group, containing a largf form, differing from the other species in
cranial characters, among which the sagittal crest is said to be more strongly
developed; and the hrcviceps group, containing forms which Merriam regards
as the most priniiti\e ot the genus.
Forms examined; tuza, biirsaiius, persoiialus, jlondtunis.
List of Xamkd Forms
(For status of " Gciimvs iiwxicaiius," Lichtenstein, nee Kerr, Anim. Kingd.,
p. 207, 1792, see Merriam, iS()5, North Amer. Fauna, no. cS, p. 201.)
tuza Group
1. GEOMYS TUZA TUZA, liarton
1806. Voigt's Mag. der Naturkunde, vol. 12, p. 488.
Georgia.
Synonym: pinetis, Rafincsquc, Amer. Monthly Mag. II, p. 45, 1817.
2. GEOMYS TUZA MOBILENSIS, Mtrnani
1895. North Amer. Fauna, no. 8, p. iig.
Point Clear, Mobile Bay, Baldwin County, Alabama.
3. GEOMYS FLORIDANUS FLORIDANUS, Audubon & Bachnian
1854. Quadr. N. Amer. vol. 3, p. 242.
St. Augustine, St. John County, Florida.
4. GEOMYS FLORIDANUS AUSTRINUS. Bancs
1898. Proc. Boston Soc. Nat. Hist. 28, p. 177.
Belleair, Hillsboro County, Florida.
5. GEOMYS COLONUS, Bangs
1898. Proc. Boston Soc. Nat. Hist. 28, p. 178.
St. Mary's, Camden County, Georgia.
6. GEOMYS CUMBERLAND! US. BanL's
1898. Proc. Boston Soc. Nat. Hist. 28. p. 180.
Stafford Place, Cumberland Island, Camden County, Georgia.
hiirsarius Group
7. GEOMYS BL RSARIUS BURSARIUS, Shaw
1800. Trans. Linn. Soc. V, p. 227.
Upper Mississippi Valley; exact locality unknown.
Svnonym; fiisco, Rafinesque, 1817, Amer. Monthly Mag. II, p. 45.
cinerca. Rafinesque, 1817. Same reference.
ianadensis, Lichtenstein. kh. .Akad. Berlin, p. 20, 1S22.
saccatus, Mitchill, N. Y'. Med. Repos. xxi, 1821.
(The above names quoted as synonyms by Trouessart.)
S. GEO.MYS BL'RS.ARIUS ILLINOENSIS, Komarck & Spencer
1931. Journ. Mamm. Baltimore. 12, p. 405.
Momence, Kankalee County, Illinois.
hievlceps Group
0. GE(JMYS l,L'Ti:SCENS, Merriam
1890. North Amer. Fauna, no. 4, p. 51.
Birdwood Creek, Lincoln County, Western Nebraska.
GEOMYS— I'APPOGEOMYS 527
10. GEOMYS RRKVICEI'S liRICVICKPS, Baird
1855. Proc. Acad. Nat. Sci. Philadelphia, VII, p. 335.
Prairie Mcr Rouge, Morehouse Parish, Louisiana.
11. GEOMYS BKIvVICEl'S SAGI TTALIS, Mcrrimi.
1895. North Amer. Fauna, no. 8, p. 134.
Clear Creek, Galveston Bay, Galveston County, Texas.
12. GEOMYS BREVICEPS ATTWATEUI, Merriam
i8<)5. North Amer. Fauna, no. 8, p. 135.
Rockport, Aransas Count>-, Texas.
13. GEOMYS BRKVICEPS LEANENSIS, Baiky
1905. North Amer. Fauna, no. 25, p. 129.
Llano, Llano County, Texas.
14. GEOMYS TEXENSIS, McTriam
1895. North Amer. Fauna, no. 8, p. 137.
Mason, Mason County, Texas.
15. GEOMYS AREN'ARIUS .^iRENARUS, Merriam
1895. North Amer. Fauna, no. 8, p. 139.
El Paso, El Paso County, Texas.
16. GEOMYS ARENARIUS BREVIRO.STRIS, Hall
1932. Proc. Biol. Soc. Washington, XLV, p. 97.
Tularosa, Otero County, New Mexico.
17. GEOMYS PERSON.aiTUS PERSONATUS, True
1889. Proc. U.S. Nat. Mus. II, 1888, p. 159.
Padre Island, Cameron County, Texas.
18. OEOMYS PERSON.VrLS FALL.\X, Merriam
1895. North Amer. Fauna, no. 8, p. 144.
South side of Nueces Bay, Cameron County, Texas.
19. GEOMYS PERSOX.^TUS TROPICALIS. Goldman
1915. Proc. Biol. Soc. Washington, XXVIII, p. 134.
.^Ita Mira, Tamaulipas, Mexico.
Genus 3. PAPPOGEOMYS, Merriam
1895. Pappogeomys, Merriam, North Amer. Fauna, no. 8, p. 145.
Type Species. — Geo7iiys bidlcri, Thomas.
Range. — Jalisco, Mexico.
Number of Forms. — Two.
Char.'\cters. — Arrangement of enamel plale on molars and premolar as
in Geotnys. M.3 an imperfectly developed double prism, a
sulcus on outer side, behind which crown narrows to form a moderate heel.
Upper incisors one-grooved. No sagittal crest developed. Zygomata slender.
"Palatoptervgoids little more than vertical lamellae. Orbitosphenoids broad,
articulating firmlv with alisphenoids" (compare Geoiiiys). For further cranial
details see p. 145 of .Merriam's inonograph.
Forms examined: bulleri.
34 — Living Rodents— I
528 PAPPOGEOMYS— CRATOGEOMYS
List of Named Forms
1. PAPl'OGi;OMVS BILLKRI, Thomas
1892. Ann. Mat;. Nat. Hist. 6, X, p. 196.
Near Talpa. Sierra de Mascota, Jalisco, Mexico.
Synonym: nchoiii, Merriam, 1S92, Proc. Biol. Soc. Washington, VII,
p. 164. Sierra Nevada of Colima, Jalisco, Mexico.
2. P.-\PP()GKOI\IVS ,\I.BINASL'S, Mirnani
1895. North .Amer. Fauna, no. 8, p. 149.
Atemajac, Ciuadalajara, Jalisco, Mexico.
Genus 4. CRATOGEOMYS, Merriam
1895. Cratogeomys, Merriam, North Amer. Fauna, no. 8, p. 150.
Type Species. — Gcoiiivs iiicrricuni, Thomas.
Range.- — "Great Plains of U.S.A., from Arkansas River in Eastern Colorado
southward, and eastern tableland of Alexico, to extreme southern
edge, in the states of Mexico and Puebla" (Merriam).
Number of Forms. — Twenty-four are now named.
Charalters. — Enamel of upper premolar as in Geomys, but M.i and M.2
with one enamel plate each, the posterior one absent. M.3
with deep sulcus on outer side. Upper incisors one-grooved. " Orbitosphenoids
short and broad, articulating with alisphenoid anteriorly." (Further cranial
details will be found in Merriam's monograph, p. 150.) Sagittal crest usually
developed, apparently.
Differing from Platv<;eoiiiys, which has a similar arrangement of enamel
plate, in the following characters : breadth of cranium posteriorly much less
than zygomatic breadth ; breadth of occipital plane not more than twice its
height ; lambdoid crest broadly convex posteriorly; mandible including incisors
longer than broad ; squamosal expansion chiefly towards median line.
Cratogcumvs is the only genus besides Thomomys and Geomys which ranges
north into the United States, the other six being entirely either Mexican or
Central American.
Forms examined: mcrriaiiii, estor, castanops (skin).
List of Named Forms
1. CRATOGEOMYS MERRIAMl MERRIAMI, Thomas
1893. Ann. Mag. Nat. Hist. 6, XII, p. 271.
Southern Mexico, probably in the Valley of Mexico.
2. CR.VroGKOAIVS MERRIAMI SACCHARALIS, Nelson & Goldman
1934. Proc. Biol. Soc. Washington, XLVII, p. 149.
Atlixco, Puebla, Mexico.
.1. CRATOGEOMYS MERRIAMI IROLONIS, Nelson & Goldman
1934. Proc. Biol. Soc. Washington, XLVII, p. 150.
Irolo, Hidalgo, Mexico.
CRATOGEOMYS S*9
4. CRATOGKOMYS PEKO'IENSIS, Mirriam
1895. North Amer. Fauna, no. 8, p. 154.
Cofrc de Perote, Vera Cruz, Mexico.
5. CRATOGKOMYS ESTOR. Mcrriam
1895. North Amer. P'auna, no. 8, p. 155.
Las Vigas, Vera Cruz, Mexico.
6. CRATOGEOMY.S OREOCETES, Mcrriam
1895. North Amer. Fauna, no. 8, p. 156.
Mt. Popocatepetl, State of Mexico, Mexico.
7. CR.^TOGEOMYS PEREGRINUS, Mcrriam
1895. North Amer. Fauna, no. 8, p. 158.
Mt. IztaccihuatI, State of Mexico, Mexico.
8. CRATOGEOMYS CAST.ANOPS CASTANOPS, Baird
1852. Report Stansbury's Exped. to Great .Salt Lake, p. 313.
Prairie Road to Bent's Fort, near present town of Las Animas, Bent
Count>', Colorado.
Synonym: ctarkii, Baird, Proc. Acad. Nat. Sci. Philadelphia, 1855,
P- 332.
g. CRATOGEOMYS CASTANOPS GOLDMANI, Merriam
1895. North Amer. Fauna, no. 8, p. 160.
Canitas, Zacatecas, Mexico.
10. CRATOGEOMYS CASTANOPS PERPLANl'S, Nelson & Goldman
1934. Proc. Biol. Soc. Washington, XLVH, p. 136.
Tascosa, Oldham Count\', Texas.
11. CRATOGEOMYS CASTANOPS LACRIMALIS, Nelson & Goldman
1934. Proc. Biol. Soc. Washington, XLVH, p. 137.
Roswell, Chaves County, New Mexico.
12. CR.ATOGEOMYS CASTANOPS HIRTUS, Nelson & Goldman
1934. Proc. Biol. Soc. Washington, XLVII, p. 138.
Albuquerque, New Mexico.
13. CRATOGEOMYS CASTANOPS ANGUSTICEPS, Nelson & Goldman
1934. Proc. Biol. Soc. W'ashington, XLVII, p. 139.
Eagle Pass, Texas.
14. CR,-\TOGEOMYS CASTANOPS CONSITUS, Nelson & Goldman
1934. Proc. Biol. Soc. Washington, XLVII, p. 140.
Gallego, Chihuahua, Mexico.
15. CRATOGEOMYS CASTANOPS TAMAl.'LIPENSIS, Nelson & Goldman
1934. Proc. Biol. Soc. Washington, XLVII, p. 141.
Matamoros, Tamaulipas, Mexico.
16. CR.ATOGEOMYS CASTANOPS EXCELSUS, Nelson & Goldman
1934. Proc. Biol. Soc. Washington. XLVII, p. 143.
San Pedro, Coahuila, Mexico.
17. CR.\TOGEO.MYS CASTANOPS SL BSIMIS, Nelson & Goldman
1934. Proc. Biol. Soc. Washington, XLVII, p. 144.
Jaral, Coahuila, Mexico.
530 CR.VrOGEOMYS— PLATYGEOMYS
iS. CRATOGi;()MVS CASTANUPS SL'BNL'BILIS, Nelson & Goldman
1934. Proc. Binl. Soc. Washington, XLVII. p. 145.
Garneros, Coahuila, Mexico.
10. CRA'l'GGEOMVS fASTANOPS I'l.AMFRGNS, Nelson & Goldman
11)34. Proc. Biol. Soc. Washington, XLVII, p. 146.
Miquihuana, Nuevo Leon, Mexico.
::o. CRATOGEOMYS CASTANOPS CONVEXUS, Nelson S; Goldman
1934. Proc. Biol. Soc. Washington, XLVII, p. 142.
Las Vacas, Coahuila, Mexico.
2T. CR.VroGKOMYS CASTANOPS PKRIDONEUS, Nelson & Gc.ldman
1934. Proc. Biol. Soc. Washington, XLVII, p. 148.
Rio Verde, San Luis Potosi, Mexico.
zz. t'RATOCJEOMYS CASTANGPS RUBELLUS, Nelson ii Goldman
1934. Proc. Biol. Soc. Washington, XLVII, p. 147.
Solfdad, San Luis Potosi, Mexico.
23. C'RAT0C;E()MYS FULVESCENS FULVESCENS, Merriam
1895. North Amer. Fauna, no. 8, p. 161.
Chalchicomula, Puebla, Mexico.
24. LR.XTOCiEOMYS FULVESCENS SCBLUTEUS, Nelson & (;oldman
1934. Proc. Biol. Soc. Washington, XLVII, p. 152.
Perote, Vera Cruz, Mexico.
Genus 5. PLATYGEOMYS, JNIerriam
1895. Platyceomvs, Merriam, North Amer. Fauna, no. 8, p. 162.
Type Species. — Geomvs gyinnunis, Merriam.
Ranc;e. — " Southf rn border of Mexican tableland ; States of C'oiima, Jalisco,
Michoaean, Mexico and Hidalgo."
Number of Forms. — Six.
Char.^cters. — Arrangement of enamel plate on upper premolar and molars
as in Cratogenmxs. Upper incisor one-grooved. "Hinder
part of cranium extraordinarily broad and flat, the great breadth chiefly due to
the lateral expansion of the squamosals, which completely arch over and conceal
the postglenoid notch. Zygomatic arches massive, broadly spreading anteriorly;
jugal normaliv large, forming an important part of the arch. Pterygoids vertical
lamellae with inferior border everted. Orbitosphenoids larger than in Crato-
gcoinvs, but not normally articulating with alisphenoid. . . . Lambdoid crest
sinuous, presenting three posterior concavities. Mandible very much broader
than long, the angular process extremely long and spreading, reaching so far out
laterally that the knob of the root of the incisor is midway between condyle
and end of angidar process."
Forms examined: juniosiis.
List of Named Forms
I. PLATVGEGMVS GVMNCRUS, Merriam
1892. Proc. Biol. Soc. Washington, VII, p. i6().
Zapotlan, Jalisco, Mexico.
PLATYGEOMYS— ORTHOGEOMYS 531
2. PLATYGKOMVS TYI.OKHINUS TYLORHINUS, Merriam
1895. North Amtr. Fauna, no. 8, p. 167.
Tula, Hidalgo, Mexico.
3. PLATYGKOMYS TYI.OKHINLS ANGUSTIROSTRIS, Mirriam
1903. Proc. Biol. Soc. Washington, XVI, p. 81.
Patamban, Michoacan, Mexico.
4. PL.IlTYGEOMYS NEGLECTUS, Merriam
1902. Proc. Biol. Soc. Washington, XV, p. 68.
Cerro de la Calentura, about 8 miles north-west of Pinal de .\moles,
Queretaro, Mexico.
5. PL.VrYGEOMYS PLANICEPS, Merriam
1895. North Amer. Fauna, no. 8, p. 168.
Volcan of Toluca, State of Mexico, Mexico.
(). PLATYGEOMYS FUMOSUS, Merriam
1895. North Amer. Fauna, no. 8, p. 170.
Colima Citj', Colima, Mexico.
Genus 6. ORTHOGEOMYS, Merriam
1895. Orthogeo.mys, Merriam, North Amer. Fauna, no. 8, p. 172.
Type Species. — Geomys scalops, Thomas.
Range. — Oaxaca and Chiapas in extreme Southern Me.xico and adjacent
parts of Guatemala. Ranging into Guerrero; and known also from
Honduras and Salvador.
Number of Forms. — Twelve.
Characters. — Upper premolar with three or four enamel plates, the posterior
one, when present (latifrons only, according to NIerriam),
restricted to the inner fourth. M.i and M.2 with two enamel plates each.
Upper incisors one-grooved. M.3 with backwardly projecting heel. "Skull
as a whole much elongated. Frontals extraordinarily broad and flat, much
broader than muzzle, with sides nearly parallel. Zygomata narrow or moder-
ately spreading. Angle of mandible short. Orbitosphenoids rather large,
articulating with anterior part of alisphenoids; . . . third endoturbinal larger
and much broader than second, a unique condition in the family. Palatoptery-
goids long and narrow, of nearly equal breadth throughout." (For further
cranial details sec p. 173 of Merriam's monograph.) Sagittal crest so far as seen
developed. The members of this genus are large forms, with coarse pelage,
apparently easily distinguishable from other genera by their unconstricted
frontals.
Forms seen : scalops, grandis.
List of Named Forms
I. ORTHOGEOMYS CUNICUI.US. Elliot
1905. Proc. Biol. Soc. Washington, XVIII, p. 234.
Yautepec, Oaxaca, Mexico.
53:: ORTHOGEOMYS— HETEROGEOMYS
z. ORTHOGEOMYS SCALOPS, Thomas
1594. Ann. Mag. Nat. Hist. 6, XIII, p. 437.
Tehuantcpec, Oaxaca, Mexico.
,^ ORTHOGIXJMYS GR.ANDIS GR.ANDIS, Thomas
i8q3. Ann. Mag. Nat. Hist. 6, XII, p. 270.
Dut'nas, Guatemala.
4. ORTHOGEOMYS GR.^NOIS .^LLENI, Nelson & Goldman
1930. Joum. Mamm. Baltimore, 11, p. 156.
.Acapulco, Guerrero, Mexico.
?. OR'I'HOGEOMYS GRANDIS FELIPENSIS, Nelson & Goldman
1930. Joum. Mamm. Baltimore, 11, p. 157.
Cerro San Felipe, Oaxaca, Mexico.
6. ORTHOGEOMYS GRANDIS G UERRERENSIS, Nelson & Goldman
1930. Joum. Mamm. Baltimore, 11, p. 158.
EI Limon, La Union, Guerrero, Mexico.
7. ORTHOGEO.MYS GR.ANUIS VLI.CANI. Nclsoji &• Goldman
1930. Proc. Biol. Soc. Washington, XLIV, p. 105.
Volcan Santa Maria, Guatemala.
8. ORTHt)GEt)MYS GRANDIS PLUTO. Lawrence
1933. Proc. New England Zool. Club, 13, p. 66.
Cerro Cantoral, Tegucigalpa, Honduras.
9. ORTHOGEOMYS GRANDIS ANNEXLS, Nelson S: Goldman
1933. Proc. Biol. Soc. Washington, XLVI, p. 195.
Tuxtla Gutierrez. Chiapas, Mexico.
10. ORTHOGE(J.MYS NELSONI, Merriam
1595. North Amer. Fauna, no. 8, p. 176.
Mount Zempoaltepec, Oaxaca, Mexico.
11. ORTHOGEOMYS LATIERO.NS, Mcrnam
1895. North Amer. Fauna, no. 8, p. 178.
Guatemala; exact locality unknown.
12. ORTHOGEOMYS PYGACANTHLS, Dickey
1928. Proc. Biol. Soc. Washington, XLI, p. 9.
Cacaguatique, San Miguel, El Salvador.
Genus 7. HETEROGEOMYS, Merriam
1895. Heterogeomys, Merriam, North Amer. Fauna, no. 8, p. 179.
Type Species. — Geomys hispidus, Le Conte.
Range. — Mexico, Vera Cruz to Campechc, extending south into Guatemala.
Number of Forms. — Seven.
Characters. — Upper premolar with four enamel plates, the posterior one
restricted to the inner half. M.i and M.2 with two enamel
plates. M.'? a double prism, crown longer than broad, the heel well developed.
"Skull as a whole high and narrow; frontal broad and flat; . . . temporal de-
pressions anteriorly defining a well-marked frontal shield. Inlerior surface of
HETEROGEOMYS— MACROGEOMYS 533
palatoptcrygoid cuneate-linguatc, long and slender, the palatal arms much
elongated; pter\'goid part small and postpalatal pits deep. Orbitosphenoids
shield-shaped, rather narrow and long, not articulating with alisphenoids. . . .
Mandible with angular process short." Pelage harsh.
Forms seen: hisptdus, torridus.
List of Named Forms
1. HETEROGEOMYS HISPIDLS HISPIDUS, Le Come
1852. Proc. Acad. Nat. Sci. Philadelphia, VI, p. 158.
Near Jalapa, Vera Cruz, Mexico.
2. HETKROCiEO.MVS HISPIDU.S CONC.>\VUS, Nelson & Goldman
1929. Proc. Biol. Soc. Washington, XLII, p. 148.
Pinal de Amoles, Queretaro, Mexico.
3. HETEROGEOMYS HISPIDUS ISTHMICUS, Nelson & Goldman
1929. Proc, Biol. Soc. Washington, XLII, p. 149.
Jaltipam, Vera Cruz, Mexico.
4. HETEROGEOMYS HISPIDUS YUCATANENSIS, Nelson & Goldman
1929. Proc. Biol. Soc. Washington, XLII, p. 150.
Campeche, Mexico.
5. HETEROGEOMYS HISPIDUS CHIAPENSIS, Nelson & Goldman
1929. Proc. Biol. Soc. Washington, XLII, p. 151.
Tenejapa, San Cristobal, Chiapas, Mexico.
6. HETEROGEOMYS LANIUS, Elliot
1905. Proc. Biol. Soc. Washington, XVIII, p. 235. •
Xuchil, Vera Cruz, Mexico.
7. HETEROGEOMYS TORRIDUS. Merriam
1895. North -Amer. Fauna, no. 8, p. 183.
Chichica.\tle, Vera Cruz, Mexico.
Genus 8. MACROGEOMYS, Merriam
1895. MACROGEOMYS, Merriam, North Amer. Fauna, no. 8, p. 185.
Type Species. — Geomys heterodtis, Peters.
Range. — Now known from Nicaragua, Costa Rica and Panama.
Number of Forms. — Nine.
Characters. — Upper premolar with four enamel plates, the posterior
restricted to the inner third. M.3 with deep outer sulcus
and elongated heel which is greatly developed, attaining the maximum size
known in the familv. Upper incisor one-grooved. "Frontals broad, flat,
depressed or concave along median line, deeply excavated laterally between
orbits, the notch immediately succeeded by a strongly developed postorbital
process. Palatoptcrygoids broad, short, and truncated posteriorly, the hori-
zontal part composed above wholly of palatal, the pterj'goid simply capping the
534 MACROGEOMYS— ZYGOGEOMYS
end and abruptly upturned at right angles. Braincase rising above posterior
root of zygoma. . . . Tlic occipital plane is flat and slopes strongly forwards as in
Hetcrvgeomvs."
There is a marked tendency apparently for the jugal to become abnormally
reduced; in more than one specimen seen it is nearly as in Zxgogeoinvs in that
the zygomatic arch appears complete without it; also in costariceinis, according
to Merriam, the zygoma is in this condition.
Species of this genus are large forms. \ sagittal crest is evidently developed
in the adult.
Forms seen : ctiviitor, lietcrudiis, (li)lithiicc[>lialus.
List of Named Forms
1. MACROGHOMYS HETERODUS, Peters
1S64. Monatsber. k. preuss. Akad. Wiss. Berlin, p. 177.
Costa Rica.
2. MACROGEOMYS DOLICHOCEPHALUS, Merriam
1895. North Amer. Fauna, no. S, p. i8g.
San Jose, Costa Rica.
3. MACROGEOMYS C.^VATOR. Bangs
1902. Bull. Mus. Comp. Zool. Harvard Coll. XXXIX, p. 42.
Boquete, Chiriqui, Panama.
4. MACROGEOMYS DARIENSIS, Goldman
1912. Smiths. Misc. Coll. LX, no. 2, p. 8.
Cana, mountains of Eastern Panama.
5. MACROGEOMYS PANSA, B.ings
1902. Bull. .Mus. Comp. Zool. Harvard Coll. XXXIX, p. 44.
Bogava, Chiriqui, Panama.
6. MACROGEOMYS COSTARICENSIS, Merriam
1895. North .'^mcr. Fauna, no. 8, p. 192.
Pacuare, Costa Rica.
7. MACROCJEOMYS CHERRIEI, Allen
1S93. Bull. .Amer. Mus. Nat. Hist. V, p. 337.
Santa Clara, Costa Rica.
S. M..\CROGE()MYS M.^TAGALPAK, Allen
1910. Bull. Amer. Mus. Nat. Hist. XXVIII, p. 97.
Pena Blanca, Matagalpa, Nicaragua.
0. MACROCJEO.MYS UNDERWOODI, Osyood
1931. Field. YIus. Publ. Zool. iS, p. 143.
Alto de Jabillo Pirris, Western Costa Rica.
Genus 9. ZYGOGEOMYS, Merriam
1895. Zygogeomvs, Merriam, North Anicr. Fauna, no. 8, p. 195.
Type Species. — Zygogeomys trichopus, Merriam.
ZYGOGEOMYS 535
Range. — Mexico: "The Sierra Madre of Michoacan, from Patzcuaro to
Nahuatzin; strictly limited to the pine zone, between altitudes of
6,800 and 9,500 feet" (Miller).
Number of Forms. — One.
Char.acters. — Upper premolar with four enamel plates, the posterior
restricted to the lingual third. M.i and M.2 with two enamel
plates each. M.3 with crown longer than broad, the heel well developed;
upper incisors two-grooved. "Cranium as a whole long and narrow; zygomata
not widely spreading; . . . zygomatic arch normally complete without the jugal,
maxillary and squamosal arms in contact above it; jugal inferior, rudimentarv,
and chiclly external. Rostrum long and narrow. . . . Pterj'goids vertical
lamellae as in Tfwmomys, meeting or nearly meeting in median line behind
palate. . . . Mandible rather long and slender, as in Geomys biirsarius. Orbito-
sphenoids relatively larger than in any other genus in the family, closmg upper
part of the sphenoidal fissure except for a foramen at apex, and ankvlosed
broadly with the alisphenoid as in some species of Tluimomxs." Sagittal crest
well developed in the one skull seen.
Forms examined: Irichopus.
List of Named Forms
I. ZYGOGEOMYS TRICHOPUS, Merriam
1895. North .^mer. Fauna, no. 8, p. 196.
Nahuatzin, Michoacan, Mexico.
GEOMYIDAE:
SPECIAL WORKS OF REFERENCE
Merriam, North American Fauna, no, 8j pp. 1 1-258, 1895. Monograph and full revision
of all forms then known of all genera except Thomomys.
Bailey, North American Fauna, no. 39, Nov. 15, 1915. Full revision of Thomomys
with figures of skulls of all leading species.
Coles, Monograph of North American Rodents, 1877: Geomyidae : p. 607.
The family Geomvidae is known from the Oligocene, but apparently not
outside the North American continent.
Superfamily ANOMALUROIDAE
As here understood this contains one family, the Anomaluridae, with two
widely separated subfamilies the .\nomalurinae and the Idiurinac, the last
regarded as of family rank by Miller & (Jidley.
Family ANOMALURIDAE
1896. Thomas: Anomaluri : Family .Anomaluridae.
1899. Tullbcrg: Sciiroon.athi : Myomorpha : .\nomaluroidei. part: Family .Anoma-
luridae.
536 ANOMALURIDAE
ii)i8. Miller &: Ciidley: Supcrfamily Dipodoidae, part, Family Anomaluridae (Anoma-
Itiriis) ; Family Idiuridae, the latter with subfamilies Idiurinae (Idiuriis), and Zen-
kercllinac (Zenkcrclla).
1924. Winge: Family Anomaluridae, part, Anomalurini.
192.S. Weber: Anomaluroidea, part, Family Anomaluridae.
Geographical Distribution. — .\frica. Western and Central : from Sierra
I.eone to Uganda, Tanganyika and Northern
Rhodesia.
Number of Genera. — Four.
Characters. — Zygomasseteric structure (so far as it afi'ects shape of skull)
essentially as defined by Miller & Gidley for their Super-
family Dipodoidae, "Masseter lateralis superficialis with anterior head not
distinct, this portion of muscle attaching along a considerable area on anterior
border of zygoma; zygomatic plate nearly horizontal, always narrow and com-
pletely beneath infraorbital foramen; angular portion of mandible not distorted
outwards to permit . . . passage of branch of masseter lateralis." Infraorbital
foramen large, transmitting muscle, extremely enlarged in the subfamily
Idiurinae; skull with no special peculiarities in the typical subfamily; zygomatic
region comparatively unmodified; jugal long; auditory bullae not excessively
inflated.
Dental formula i. }, c. \\, p. ], m. a=20, the cheekteeth rooted, flatcrowned,
relatively brachyodont, characterized by a pattern of narrow cross ridges
separating wide recurrent spaces.
Externally considerably modified for arboreal life; form usually Pteromyine;
a flying-membrane usually attached to sides; underside of tail with scaly
outgrowths on posterior portion near the body. Tibia and fibula (so far as
known) not fully fused.
Remarks. — The Anomaluridae have by some authors been placed in the
neighbourhood of the Squirrels. But so far as zygomasseteric
structure is concerned there exists between the two families a very wide
distinction.
Divisions. — Two well-marked subfamilies may be recognized, as indicated
above. Although the Idiurinae are very much more specialized
in cranial characters, the two groups present many features in common, so that
it seems undesirable to refer them to two separate families.
Key to the Subfamilies of ANOMALimiDAE
Infraorbital foramen moderate in size, and zygomatic plate not projected
forwards conspicuously, the upper and lower zygomatic roots
above one another. Cheekteeth not reduced in size, less brachy-
odont. Incisors not greatly thickened. Palate not excessively
narrowed. Bullae more inflated. Anterior point of masseteric
insertion on mandible beneath hinder part of M.i ( Miller & Gidley).
Subfamily Anomalurinae
[Anoinalunis, Anomalurops)
ANOMALURINAE: ANOMALURUS 537
Infraorbital foramen extremely enlarged, owing to zygomatic plate being
projected forwards to a point nearly immediately behind the in-
cisors. Cheekteeth extremely brachyodont, greatly reduced in size.
Incisors much thickened from before backwards. Palate much
narrowed. Bullae less inflated. Anterior point of masseteric
insertion on mandible in front of P. 4 (Miller & Gidley).
Subfamily Idilrinae
{Zenkerella, Idiiirus)
Subfamily ANOMALURINAE
Geographical Distribution. — As in the family Anomaluridae.
Number of Genera. — Two.
Characters. — Cranial characters as indicated in the above key. Flying-
membrane always present, the bony outgrowth supporting
it anteriorly rising from the elbow; the membrane extending to the hindfoot;
a well-developed interfemoral membrane present. Tail well haired, relatively
long though usually somewhat shorter than head and body; usually thickly
bushv terminally, and well haired; two thick rows of jagged scales are present
on underside near the body and extend downwards for about a quarter or a
third of its length. Digits of hindfoot five, the hallux shortest, the others sub-
equal, each digit bearing a prominent curved claw; functional digits of forefoot
four, all well developed, and with prominent claws.
This group has been revised recently by Riimmler. He recognizes two
genera, as here retained, and four distinct species only.
Key to the Gener.\ of the .Anomalurinae
Cheekteeth with three transverse ridges surrounded by four depressions;
tail broader, terminal tuft stronger. Anomalurus
Cheekteeth with two transverse ridges surrounded by three depressions;
tail narrowed, terminal tuft weaker. Anomalurops
Genus i. ANOMALURUS, Waterhouse
1842. Anomalurvs, Waterhouse, Proc. Zool. Soc. London, p. 124.
1915. Anomalubodon, Matschie, S.B. Ges. Nat. Fr. Berlin, p. 350. (A. auzembergeri,
Matschie --= A. peli, Temminck.)
1915. Anomalurella, Matschie, S.B. Ges. N'at. Fr. Berlin, p. 350. (A. pusillus,
Thomas.)
Type Species. — Anomalurus fraseri, Waterhouse.
Range. — About as in the family Anomaluridae; perhaps not extending
farther w'est than the Gold Coast.
Nltmber of Forms.— Sixteen.
Fig. 132. Akcimah Ris kraseri jacksoni, de Winton.
B.M. No. 35.1.6.82, V; ■ it.
Fig. 133. Anomalurus fr.iseri jacksoni, de Wiiiton.
B.M. No. 35.1.6.82, $; X li.
ANOMALURUS
539
Characters. — -Skull with moderate frontals, little constricted; short nasals
thick, widely open anteriorly; frontals depressed, and bor-
dered by moderately developed ridges which may appear as a small postorbital
process, behind which the ridges tend to extend over the braincase, but show
no signs of coming together. Incisive foramina medium, in front of toothrows.
Palate tends to be slightlv constricted anteriorly. Bullae relatively large, well
inflated. Jugal long, forming most of zygoma, but not extending to lachrymal,
its posterior upper border somewhat raised up. Infraorbital foramen moder-
ately large, well open; zygomatic plate completely beneath it, and narrow.
Mandible without special peculiarities.
Fic. 134. .\nomah:rus fraseri jacksoni, de Winton.
Cheekteeth; B.M. No. 35.1.6.82, ?; X 5.
Upper cheekteeth with three narrow transverse ridges cutting the tooth
into four wide depressions ; flatcrovvned in adult, the pattern ultimately obliter-
ated. Lower cheekteeth like the upper series, but also with one prominent
external fold to each tooth.
Externally as described above ; fur soft ; ear prominent.
'I'lillberg iTientions that the pairs of ribs in the specimens examined bv him
were nine (the highest number he quotes tor any Rodent), and that the palmar
and plantar tubercles are more numerous than in any other Rodent he examined.
Anomalurus lias been revised by Riimmlcr (Sitz. Ber. Ges. Nat. Fr. Berlin,
1933, p. 389). He recognizes three species, which in his key are based entirely
on size.
540 ANOMALURUS
Hindfoot more than 65 : peli.
Hindfoot less than 40: pusilltis.
Hindfoot longer than 40, shorter than by. Jriiseri.
Measurements of condylobasal length, total length, and upper toothrow
will he found for the three species in the above-mentioned paper.
A. peli may be noted for its specialized black and white colour pattern.
A. hatesi, de Winton, he synonyniizes with piisilliis; there appears a tangible
difference in the size of the bullae in the type skulls of the two species ; and also
apparently in the colour. Mr. R. W. Hayman has suggested to me that in his
opinion batesi should not be regarded as a synonym, and I propose to retain it
here as a valid race.
Forms seen: batesi, cinereiis, erythrotiotus, fraseri, griselda, impcrator, jacksoni,
jordani, ncavci, iiigrensis, orientalis, peli, perustiis, pusilliis.
List of Named Forms
(The references and type localities for all members of the Anomaluridae are
the work of Mr. R. W. Hayman.)
fraseri Group
1. ANOMAIA'RLS I-'RASERl FRASERI, Waterhouse
1S42. Proc. Zool. Soc. London, p. 124.
Fernando Po.
Synonym: dcrbianus. Gray, 1S42, .Ann. Mag. Nat. Hist. X, p. 262.
squamicamlus. Schinz, 1845, Syn. Mamm. 2, p. 58.
chrysop/iacmis, Dubois, 1S8S, Bull. Soc. Zool. Paris, XIII,
p. 23-
beldeiii, du Chaillu, 1S61, Proc. Boston Soc. Nat. Hist. VII,
P- 3°3-
2. ANOMALURUS FRASERI L.'^TICKPS. Aguilar-Amat
1922. Bull. Inst. Catal. N.H. Barcelona, 2, 2, p. 52, pi. i.
Fernando Po.
(A synomym oi fraseri fraseii according to G. M. .Allen. 1939.)
3. ANOMALURUS FRASERI GRISELDA, Dollman
1 914. Ann. Mag. Nat. Hist. 8, XIV, p. 490.
Bitye, South Cameroons.
4. .-XNO.MALURUS FRASERI KRYTHRONOTUS, Milne-Edwards
1879. C.R. Acad. Sci. Paris, LXXXIX, p. 771.
Gaboon.
5. ANOMALURUS FRASERI NIGRENSIS, Thomas
1904. Abstr. Proc. Zool. Soc. London, no. 10, p. 12.
.Abutschi, Lower Niger.
6. .\NOM.\LURUS FRASI-RI IMPKR..\T(JR, Dollman
191 1. .Ann. Mag. Nat. Hist. 8, VIII. p. 257.
Bibianaha, Gold Coast.
7. ANOMALURUS FRASERI FORTIOR, Lonnberg
1917. Stockholm Vet. Akad. Handl. 58, no. 2, p. 66.
Central Africa: no exact locality; specimens quoted from Masisi, near
Kivu, and forest west of Beni.
ANOMALURUS— ANOMALUROPS 541
8. ANOMALURUS FRASF.IU PKRUSTUS, Thomas
1914. Ann. Mag. Nat. Hist. 8, XVIII, p. 235.
River Lubefu, 75 miles north of Lusambo, S. Congo.
9. ANOMALURUS FRASKRI NKAVKI, DoUman
1909. Ann. Mag. Nat. Hist. 8, III, p. 351.
Katanga, South Congo.
10. ANOMALURUS FRASKRI JORDAXI, St. I.l-kct
1935. Nov. Zool., XXXIX, p. 251.
Near Ainboin, Angola.
11. ANOMALURUS FRASKRI JACKSONI, de Winton
1898. Ann. Mag. Nat. Hist. 7, I, p. 251.
Entebbe, Uganda.
12. ANOMALURUS FRASKRI ORIKNTALIS, Peters
1880. Monats. Bcr. .\kad. Berlin, XLV, p. 164.
Zanzibar (? Mainland).
13. ANOMALURUS FRASKRI CINKRKUS, Thomas
1895. Ann. Mag. Nat. Hist. 6, XV, p. 188.
Upper Rovuma River, near Lake Nyasa.
pelii (iroup-
14. ANO\LALURUS PliLII, Temminck
1845. Verhandl. Nat. Ges. Ned. Bez. I, 2, p. ioq.
Dabocrom, Clold Coast.
Synonym: auzembergeri, Matschie, IQ14, S.B. Ges. Nat. Fr. Berlin,
p. 350. Near boundary between Liberia and Ivory
Coast; middle Ca\alle River.
pusillns Group
15. ANOMALURUS PUSILLUS I'USILLUS, Thomas
1887. Ann. Mag. Nat. Hist. 5, XX, p. 440.
Bellima, Monbuttu, N.-E. Congo.
16. ANOMALURUS PUSILLUS B.ATESI, de Winton
1897. .\nn. Mag. Nat. Hist. 6, XX, p. 524.
Como River, Gaboon.
Genus 2. ANOM.ALUROPS, Matschie
1914. Anomalurops, Matschie, Sitz. Ber. Ges. Nat. Fr. Berlin, p. 351.
Type Species. — Anomalurus beecrofti, Fraser.
R.\NGE. — Africa: Sierra Leone to Consjo (Ituri).
Number of Forms. — Four.
Characters. — Liice Anomalurus, but cheekteeth with, in the upper series,
only two transverse ridges and three depressions, the anterior
and posterior depression isolated, but the centre one remaining widely open.
Lower cheekteeth with four depressions, the anterior and posterior ones isolated ;
the other two caused bv one external and one internal fold.
542 ANOMALUROPS— IDIURINAE
Externally difftring IVdiii Anoinaliirus in the tail, which is much narrower,
and less bushv at the end.
Remarks. — The considerable difference in the pattern of the cheekteeth
seems to warrant the separation of the two genera.
Forms seen: biccrofii, ar^eiitenSy citritius, " lainger," "fiilgens."
(Revised hv Riinimler, Sitz. Ber. Ges. Nat. Fr. Berlin, 1933, p. 1S9.)
List of Named Forms
1. .\XOMALUROPS BEECROFTI BF.IX'ROFTI, Fraser
1S52. Proc, Zool. Soc. London, p. 17, pi. 32.
Fernando Po.
Synonym: fiilgens. Gray, Ann. Mag. Nat. Hist. 4, III, p. 467, 1869.
Ciabnon.
/anigt'f, IVinminck, 1853, Esq. Zool. Cote de Guine, p. 149.
Gold Coast.
2. ANO.MALLROPS BEECROFTI CHAPINI, Alien
1922. Bull. Amer. Mus. Nat. Hist. XLVH, p. 65.
Medje, Ituri.
:,. ANOMALUROPS BEECROFTI CFFRINUS, Thomas
1916. .Ann. Mag. Nat. Hist. 8, XVHI, p. 236.
Benito River, Spanish Guinea.
4. .-WOMAFUROPS BEECROFTI ARGENTEU.S, Schwann
1904. Ann. Mag. Nat. Hist. 7, XIII, p. 70.
.-\butschi, Ri\'er Niger.
Subfamily IDIURINAE
Geographr'AL Distribution. — Tropical Africa: Cameroons and Congo,
east to Lake Kivu.
Number of Genera. — Two.
Characters. — Size smaller than in Anomalurinae; intraorbital foramen
greatly enlarged owing to anterior prolongation of the zygo-
matic plate, which projects far forwards to a level immediately behind the
incisors, and is extremely narrow. Incisors greatly thickened from before back-
wards, and with prominent subapical notch. Cheekteeth extremely reduced
and brachyodont. Palate greatly narrowed. Bullae less inflated. Flving-
membrane present or absent.
Remarks. — This subfamily was regarded as a distinct family by .Miller &
Gidley, and further divided into two subfamilies the Idiurinae
and the Zenkerellinae mainly on account of the presence or absence of the
flying-membrane.
This division hides the close relationship obviously existing between
Zenkcrella and Id/urns; further, the presence or absence of a flying-membrane
certainly does not seem indicative of subfamily distinctions if one believes what
one has read about the relationships occurring in the family Phalangeridae
IDIURINAE: IDIURUS 543
(Marsupialia). In this group, apparently, a flying-membrane has been de-
veloped, independently in three cases; and each of these three genera appear
more closely allied to a non-flying member than to each other; indeed the large
Flying-phalanger, Petauroiiles, is sometimes referred to a distinct subfamily, the
Pseudochirinae, containing also the non-flying Pseudochirus; while the smaller
flying genera Acrobates and Petaurus are usually held to be related not to each
other, but more to Dislaechurus and Gymnobclideus respectively, neither of which
has a flying-membrane.
If cranial characters or resemblances are to be trusted, Idiurus is certainly
so closely allied to Zenkerella that there is no need for referring them to two
distinct subfamilies.
Key to the Genera of Idiurinae
Cheekteeth with two complete transverse ridges; infraorbital foramen
less widely open ; flying-membrane present. Idicrus
Cheekteeth with one complete transverse ridge; infraorbital foramen
more widely open; flying-membrane absent. Zenkerella
Genus i. IDIURUS, Matschie
1894. Idiurus, Matschie, Sitz. Ber. Ges. Nat. Fr. Berlin, p. 194.
Type Species. — Idiurus zenkeri, Matschie.
Range. — As in the subfamily.
Nu.mber of Forms. — Five.
Characters. — Anterior portion of skull rendered abnormal by the great
anterior prolongation of the zygomatic plate, which slants
downwards and forwards from the ascending root of the zygomatic process of
the maxillarv- to a level just behind the incisors. Infraorbital foramen with no
canal for ner\-e transmission. The bones forming the upper margin of the infra-
orbital foramen broadened, so that the foramen is less open than in Zenkerella.
Nasals short, narrow, well open anteriorly. Frontals with moderately developed
supraorbital ridges. No sign of postorbital process. Braincase smooth and
rounded. Zygoma not essentially different from Anomalurus. Bullae medium,
smaller than in Anomalurus relatively. Palate very narrow', extending back to
M.3, continued forwards as a straight shelf far in front of level of premolars.
Incisive foramina verv small, situated far forwards, between zygomatic plates.
Incisors greatly thickened from before backwards, much compressed.
Toothrow extremely reduced; cheekteeth very small; M.3 considerably
reduced in all skulls seen. Upper teeth cut into three subequal depressions by
two narrow transverse ridges; lower teeth similar in pattern to the upper series,
but apparently the outer fold present in Anomalurus can be present, and M.3 is
not specially reduced. Mandible high in proportion to its length.
Size very small for the family, head and body not exceeding 116 mm. in
those examined. Fur soft. Ear large, "its form strongly suggestive of that of
35 — Living Rodents — I
544
IDIURUS
some of the smaller Rats" (Miller: Idiurus macrotis). Tail considerably longer
than head and body, the upper part moderately or well haired, and long hairs
present throughout its length, the tail ending in a moderate brush. Scales on
underside moderatelv developed, less so than in ZcnkcrcUa and Anomaliirus
evidently. Flying-membrane present, its formation apparently similar to that
of Anonitiliinis, but the interfemoral membrane appears less developed than
Fig. 135. Idii Rus m.^crotis. Miller.
B..M. No. 3.2.4.16, i: ■ 3*.
in that genus. Functional digits of forefoot four, the pollex not traceable in
dried skins; claws prominent, curved. Hindfoot with five digits, the hallux
shorter than the others; claws as in forefoot.
Forms examined : senkcri, macrotis.
Two well-marked species are represented at the British Museum, the type,
much smaller, head and body about Si; -90 or less, and macrotis, Miller, with
head and body measurement of 105-116 or more.
But Allen, 1922, described two new forms, of the macrotis group, one of
which has a head and body measurement of only 73, or is about the size of
senkeri.
Fig. 136. Idiurus macrotis, Miller.
B.M. No. 3.2.4.16, 5; X 3J.
Fig. 137. Idiirus .m.acrotis, .Miller.
Cheekteeth: B,.M. No. 3.2.4.16, i; .-. ,-.
546 IDIURUS— ZENKERELLA
The difference between the two groups, if Allen's species are to he regarded
as species and not races, therefore lies in the measurement of the car and the
tail. The tail in Allen's measurements is at the highest io8 in zenkeri (none
of our specimens exceed 93); while Allen's lowest tail measurement for the
macrotis group is 117. The ear averages 13 -6 in a series of male zenkeri, and
13-4 in a series of females (the highest measurement being 14), whereas in the
macrotis group the lowest measurement is 14, the average being in langi 15-7,
in pcinga 17-3 and in macrotis (two specimens quoted), 16 or 15-5.
The total length measurements quoted for the various species are in Allen,
1S7 highest, zenkeri (160-187); -^'^ (207-224) for langi; 20b average (199-212)
pangii; and 228-241 for macrotis.
Full details will be found in Allen's paper.
List of Named Forms
zenkeri Group
1. IDRRLS ZENK1:RI ZENKERI, Matschie
1894. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 197.
Yaunde, S. Cameroons.
2. IDIURUS ZENKERI KIVUENSIS. I.onnbert;
1917. Stockholm Vet. Akad. Handl. 58, no. 2, p. 67.
Masisi, about 40 miles north-west of Lake Kivu, Belgian Congo.
macrotis Group
3. IDIURUS MACROTIS, Miller
1898. Proc. Biol. .Sec. Washington, XII, p. 73, figs. 15-19.
Efulen, Cameroons.
4. IDIURUS LANGI, Allen
1922. Bull. .\mer. Mus. Nat. Hist., XLVII, p. 69.
Medje, Ituri.
5. IDIURUS PAXC;A, Allen
1922. Bull. Amcr. Mus. Nat. Hist., XLVII, p. 70.
Panga, Ituri.
Genus 2. ZEXKHRELLA, Matschie
1898. Zenkerell.\, Matschie, Sitz. Ber. Ges. Nat. Fr. Berlin, no. 4, p. 23, no. 5, p. 63.
1 898. .AETHrRus, de Winton, Proc. Zool. Soc. London, p. 450, pis. XXXIV. XXXV.
(Aethunts gliriuus, de Winton - Zenkerella iusignis, Matschie.)
Type Species. — Zenkerella insignis, Matschie.
Range. — West Africa: Cameroons.
Number of Forms. — One.
Char.\cters. — (jeneral cranial characters much as Iclinriis; frontals appear
relativelv narrower, straight and well ridged, with no post-
orbital process. Infraorbital foramen more widely open. Mandible high in
i
ZENKERELLA— PEDETIDAE 547
proportion to its length, the ascending portion from the incisors to the coronoid
being straight; angular process relatively small. Jugal and lachrymal in contact
in the one skull seen.
P. 4 and 1M.3 both considerai>ly reduced in size. Cheekteeth simpler than
Idiurtis; one transverse ridge divides the tooth into two lobes, most of which are
occupied by a deep depression; lower molars like the upper series, P. 4 much
reduced.
Size larger than IiHiirus; tail long, naked for about a fifth of its length near
body, then thickly haired and bushy for the remainder of its length. Scales on
underside very coarse and large. Digits of forefoot four; hindfoot broad, with
five digits; general digit arrangement as in other genera; a tuft of brush-like
hairs present on ankles. Ears large. Flying-membrane absent.
The genus is evidently very rare.
Forms seen: insigfiis.
List of Named Forms
I. ZENKERELL.-Si INSIGNIS, Matschie
1898. Sitz. Ber. Ges. Nat. Fr. Berlin, no. 4, p. 24.
Yaunde, Cameroons.
Synonym: glirinus, de Winton, i8g8, Proc. Zool. Soc. London, p. 450.
Benito River, French Congo.
Nothing appears to be known of the fossil history of the family.
ANOMALURIDAE :
SPECIAL WORKS OF REFERENCE
TuLLBERG, Nova Acta Reg. Soc. Sci. Upsaliensis, XVIII, ser. 3, no. i, 1899. (Anoma-
lurus.)
RCmmler, Sitz. Ber. Ges. Nat. Fr. Berlin, p. 389, 1933. Revision of Anomalurus and
Anomalurops,
Miller, Proc. Biol. Soc. Washington, XII, 1898, p. 73. Description of a new Rodent
of the genus Idiurus.
De Winton, Proc. Zool. Soc. London, 1898, p. 450. (Description of Aethurns =
Zenkerella.)
Alston, On Anomalurus, its structure and position : Proc. Zool. Soc. London, 1875, p. 88.
Superfamily PEDETOIDAE
As here understood this contains one living family.
Family PEDETIDAE
1896. Thomas : Hystricomorpha. part. Family Pedetidae.
1899. Tullberg: Sciurognathi: Myomorpha : Anomaluroidei, part: Family Pedetidae.
1918. Miller & Gidley: Superfamily Dipodoid.^e, part. Family Pedetidae.
1924. Winge: Family .Anom^luridae, part : Pcdetini.
1928. Weber: Anomau'roidea, part: Family Pedetidae.
Geogr.\phical Distribution. — Central and Southern Africa: from Kenya
and Angola to Cape Province.
548 I'EDETIDAE
Number of Genera. — One.
Ch.^r.'\cters. — Zygomasscteric structure essentially as in Anomaluroidae,
so far as it concerns the shape of the mandible, infraorbital
foramen (which is tjreatly enlarged), and zygomatic plate. According to
Tullberg's figures, the temporalis muscles of Pcdctes are much more reduced
than in Anomalurus; and the pterygoid muscles and pits for their insertion are
much more extensive in Pedetcs (being apparently very weak in all Anomalur-
idae). Skull massive and Hystricoid in general appearance, but mandible with
angular portion small, not distorted outwards; zygomatic region much thick-
ened ; mastoids extremely inflated ; cheekteeth evergrowing, simplified in pattern ;
dental formula i. ] , c. 2, p. 1, m. ji = 20, the premolars not reduced in size. Fibula
reduced and fullv fused with the tibia in adult. External form modified for
bipedal saltatorial life; digits of hindfoot reduced to four, the metatarsals
normal, not becoming fused (compare specialized Dipodidae).
Remarks. — Great diversity of opinion has prevailed on the systematic
position of this tamily. Most authorities are now agreed that
the relationship with the Dipodidae, in which it was formerly classed, is
remote. Winge, TuUberg, and Weber place the family in the Anomaluroid
division. But there seems to be remarkably little in common between Pedetes
and the Anomaluridae as known to-day other than that both are clearly somewhat
archaic groups of Rodents. Although the zygomasscteric structure is essentially
the same in the two groups, the following characters appear to me to indicate
a rather wide gap between the two families, and show that the Pedetidae are
very much more specialized in many ways than the Anomaluridae :
Cheekteeth evergrowing in Pedetes; rooted and brachyodont in the Anoma-
luridae.
Pattern of cheekteeth simplified in Pedetes; very rarely showing any sign of
simplification in the Anomaluridae, usually of the rather primitive
complex type found in some Sciuridae, Erethizontinae. (Only in
Zeiikerellii, in which the whole toothrow is extremely reduced, is there
sign of simphfii'ation in the one skull seen.)
Fihiilii fused with the tibia in Pedetes, as far as known separate or not tuliy
fused in Anomaluridae.
Dit;its reduced in the pes to four in Pedetes; not so in Anomaluridae.
Skull specialized, characterized by large inflated mastoids, thickened zygoma,
massive frontals, heavy rostrum, deepened pterygoid fossae in Pedetes;
not so in Anomaluridae.
Some of the above may be adaptive characters. But taken altogether they
seem to indicate a wide difference to-day, whatever the ancestors of the two
groups may have had in common.
The two families cannot in my opinion be regarded as so closely allied
to each other as, say, any two families of Hystricoidae, or any two families of
Muroidae.
On the other hand 'I'ullberp lists a number of points which are shared by
PEDETIDAE: PEDETES 549
Pedetes and the Anomaluridac, some of which arc in the ahmentary canal;
the lachrymal foramen is placed hit^h up; the great similarity in the hyoid bone
of the two genera (he evidently did not examine Idiurus and Zetikcrella); the
number of cheekteeth (but this also applies, for instance, to Muscardinidae !);
the large infraorbital foramen (as Dipodidae, Ctenodactylidae, etc.); and the
absence of a transverse canal in the corpus of the sphenoideum.
Thomas transferred the family to the Hystricomorpha, remarking: "while
many naturalists have noticed the Hystricomorph affinities of Pedetes, no one in
modern times (except Dobson, who transferred the whole of the Dipodidae)
seems to have thought of actually placing it among them. To me this appears
to be clearly the proper course as there seems to be scarcely a character in its
skull or teeth which is not found in one member or another of that group."
But the "Hystricomorpha" of Alston, on which Thomas's classification is based,
are defined as with the tibia and fibula persisting as free bones, whereas in
Pedetes they fuse. Taking into account that this structure is not a sufficientlv
important one on which to base major groups, the fact remains that though there
may be a suggestion of the "twisted" lower jaw characteristic of the Hystricoidae
in Pedetes, it is nothing like fully developed; indeed, the angular process is in
this genus reduced; and in Tullberg's figures of the zygomasseteric structure of
Pedetes, on the lower jaw it will be seen that the portion marked mis. (masseter
lateralis superficialis), which in every member figured of his Hystricognathi
( = Bathyergidae +Hystricoidea) except the Caviidae, takes up the greater
portion of the jaw, is in Pedetes, as in Ctenodactylus, Anomalunis and others,
small and unimportant.
There appears to be to me no alternative to the classification of Pedetes as
a superfamily distinct from all others in living Rodentia.
Genus i. PEDETES, Illiger
181 1. Pedetes, Illiger, Prodr. Syst. Mamm. & Avium, p. 81.
Type Species. — Mus cafer, Pallas.
Range. — As in the family Pedetidae.
Number of Forms. — Ten or eleven.
Characters. — Skull with broad nasals, extremely broad frontals; mastoid
portion of bullae appearing conspicuously in superior aspect
of skull; lower zygomatic root projected forwards to a point immediately behind
the incisors, the zygomatic plate continued forwards and forming nearly a right
angle with the ascending portion of the maxillary process of zygoma, as a conse-
quence of which the infraorbital foramen is greatly enlarged. No canal for
nerve transmission in the infraorbital foramen. Jugal immensely broadened
anteriorly, and in contact with the lachrymal. Bony palate short, extending
backwards only to M.i or the front portion of M.2. Basisphenoid much
narrowed; pterygoid fossae deep. A pitlike depression is situated in front of
the toothrows, which extends forward to incisors and lies between the zygomatic
plates.
Fig. 138. Pedetes siirdaster larvalis, Hollister.
B.M. No. 28. 1 2.7. 14; I.
Fig. 139. Pedetes surdaster larvalis, Hollister.
B.M No. 28. 12. 7. 14; / I.
PEDETES
551
Mandible with angular portion short and somewhat reduced; coronoid
process much reduced; a strong ridge for muscle attachment on lower border of
angular portion, which is, however, not "lifted outwards" as it is in Hystricoidae
and Bathyergidae.
Incisors thick. Cheekteeth evergrowing, each tooth divided into two lobes
by a re-entrant angle, in the upper series externally, in the lower series internally.
The teeth when cut are not entirely simplified, and traces of more than one
inner cusp may be seen in the inner side of the centre of the upper molars;
but the lower molars of the one very young specimen examined are practically
simple.
Fig.
-r_
140. Pedetes surdaster larvaljs, Hollister.
Cheekteeth: B.M. No. 28. 12.7. 14; x 4.
Externally like a giant Jerboa. Hindfoot perissodactyle in arrangement of
digits, elongated; claws more or less hooflike; hallux absent; D.5 short but well
developed. Tail about as long as head and body or frequently longer than this
measurement, heavily haired, with a thick black brush terminally. Ears
prominent. Alanus with five digits, the claws strong, that of the poUex appar-
ently in no way reduced and as large as the others (a rare feature in the Order).
Pocock states, with reference to the fact that this genus is not to be associated
in the Ilvstricoidae, that "the penis is elongated . . . but there is no trace of the
glandular pouch which is so characteristic of the Hj'stricomorphs."
The family is, as far as I have traced, not known fossil outside Africa, though
a related fossil genus, Parapedetes, has been described from that continent.
The species admitted are unquestionably very closely related to each other.
Forms examined: an^olae. cafer, orangiae, salinae, surdaster.
552 PEDETES
SPECIAL WORKS OF REFERENCE
Ti'LLBERC, Nova Acta Rep. Soc. Sci. Upsaliensis, XVIII, 3, no. i, 1899.
HoLHSTER, East African Mammals in the United States National Museum: Smiths.
Inst. Bull. 99, p. 156, 1919.
PococK, External Characters of Scarlurus and other Jerboas compared with Ztipiis and
Pedetes: Proc. Zool. Soc. London, p. 659, 1922.
List of Named Forms
(The references and type localities have been collected by Mr. R. W. Hayman.)
1. PEDETKS CAFKR CAFER, Pallas
1778. Nov. Spec. Quadr. Glir. Ord., p. 87.
Cape Colony.
Synonym: capensis, Forster, Svensk. Vet. Acad., p. loS, pi. Ill, 1778.
typicus. Smith, 111. S. Afr. Zool., 1849, p. 20,
2. PEDETES CAFER OR.'^NGIAE, WrouKhton
1907. Ann. Mag. Nat. Hist. 7, XX, p. 32.
.Aberfeldy district, Orange River Colony.
3. PEDETES CAFER SAMNAE, Wrouchton
1907. Ann. Mag. Nat. Hist. 7, XX, p. 33.
Woodbush, Zoutspansberg district, N.-E. Transvaal.
4. PEDETES CAFER DAMARENSIS, Roberts
1926. Ann. Transvaal Mus. XI, p. 261.
Okahandja, S.-W. Africa.
5. PEDETES CAFER TABORAE, Allen & I.overidge
1927. Proc. Boston Nat. Hist. Soc. 38, p. 438.
Tabora, Tanganyika.
6. PEDETES CAFER DENTATUS, Miller
1927. Proc. Biol. Soc. Washington, XL, p. 113.
D(jdoma, Tanganyika.
7. PEDETES CAFER ANGOLAE, Hinton
1920. Ann. Mag. Nat. Hist. 9, VI, p. 102.
About 20 miles north-east of Bihe, Angola.
8. PEDETES SURDASTER SURDASTER, Thomas
1902. Ann. Mag. Nat. Hist. 7, IX, p. 440.
Morendat, mile 365 of Uganda Railway, Naivasha. Kenya.
0. PEDETES SLRDASTER CURR.\X, HoUister
191S. Smiths. Misc. Coll. LXVIII, no. 10, p. 3.
Kabalalot Hill, Sotik, Kenya.
10. PEDETES SCRDAST1:R LARVALIS, Hollister
1918. Smiths. Misc. Coll. LXVIII, no. 10, p. 2.
Ulukenia Hills, Athi Plains, Kenya.
There are specimens at the British Museum labelled " Peih'tes cafer brad-
fieldi," from the Kalahari. The reference to this form has not been traced.
CTENODACTYLIDAE 553
Superfamily CTEXODACTYLOIDAE
As here understood this contains one hving family.
Family CTENODACTYLIDAE
1896. Thomas: Hystricomorpha : Family Octodontidae, part, Subfamily Ctenodact>'l-
inae, part, included Petromys.
1899. Tullberg: Scilrognathi : Myomorpha: Ctenodactyloidei : Family CtenodamM-
idae.
1918. Miller & Gidley: Superfamily Dipodoidae, part. Family Ctenodact\lidae.
1924. Winge: Hystricidae, part, Ctenodactylini, part, included Petromys.
192K. Weber: Hystricoidea : part, Family Ctenodact\-lidae.
Geographical Distribution. — Northern Africa: from Senegal and Morocco
(including Palaearctic coastal area) to
Somaliland.
Number of GENERA.^Four.
Characters. — Zygomasseteric structure, so far as it affects the shape of the
skull, about as in Dipodidae, though the mandible is slightly
more modified than in that family. Dental formula i. {, c. i^, p. { or a, m. | = 20
or 24, at full dentition. The extra upper and lower premolars in the genus
Pectitiator are shed before the posterior molars are cut ; in the remaining genera,
with cheekteeth at full dentition normally *, the premolar is shed early during
life, and is rarely present in the specimens examined. Cheekteeth evergrowing,
practically or completely simplified in pattern. Auditory bullae and mastoids
considerably inflated, the mastoids normally a prominent feature in the superior
aspect of the skull. Jugal divided into two portions, a horizontal and a vertical,
as in the subfamily Dipodinae. Lachrymal large. Mandible with no coronoid
process, and usually a weak ridge present for (presumably) attachment of
masseter medialis, this structure foreshadowing that present in Caviidae but
very much less developed than in that family. Angular portion of mandible
produced backwards to a greater or lesser degree, not excessively so. Digits of
both fore and hindfoot reduced to four. Tail fully haired, much shortened.
Tibia and fibula not fused.
Remarks. — The Ctenodactylidae have been associated with the Hystricoid
Rodents by Thomas, Weber, and Winge. Miller & Gidley
place the family among the Dipodoidae, in the neighbourhood of Dipodidae
and Pedetidae; Tullberg regards them as more nearly related to Muroid (and
Dipodoid) Rodents than to the Hvstricoids. Pocock states : "The claims of
Ctenodactylus, indeed, to a place in the Ilystricomorphs seem to me to be more
than questionable."
Peters in an extensive paper on the genus Pectitiator came to the following
conclusions: "The Ctenodactyli cannot be associated with the Dipodes, their
relation to them being not greater than that of the Chinchillae, Octodontes, and
554 CTENODACTYLIDAE
Echinomyes; they show in nearly every part of their structure their near relation-
ship with the last-named groups, and deviate from them only in a very few
points, the form of the hyoid bone, of the sacral and caudal vertebral column,
of the development of the crest of the humerus and femur, in which they do not
show any inclination towards the Dipodes, but rather some affinity with the
Murinae; they form a peculiar group of the Hystricidae as understood by
Waterhouse, which in some points is more allied to the Chinchillae, in other
points to the Octodontes. Pelromys is not to be associated with the Cteno-
dactyli, but with the Octodontes."
(It may be noted that Waterhouse divided his Hystricidae into six groups,
the Hystricina, Echimyina, Octodontina, Dasyproctina, Chinchillina, and
Caviina.)
None of the authors who place the Ctenodactylidae in the Hystricomorph
series note, however, that the form of the mandible in Ctenodactylidae does not
agree with either that present in Hystricoidae, as here understood, or with that
of the Caviidae. It may be transitionary towards the latter, but it certainly,
so far as I have had occasion to examine, does not agree with it in structure.
This being the case, the zygomasseteric structure of this family cannot be
said to agree with that of the Hystricoidae. I do not think that the faint resem-
blances between these animals and the Caviidae in lower jaw structure need
indicate any close relationship.
Tullberg has dealt extensively with the relationships of this group, and
comes to the conclusion that on account of the formation of the mandible they
cannot he regarded as Hystricoid Rodents, a view which I fully support. He
is evidently of the opinion that they may not be distantly related to the Anomalu-
roid-Pedetoid branch of the Order. He writes extensively on the parallel
evolution of this group and the Chinchillidae.
The separation of Ctenodactvlidae from the Hystricoidae is supported by
Miller &: Ciidley, who rightly place it in a superfamily (Dipodoidae) in which the
angular portion of the mandible is not distorted outwards.
Further characters which should be mentioned are that according to Tullberg
the radiale and intermedium are separate, alone in Rodents (as examined by
him) excepting the Bathyergidae; and that the malleus and incus are fused, as
in Hystricoidae and Bathyergoidae, but unlike the remainder of the Rodents.
Four rather closely allied genera are now admitted.
Of these Pcctinator seems to be the most primitive, in that the full dentition
is J instead of |, the palate is relatively shorter, and the tail appears rather less
reduced than in allied forms. iMiissoiitlcm and Felovia have the "eight-shaped"
type of cheekteeth found in South American Octodontinae in the genera
Octonixs, Acunuemys and Spalacopiis; Ctenodactvlus parallels the Octodont
genera Ctenomys and Octodon in having "kidney-shaped" cheekteeth, and in
this genus the tail reaches its greatest reduction in the family; also Ctenodactyliis
appears to tend to be a little larger than other members of the family.
Skull CiiARACTr.RS. — The following skull characters appear constant in the
family ;
The skull is flattened, with broad frontals; it is wider posteriorly than
CTENODACTYLIDAE: PECTINATOR 555
anteriorly ; the rostrum is moderate or narrow and inclined to be bowed down-
wards. The supraorbital ridges are comparatively well developed; and a small
postorbital-like ridge on the parietal is situated immediately in front of and
above the squamosal roots ot the zygoma, similar to that present in Jaculus.
The apices of the bullae are not in contact. Jugal extending up to the lachrvmal.
General scheme of zygoma like that of Jaculus, but horizontal portion about
as broad as the vertical portion, and the vertical part does not rise to such a
high degree, due perhaps to the much lower skull. Incisive foramina large and
well open, extending about to the toothrows, broader posteriorly. Paroccipital
process closely applied to bullae, and quite large. A prominent canal on
maxillae running through the infraorbital foramen, the latter much enlarged.
Except in the genus Pectinator the toothrows tend to converge in front, and the
palate projects behind M.3. In all genera the upper incisors are opisthodont.
Key to the Genera of Ctenodactylidae
Cheekteeth at full dentition j; lower cheekteeth less simplified, with two
well-marked inner folds. Palate not extending behind toothrows.
Pectinator
Cheekteeth at full dentition *. Lower cheekteeth simpler, with one inner
fold only. Palate normally extending behind toothrows.
Upper cheekteeth simpler, with no inner fold, the general effect
"kidney-shaped." Ctenod.ictylus
Upper cheekteeth less simple, with inner fold present, general effect
becoming "eight-shaped."
Inflation of bullae and mastoids at maximum for the family;
outer fold of upper cheekteeth very narrow. Massoltier.'v
Inflation of bullae and mastoids at minimum for the family;
outer fold of upper cheekteeth remaining widely open. Felovia
Genus I. PECTINATOR, Blyth
1855. Pectin.\tor, Blyth, Journ. Asiat. Soc. Bengal, XXIV, p. 294.
Type Species. — Pectinator spekei, Blyth.
Range. — Africa : Abyssinia, Somaliland, Eritrea.
Number of Forms.— Three.
Characters. — Palate shorter than Ctenodactylus, not extending behind last
molars. Bullae and mastoids considcrablv inflated. Cheek-
teeth ; at full dentition (it may be mentioned that this genus and the Bathver-
goiti Ueliop/iobius are the onlv living genera of Rodents with a cheekteeth
formula of more than J). P. 5 minute; M. ij cut late in life; the small extra
premolars shed before the posterior molars are cut as a rule. The upper cheek-
teeth are tolerablv similar to those of Ctenodactvlus, but in addition to the shallow
556 PECTINATOR— CTENODACTYLUS
outer open fold there is a verv small inner fold. The lower cheekteeth are more
complicated than in other members of the family; there are two sharp well-
marked inner folds, and one outer one.
Form more or less Guineapig-like. Fur soft. Digits of forefoot four, sub-
equal, the claws not large. Digits of hindfoot four, the two inner digits with
well-developed brush of comb-like bristles, the outer digits with similar structure
rather less developed. Ilindfoot relatively narrow; tail bushy, a little longer
than hindfoot.
Pocock, writing of this genus and Ctenodactvhis, states : "Assuming that their
ears are similar, these two genera differ markedlv not only from all the Hystri-
comorphs but from all other Rodents known to me in the structure of this
organ."
Forms examined : spckci.
List of \.\med Forms
1. PECTINATOR SPP;KEI SPEKEI, Blvth
1S55. Joum. .Asiat. Soc. Bengal, XXIV, p. 294, pi. II, fig. i.
Somaliland, 11" 40' N.
2. PECTIN.ATOR SPEKEI MERIDIONALIS, du Beaux
1922. Atti. Soc. Ital. Sci. Nat. 61, p. 27.
Dolo, Somaliland.
3. PECTINATOR SPI-KEI LEGERAE, de Beaux
1934. Atti. Snc. Ital. Sci. Nat. 73, p. 293.
Assab, Eritrea.
Genus 2. CTENODACTYLUS, Gray
1828. CTENODACTYLUS, Gray, Spicil. Zool., p. 10.
Type Species. — Ctenodactylus massonii, Gray.
Range. — Northern Africa : Morocco, Algeria and Tripoli.
Number of Forms. — Four.
Characters. — Cheekteeth \ at full dentition. In nearly all specimens
examined there are only %\ but as pointed out by Lataste,
the premolar is present, though shed early. This author takes from a series of
specimens seven skulls, each with different teeth in place, and describes first
the newly-born, with two teeth, P. 4 and M.i, the teeth described as tubercu-
lated; next a young skull, with the four teeth in place, the posterior one being
cut; the next stage with only three, the front premolar having been shed; then
with the same teeth present, but P.4 being cut; then with the four teeth in place;
then an older animal with three teeth but with a scar marking where the pre-
molar had been shed; finally the adult, with three cheekteeth only, and all
trace of P.4 lost.
Cheekteeth simple, reminiscent to a certain degree of those of Ctenomys;
with no inner fold in the upper series, but with a shallow widely open outer
Fig. 141. Ctenodactylus gundi cundi, Rothman.
B.M. No. 97.6.9.19, 5; X ij.
^^_;mx^
\.
Fig. 142. Cte.sod.^ctvxus gundi gundi, Rothman.
B.M. No. 97.6.9.19, J; X ij.
5S8
CTENODACTYLUS
fold in all teeth. Lower cheekteeth with one widely open outer and inner
re-entrant fold in all teeth.
Palate longer than in Peciinator. Bullae and mastoids considerably inflated,
most so in C. vali.
Externally like Pcctinator except that the tail is considerably shorter than
the hindfoot, and almost obsolete.
These animals are described as being born hairy, and not blind, and able
to run at birth.
Fig. 143. Cte.xodactyll's gindi glndi, Rothman.
Mandible from below, ij; Cheekteeth ■ 7; B.M. Xo. 97.6.9.19, ?.
Two species are at present admitted, which as indicated above differ in the
size of the mastoids and bullae; whether such ditierences would be valid in
the event of a really large number ot specimens coming to hand is at the moment
not known.
Forms examined : giindi, massonii, vali.
List of Named Forms
1. CTKNOD.ACrVLUS GUNDI GUNDI, Rothman
1776. Schloezer's Briefwechsel, p. 339.
Gharian, 80 km. south of Tripoli.
Synonym: nrahicus, Shaw, Gen. Zool., II, 1801, p. 123.
2. CTENOD.ACTYLUS GUNDI MASSONII, Gray
1S28. Spicil. Zool., p. 10, pi. 10.
Biskra, south slope of Atlas Mountains.
CTENODACTYLUS— MASSOUTIERA 559
J. CTENODACTYLUS VALI, Thomas
1902. Proc. Zool. Soc. London, p. 11.
Wadi Bey, north-west of Bonjem, Tripoli.
4. CTIiNODACTYLL'S JOLIiAUDI, Huim de Balsac
1936. Suppl. Bull. Biol, de France et de Belgique, Paris, 21, p. 315.
Beni Ounif, Jebel Melias, Algeria.
Genus 3. MASSOUTIER.\, Lataste
1885. Massoutiera, Lataste, Le Naturaliste, no. 3, p. 21.
Type Species. — Massoutiera msahi, Lataste.
Range. — Africa : Central and Western Sahara.
Number of Forms. — Three.
Characters. — Like Ctenodactylus, but mastoids typically more inflated, and
cheekteeth less simplified ; those of upper series divided into
two lohes by narrow folds, the folds meeting in the middle of the teeth, the
structure in general reminiscent of that present in the South American
Aconaemys. iSL3 the largest tooth. Lower cheekteeth with the same elements,
but rather more angular, the folds more open. Tail less reduced than in
Ctenodactylus, about as long as hindfoot; bushv; other essential external
characters as in Ctenodactylus.
Three species are admitted in this genus at present : harterti, with mastoids
and bullae at maximum inflation for the family; rothschildi, in which the bullae
are more moderate; and mzabi, in which they are smallest for the genus, but
still relatively very large. Each appears to be known, so far as the London
collection is representative, by a comparatively small number of specimens.
Forms examined : harterti, rothschildi, mzabi.
List of Named Forms
1. MASSOUTIERA MZABL Latastc
1881. Bull. Soc. Zool. France, VI, p. 314.
Ghardaia, .\lgerian Sahara.
2. MASSOUTIERA ROTHSCHILDI, Thomas & Hinton
1921. Nov. Zool., XXVIII, p. II.
Mount Baguezan, -Asben, Sahara.
3. MASSOUTIERA HARTERTI, Thomas
1913. Nov. Zool., XX, p. 31.
Oued Mya, south of Fort Mirabel, Western Sahara, about 28' 30' N. 3' E.
Genus 4. FELOVIA, Lataste
1886. Felovia, Lataste, Le Naturaliste, iii, p. 287.
Type Species. — Felovia vae, Lataste.
Range. — Known from Senegal (\.-W. Africa).
Number of Forms. — One.
36 — Living Rodents — I
56o FELOVIA— DH'ODIDAE
Characters. — Like Massoutiera, but bullae and mastoids considerably less
inflated, appearing on the top of the skull to a lesser degree
than in any other member of the family.
Cheekteeth differing from MassouUera in that the outer fold of the upper
series remains widely open. In the lower series M.3 has a short backwardly
pointing heel. The upper incisors are faintly grooved. Externally like Massoti-
tiera.
Remarks. — Though closely allied to Massoutiera, the differences indicated
above seem sufficient to warrant the retention of this genus.
Forms examined : vae.
List of Named Forms
I. FELO\'lA VAF., Lataste
1 886. Le Naturaliste, iii, p. 287.
Felou. Medina district, Upper Senegal River, Senegal.
The references and type localities to all named forms of this family have
been collected for me by Mr. R. W. Hayman.
The Family Ctenodactylidae is known from the Pliocene, from the Mediter-
ranean region. Pectinator has been described fossil from the Miocene of India.
CTENODACTYLIDAE :
SPECIAL WORKS OF REFERENCE
TuLLBERG. Nova Acta Regiale Soc. Sci. Upsaliensis, XVIII, ser. 3, no. i. 1899. (Cteno-
dactylus).
Peters, Contribution to the knowledge of Pectinator, a genus of Rodent Mammalia
from North-eastern Africa: Trans. Zool. Soc. London, VII, p. 397, 1871.
Lataste, Le NaturaUste, 1885, p. i. Sur le systeme dentaire du genre Ctenodactyhis,
Gray.
St. Leger, Key to Families and Genera of African Rodentia: Proc. Zool. Soc. London,
1931, p. 978.
Superfamily DIPODOIDAE
This contains, as here understood, one family, the Dipodidae, with sub-
families Sicistinae, Zapodinae, Cardiocraniinae, Euchoreutinae, and Dipodinae,
the relationships and characters of which are fully discussed below-
Family DIPODIDAE
1896. Thomas: Myomorpha, part: F'amily Dipodidae. Subfamilies Sminthinae
(=Sicistinae), Zapodinae, and Dipodinae.
1899. Tullberg: Mvomorpha; Dipodiformes, Family Dipodidae.
1918. Miller & Gidlcy: Superfamily Dipodoid.\i;: F"amily Zapodidae (Subfamilies
Sicistinae, Theridomyinae (fossil), and Zapodinae); Family Dipodidae (Subfamilies
Protoptychinae (fossil), and Dipodinae).
1924. Winge: F'amily Dipodidae, part (includes Spalax): Dipodini.
1928. Weber: DlPODorDE.\: Family Sicistidae (Sicista only), and F'amily Dipodidae.
DIPODIDAE S6i
Geographical Distribution. — Holarctic region: Europe from Norway,
Denmark, Hungary and Bulgaria, east-
wards across Asia south to Kashmir, Szechuan, Afghanistan, Arabia; east to
Sakhalin. Africa, northern, from Senegambia and Morocco to Egjpt and
Somaliland; North America from Alaska to the Atlantic coast, covering the
greater part of Canada and the United States but evidently not occurring in
Mexico.
Number of Genera. — Fifteen are here retained, based as far as possible
on Vinogradov's papers on the family. The only
modifications I make are that Scarturus is here considered a synonym of Allac-
taga ; and that Napaeozapiis and Eozapus are, following American authors, here
given full generic rank, chiefly on account of their dental peculiarities.
Characters. — Infraorbital foramen greatly enlarged for muscle trans-
mission; zvgomatic plate narrow and remaining completely
below it; mandible weak, the angular process not distorted outwards, this part
of the jaw frequently with a perforation present.
Dental formula i. \, c. g, p. i or jj, m. f = i6 or i8. When present, the extra
premolar usually very small. It should be noted that in a skull in the British
Museum of Allactaga elater there is a minute extra tooth at the back of the
series. This might suggest that apparently, as in Muridae, reduction has taken
place from behind so that the formula might be more correctly written as
p. J or \, m. s. But for convenience I adopt the notation given above.
Cheekteeth rooted, usuallv cuspidate, with broad re-entrant folds, the
pattern often reminiscent of that of Cricetinae; in Zapodinae tending to become
flatcrowned, in which case the re-entrant angles in progressive forms become
narrow.
Tibia and fibula fused high on the leg, the fibula reduced, thread-like.
Externally showing a progressive series of adaptations towards bipedal
saltatorial life, except in the genus Sicista ; at highest development more specialized
for saltatorial life than in any other Rodents, the three central metatarsal bones
fusing to form a cannonbone (Dipodinae (including Allactaga group), Eucho-
reutinae). Some of the cer\ical vertebrae tend to become fused in Dipodinae.
Zygoma in progressive forms divided into two portions, a horizontal and a
vertical, these portions forming a sharp angle with each other. Infraorbital
foramen always with a separate canal for nerve-transmission. Incisors com-
pressed, frequently grooved. Hindfoot with hallux reduced or absent; three
functional digits only present in all members of the family except Cardiocraidus
(not seen), Sicista, and the Zapodinae. In progressive genera, skull extremely
specialized, by modification of zygomatic region, as indicated above, broadening
of frontals and still more of braincasc, and in certain abnormal cases by extreme
inflation of mastoids and bullae.
RE.MARKS. — The Dipodidac have frequently been referred to (Forsyth
Major, Winge, etc.), as being the forerunners of the Muridae;
in the opinion of Forsjth Major evidently they are to be considered more
primitive than the Muridae.
S62 DIPODIDAE
All members of the present family with the exception of Sicista and prob-
ably Zapiis seem to me to be so very much more highly specialized than any
member of the Muridae or any member of the Superfamily Muroidae as
here understood, in so many ways, that I very much question if this assump-
tion is correct, and would regard them as at most parallel offshoots of a
common ancestor in which the present group has become much more highlv
developed.
Divisions. — In Vinogradov's earlier paper, on the genital organs of the
Dipodidae, he recognizes two families, Zapodidae and Dipo-
didae, and seven subiamilies. In his later paper, (jn cranial characters, he states
that the Zapodidae are best referred to the Dipodidae, and he reduces his sub-
families to five, by referring Sulpingotiis to the Cardiocraniinae, and Sicista to
the Zapodinae. His classification is in the main followed, though I prefer to
keep Sicista apart as type of a subfamily from the Zapodinae, and I think that
the AUactaginae and Dipodinae of Vinogradov, though sharply separated as
"groups," have too many characters in common for it to be necessary to refer
them to separate subfamilies.
Miller & Gidlev, and many American authors, referred the Zapodinae to
a separate family chiefly on account of the lack of fusion of the three central
metatarsals of the hindfoot. The discovery that the metatarsals are free in the
exceedingly rare Palaearctic Pygmv Jerboas, Salpiugotus and Cardiocranius is
of great interest, and seems to render this course no longer necessary, striking
as the diff^erences are between the specialized fused metatarsals of higher
members of the group and the more normal lower type found in Sicista,
Zapodinae, and Cardiocraniinae.
Salpiugotus and Cardiocranius appear to be true members of the Dipodidae
as currently understood, agreeing with the higher forms, as far as one reads, in
dental structure, foreshadowing them in the structure of their zygoma, and
even exceeding them in inflation of the bullae and mastoids.
F.uchoreutes, on the other hand, agreeing with AHactaga and Dipus in the
fusion of the metatarsals, presents seveial cranial resemblances to the Zapodinae,
as, for instance, the normal zygoma.
It would seem therefore that there are two alternatives, to recognize three
families, based on metatarsal structure alone, the Zapodidae, Cardiocraniidae,
and Dipodidae, the last including Euchoreutes, or to unite them altogether as
a family, as do most Russian authors of to-day. Personally I am in agree-
ment with the latter course. It must be noted, however, that the differences
between the most generalized member of the present group, Sicista,
and the most specialized member, say Jacuhis or AHactaga, are greater than
between those of the lowest and highest members ot the other families in the
Order.
According to Tullberg the malleus and incus of Sicista agree with or
resemble those of Cricetus and the Muridae, but those of Zapus differ not only
from Sicista but also from AHactaga and Jaculus.
DIPODIDAE 563
Kr.Y TO THE Subfamilies of Dipodidae
The three central metatarsal bones not fused to form cannonbone.
Auditory bullae not inflated, relatively small. Jugal slanting gradually
upwards to the lachrymal, zygoma simple.
Cheekteeth brachyodont, cuspidate, quadritubercular, with
moderately marked re-entrant folds, the teeth not showing
tendency to become flatcrowned. Hindfoot not lengthened;
externally not specialized for saltatorial life. Subfamily Sicistinae
(Sia'sta)
Cheekteeth semihypsodont, not quadritubercular; primitively
with strongly marked re-entrant folds; showing a tendency
to become flatcrowned, in which case the re-entrant folds
isolate on crown surface, or become narrowed. External
form considerably modified for saltatorial life, the hindfoot
lengthened. Subfamily Zapodinae
(Eozapus, Zapus, Napaeozapus)
Auditory bullae and mastoids abnormally inflated, about at maximum
development for the whole Order; occupying about a third of
upper surface of skull. Jugal in two portions, a horizontal and
a vertical, these portions connected by a curvature. (External
form saltatorial.) Subfamily C.\rdiocraniinae
{Cardiocranius (not seen), Salpingotus)
The three central metatarsal bones fused to form a cannonbone.
Jugal slanting gradually upwards towards lachrymal, the zygoma
simple; cheekteeth with extremely high cusps and shallow
re-entrant folds; !\1.3 vestigial (so far as ascertainable); skull
narrowed at a point considerably behind lachrymals; rostrum
elongated. (Ear abnormally enlarged; mastoids well inflated.)
Subfamily Euchoreutinae
(Euchoreutes)
Jugal in two portions, a horizontal and a vertical, these portions form-
ing a sharp angle with each other, and not connected by a
curvature. Cheekteeth with moderate or low cusps, and well-
marked re-entrant folds. .M.3 not vestigial. Skull not narrowed,
or very slightly so immediately behind lachrymals; rostrum
not elongated. (Ear less enlarged; mastoids well to extremely
inflated.) Subfamily Dipodinae
Mastoids and bullae little inflated (comparatively); upper incisors
pro-odont; ears usuallv larger; functionless lateral digits of
hindfoot not suppressed; infraorbital foramen more widely
open, and anterior vertical portion of jugal not greatly broad-
ened; OS penis absent (Vinogradov). ,^//ac?a^a Group (Allactagae)
{Allactaaa, Alactagiilus, Pygeretmus)
564 SICISTIXAE: SICISTA
Mastoids and bullae considerably to extremely inflated; upper
incisors not pro-odont; ears usually smaller; lateral digits of
hindtoot suppressed; intraorbital foramen less widely open,
and anterior vertical portion of zvgoma greatly broadened;
OS penis present (Vinogradov). Dipus Group (Dipodes)
(Paradipus. Dipus, Scirtopotia, Jaciilns, Ereinodipus (not seen))
Subfamily SICISTINAE
Geographical Distribution. — Palaearctic: Central Norway, Denmark,
Finland, Eastern Roumania, Bulgaria,
Hungarv; European Russia (Ukraine, Crimea, the Caucasus; surroundings of
Leningrad, Moscow, former Tver govt., lower part of River Pechora, former
.\rchangel district, North I'ral); Asiatic Russia; Kazak.stan east to former
Kusnetzk, Krasnoiar, Minusinsk and Irkutsk districts, Altai; East coast Lake
Baikal; Tianshan mountains; Sakhalin, and L'ssuri region (all Russian localities
as quoted bv Vinogradov, 1933). Also known from Chinese Turkestan, Man-
churia, Szechuan, Kansu, Kashmir.
Number of Genera. — One.
Characters. — As indicated in the above key. Skull little specialized;
braincase rounded; bullae relativelv small; upper and lower
zygomatic roots above one another; jugal slanting gradually towards lachrvmal,
but not in contact with it; rostrum moderately long. External form Murine,
limbs not lengthened, not modified for saltatorial life. Size very small. Cheek-
teeth |.
As compared with Graphiurus and Deom\s, the only two genera included
here in the Muroidae which present a similar arrangement of the zvgomatic
plate and infraorbital foramen, it appears that the infraorbital foramen of the
present genus is relativelv more widely open, and shaped difFerentlv, being
considerably broader below than in either of the two genera mentioned above,
and more or less triangular in shape. The intraorbital foramen oi Zapiis agrees
essentially with that of Sicista.
Genus i. SICISTA, Gray
1827. Sicista, Gray, Griffith's .Anim. Kingd., V, p. 227.
1840. Sminthis, Xathusius, A.V. Xnrdmann in DemidofF Voy. Russie. iii, p. 49.
Smintlms longer, Xathusius.
Type Species. — Mus suhtilis, Pallas.
Range. — As in the subfamily Sicistinae.
Number of Forms. — About seventeen.
Characters. — Frontals constricted at considerable distance behind
lachrymals, degree of constriction moderate. Palate pro-
jecting beyond M.3, terminating in a spinous process, the palate broad.
Nasals not projecting beyond premaxillae. Skull without supraorbital ridges.
Incisive foramina large, well open, extending about to toothrovv.
T^
Fig. 144. SicisTA slbtilis lobiger, Nathusius.
B.M. No. 12. 12. 17.13, ?; X 4.
Fig. 145. SicrsTA subtilis loriger, Nathusius.
B.M. No. 12. 12. 17.13, 5; X 4.
566
SICISTA
P. 4 very small; I\T.3 small. M.i and M.2 with four main cusps, the outer
of which tend to be larger than the inner, the cusps, which are situated roughly
at the corners of the teeth, separated by relatively wide folds. The cusps are
evidently long retained, and the general effect is complex. Lower molars like
those of the upper series, but M.3 less reduced, and the folds separating the
cusps appear as a relatively more important element in the tooth. The general
dental effect is reminiscent to a degree of that present in Cricetinae.
Incisors not grooved. Mandible without per-
foration in the angular process.
Size very small indeed, under 100 mm. head
and body length as far as seen. Tail considerably
longer than head and bodv, moderatelv haired.
Forefoot without peculiarities. Hindtoot with
very short hallux, D.5 somewhat reduced, the
foot very narrow.
There are 8 mammae in a specimen of S.
norvegica given to me by Mr. J. L. Chaworth-
Musters, who tells me that this species hibernates
for a considerable portion of the year in Norway.
I am much obliged to ^f r. Chaworth-Musters for
much information concerning this genus.
Forms seen: beluliiia, camiatii, coiicolor, leatheini, Imiger, moiitana, nomgica,
subtilis, tiansc/iaiiica, trizona.
Two groups are currentlv recognized, those species in which there is a
middorsal stripe present, tvpified by subtilis, and those in which this is absent,
typified by concolor.
The synonymy of the subtilis group has been published by Chaworth-
Musters, Ann. Mag. Nat. Hist. ser. 10, vol. xiv, p. 554, 1934; in this paper the
range of some of the races will be found.
S. bctuliiia differs from 5. subtilis in the relatively longer hindtoot and tail.
The concolor group appears to consist of forms which are rather doubtfully
separable from each other as full species, with the exception of napaea,
which was described as near flavus, but which, according to Vinogradov,
has manv characters which separate it from the concolor group, including
the genitalia.
Fig. 146
SlCIST.Ji SUBTILIS LORIGER.
Cheekteeth: X g.
List of Named Forms
(The references and type localities for all members of the family Dipodidae
are the work of Mr. G. W. C. Holt.)
Not allocated to group
I. .SICIST.^ X.AP.\E.\. Hollistcr
igi2. Smiths. Misc. Coll., LX, no. 14, p. 2.
Tapucha, Altai Mountains, Siberia.
SICISTA 567
subtilis Group
2. SICISTA SUBTILIS SUBTII.IS, Pallas
1773. Reise, ii, p. 705.
Mouth of Ural River, Siberia.
Synonym: vagus, Pallas, 1778, Nov. Spec. Quadr. Glir, Ord. p. 327.
Russia.
Uiteatiis, Lichtenstein, 1823, Eversmann's Reis. Buch., p. 123.
3. SICISTA SUBTILIS LORIGIvR, Nathusius
1840. Nordm. Voy. Demidoff, iii, p. 49.
Odessa, South Russia.
Synonym: nordmanni, Keyserling & Blasius, 1840, Wirb. Europ., X,
p. 38. Crimea.
4. SICISTA SUBTILIS PALLIDA, Kaschkaroff, ex Vinogradov
1926. Rodents of Turkestan, p. 1 1, in Usbekistan Exp. Plant. Prot.
Djetysu, Turkestan. •
5. SICISTA SUBTILIS TRIZONA, Petenyi
1882. Termeszetrajzi Fuzetek, V, p. 103.
Hungary.
With alternatives interzomis, inter striatus, tripartitus, virgulosus and
tristriatus: same reference.
6. SICISTA SUBTILIS SIBIRICA, Ognev
1935. Abstr. Works. Zool. Inst. Moscow, 2, p. 54.
Central part of Russian .-Mtai.
7. SICISTA SUBTILIS SEVERTZOWI, Ognev
■935- -■^bstr. Works. Zool. Inst. Moscow, 2, p. 54.
Kamennaja Steppe, Bobrow, Voronesh, South Russia.
8. SICISTA BETULINA BETULIN'A, Pallas
1778. Nov. Spec. Quad. Glir. Ord., p. 90.
Banks of River Ischim, Siberia.
9. SICISTA BETULINA MONTANA, M^hely
1913. .AUattani Kozlem, 12, p. 69.
Zuberecz, Hungary.
10. SICISTA BETULINA .STR.ANDI, Formozov
193 1. Folia Zool. Hydrob. Riga, 3, p. 79.
Caucasus, Ortschaft Igera Hohe, 2,100 m. Distrikt Utschkulak,
Karatschai.
11. SICISTA BETULINA NORVEGICA, Chaworth-Musters
1927. Ann. Mag. Xat. Hist. 9, XIX, p. 542.
Surendal, Xordmore, Norway.
concolor Group
12. SICISTA CONCOLOR, Buchner
1892. M^l. Biol. Acad. St. Petersb., xiii, p. 268.
Si-ning, Kansu, China.
Synonym: (?)ti'«goWi, Jacobi, i923,.-\bh. Mus. Dresden, 16, no. i, p. 15.
Hsueschau, West China.
S68 SICISTA— ZAPODINAE: EOZAPUS
i.V SICISTA LEATHKMI, Thomas
1S93. Ann. Map. Nat. Hist. 6, XI, p. 184.
Krishnye Valley, Wardwan, Kashmir.
14, SICISTA FLAVCS, True
1894. Proc. U.S. Nat. Mus. Washington, X\'II, p. 341.
Kashmir.
15, SICISTA TIANSCHANICA, Salensky
1903. Ann. Mus. St. Pctersb., viii, p. 17.
Thian Shan, Chinese Turkestan.
16. SICISTA C.\L"DATA, Thoma.s
1907. Proc. Zool. Soc. London, p. 413.
Korsakoff, Saghalien.
17. SICISTA CAUCASICA, VinoKradov
1925. Proc. Zool. Soc. London, p. 584.
Kuban, North Caucasus.
Subfamily ZAPODINAE
Geogr.^phic.al Distribution. — The greater part of Canada and the United
States; China, statesof KansuandSzechuan.
Number of Genera. — Three.
Characters. — Skull not essentially different from the Sicistinae; interorbital
constriction moderate; jugal in contact with lachrymal;
anterior end of nasals projecting beyond premaxillae; palate not continued
backwards behind M.3; and not terminating in spinous process. Cheekteeth
semihypsodont, becoming flatcrowned in American genera, but with relatively
high cusps in the Asiatic genus, which has also wide re-entrant folds. External
form considerably modified for bipedal saltatorial life; hindlimbs and tail
lengthened. Cheekteeth i or 'J. Bullae relatively small, not inflated. (Meta-
tarsals normal. Zygoma simple.)
Key to the Genera of Zapodinae
Cheekteeth with high cusps and broad re-entrant folds, showing no sign
of becoming flatcrowned (so far as ascertainable). Eozapus
Cheekteeth with low cusps and moderate re-entrant folds, the folds con-
siderably narrowed; or the teeth becoming flatcrowned.
Cheekteeth nearly completely flatcrowned, with numerous narrow-
isolated islands on crown surface in adult. Napaeozapus
Cheekteeth less flatcrowned, without numerous small isolated islands
on crown surface in adult. Zapus
Genus i. EOZAPUS, Preble
1899. Eozapus, Preble, North Amer. Fauna, o. 15, p. 37.
EOZAPUS— ZAPUS 569
Type Species. — Zapus setchuanus, Pousargues.
Range. — China: Szechuan and Kansu.
Number of Forms. — Two.
Char.\cters. — Like Zapus, ne.xt to be described, but with, in the three
skulls examined, a considerably different dental pattern.
The cheekteeth have very wide re-entrant folds separating high cusps and
ridges. Four outer, one inner folds in the upper main teeth (M.i and M.2);
in the lower teeth there is one very wide main outer fold, and a small extra one
in front of it; three inner folds, and in M.i also an anterior fold. Teeth more
or less prismatic, showing no sign of becoming flatcrowned ; it would be desirable
to examine a much larger number before giving any guarantee that these
characters are constant. In the upper jaw, M.3 is considerably reduced; P. 4 is
present, and minute.
Forms seen : selchuaniis, vicimis.
List of Named Forms
1. EOZAPUS SETCHUANX'.S SETCHUANUS, Pousargues
1896. Bull. Mus. Paris, no. 2, p. 13.
Szechuan, China.
2. EOZAPUS SETCHUANUS VICINUS, Thomas
1912. Ann. Mag. Nat. Hist. 8, X, p. 402.
Kansu, China; 46 miles south-east of Tao-chou.
Genus 2. ZAPUS, Coues
1876. Bull. U.S. Geol. & Geogr. Surv. Terr., ser. 2, vol. i, p. 253.
Type Species. — Diptis hiidsonius, Zimmermann.
Range. — Canada and the United States; forms named from Alaska, Hudson
Bay, Labrador, Saskatchewan, British Columbia, Oregon, Idaho,
Wyoming, California, Nevada, Utah, Colorado, New' Mexico, Pennsvlvania.
Number of Forms. — Twenty-si.x.
Char.\cters. — Superior margin of canal for nerve-transmission frequentlv
fused to maxilla in adult. Angular portion of mandible with
no perforation, strongly ridged for muscle attachment, the inner side of the
angular portion pulled inwards, after the manner of that of Aplodontia though
less extreme than in that genus. Incisive foramina large, well open, generallv
extending to toothrows. Upper incisors one-grooved, lower incisors plain.
Cheekteeth 5, the premolar minute; M.3 smaller than other molars. In the
upper series, there is one main fold internally, curbing forwards, and apparently
originally four outer folds, some of which tend to isolate as islands in the adult.
The folds are wider and more definite than in Napaeozapus, and never so far
as seen isolate to the same extent. Lower molars with two outer, four inner
folds.
Fig. 147. Zapus hudsonius hudsonius, Zimmemiann.
B.M. No. 135.1.7.95, 3; .-. 3*.
Fig. 148. Zapus hudsonius hudsonius, Zimmcrmann.
B.M. No. 95.1.7.95, o; X 3*-
ZAPUS
571
Mammae normally 8 (Preble). Hindlimbs elongated, the hindfeet narrow,
with the three central digits long, D.5 reaching about to base of D.4, the hallux
much shortened. Forefoot narrow; D.3 and D.4 slightly longer than D.2 and
D.5 ; pollex rudimentary. Tail very long, moderately or poorly haired, the
scales visible; a small pencil at the end. Cheek-pouches present.
Forms seen : htidsonius, tnnotatiis, ciimpestris, ladas.
The genus is revised by Prelilc (North .\mer. Fauna, 15, p. 13, 1899). -'^1
species admitted appear closely allied to each other.
Fig. 149. Z.\PLS hudsonius hudsonils, Zimmermann.
Cheekteeth: B.M. No. 95.1.7.95, c?; >'■ 15.
List of Named For.ms
1. Z.A.PUS HUDSONIUS HUDSONIUS, Zimmermann
1780. Geogr. Gesch., vol. 2, p. 358.
Hudson Bay.
Synonym: hudsonius hardyi, Batchelder, Proc. New England Zool.
Club, I, p. 5, 1899. Mount Desert Island, Hancock
County, .Maine.
Trouessart quotes as synon\-ms :
longipes, Zimmermann, Pennants Arctic Zool., 1787, p. 131.
canadensis, Davies, Trans. Linn. Soc, 1798, IV, p. 157.
labradorius, Turton, 1806, Syst. Nat., I, p. 99.
nemoralis, Geoffrov, Diet. Class. Hist. Nat., VII, p. 323,
1825.
daviesi and soricinus, Rafinesque, Somiol., p. 14, 1810.
leonurus and megalops, Rafinesque, Amer. Monthly Mag.,
1818. p. 446.
microcephalus, Harlan, 1839, Proc. Zool. Soc. London, VII,
p. I.
2. ZAPUS HUDSONIUS LAD.\S, Banes
1899. Proc. New England Zool. Club, I, p. 10.
Rigolet, Hamilton Inlet, Labrador, Canada.
572 ZAPUS
3. ZAPL'S HUDSOXIfS ALASCF.NSIS, Merriam
1897. Proc. Biol. Soc. Washington, XI, p. 223.
Yakutat Bay, Alaska.
4. ZAPL'S HUDSOML'S AMERICANUS, Barton
1799. Trans. Amer. Philos. Soc, IV, p. 115.
Near Philadelphia, Pennsylvania.
5. ZAPUS HUDSUNIUS CAMPESTRIS, Preble
1899. North Amer. Fauna, no. 15, p. 20.
Bear Lodge Mountains, Crook County, Wyoming.
6. ZAPUS TENELLUS, Merriam
1S97. Proc. Biol. Soc. Washington, XI, p. 103.
Kamloops, British Columbia, Canada.
7. ZAPUS PRINCEPS PRINCEPS, Allen
1893. Bull. Amer. Mus. Nat. Hist., V, p. 71.
Florida, La Plata County, Colorado.
S. Z.-\PUS PRINXEPS MINOR, Preble
1899. North Amer. Fauna, no. 15, p. 23.
Wingard, Carlton House, Saskatchewan, Canada.
9. ZAPUS PRINCEPS OREGONUS, Preble
1899. North .Amer. Fauna, no. 15, p. 24.
Elgin, Union County', Oregon.
10. ZAPUS PRINCEPS CINHREUS, Hall
1931. Univ. Calif. Publ. Zool. XXXVII, p. 7.
Pine Canyon, Raft River Mountains, Boxelder County, Utah.
11. ZAPUS PRINCEPS CURTATUS, Hall
193 1. Univ. Calif. Publ. Zool. XXXVII, p. 7.
Big Creek, Humboldt County, Nevada.
12. ZAPUS PRINCEPS P.AL.ATINUS, Hall
1931. Univ. Calif. Publ. Zool. XXXVII, p. 8.
Wisconsin Creek Toyabe Mountains, Nye County, Nevada.
13. ZAPUS PRINCEPS KOOTE.NAYENSIS, Anderson
1933. Bull. Nat. Mus. Canada, no. 70, p. 108.
British Columbia, Canada, near summit of Green Mountain, head of
Murphy Creek, about 10 miles north (jf Rossland West Kootenay
district.
14. ZAPUS PRINCEPS IDAHOENSIS, Davis
1934. Joum. Mamm. Baltimore, 15, p. 221.
Valley County, Idaho, 5 miles east of Warm Lake.
15. Z.-\PUS PRINCEPS UTAHENSIS, Hall
1934. Occ. Pap. Mus. Zool. Mich., no. 296, p. 3.
Beaver Creek, Manilla, Daggett County, Utah.
16. ZAPUS MAJOR, Preble
1899. North Amer. Fauna, no. 15, p. 24.
Warner Mountains, Lake County, Oregon.
17. ZAPL'S NI-VADENSIS, Preble
1899. North Amer. Fauna, no, 15, p. 25.
Rubv Mountains, Elko County, Nevada.
I
ZAPUS— NAPAEOZAPUS 573
i8. ZAPUS TRINOTATUS TRINOTATUS, Rhoads
1894. Proc. Acad. Nat. Sci. Philadelphia, p. 421.
Lulu Island, Fraser River, British Columbia, Canada.
Synonym: imperator, Elliot, 1899, Field Columb. Mus. Publ. 30, zool.
ser., vol. I, p. 228. Sieg's Range, Elwah R., Clallam
County, Washington.
19. ZAPUS TRINOTATUS ALLENI, Elliot
1898. Field Columb. Mus. Publ. 27, zool. ser., vol. i, p. 2x2.
Pyramid Peak, Lake Tahoe, Eldorado County, California.
20. ZAPLS TRINOTATUS EUREKA, Howell
1920. Univ. Calif. Publ. Zool., XXI, p. 229.
Fair Oaks, Humboldt County, California.
21. ZAPUS LUTEUS LUTEUS, Miller
1911. Proc. Biol. Soc. Washington, XXIV, p. 253.
Espanola, Santa Fe County, New Mexico.
22. ZAPUS LUTEUS AUSTRALIS, Bailey
1913. Proc. Biol. Soc. Washington, XXVI, p. 132.
Socorro, Socorro County, New Mexico.
23. ZAPUS MONTANUS, Merriam
1897. Proc. Biol. Soc. Washington, XI, p. 104.
Crater Lake, NIount Mazama, Klamath County, Oregon.
24. Z.'^PUS OR.'VRIUS, Preble
1899. North Amer. Fauna, no. 15, p. 29.
Point Reyes, Marin County, California.
25. ZAPUS PACIFICUS, Merriam
1897. Proc. Biol. Soc. Washington, XI, p. 104.
Prospect, Rogue River Valley, Jackson County, Oregon.
26. ZAPUS S.-^LT.^^TOR, .\llt-n
1899. Bull. Amer. Mus. Nat. Hist.. XII, p. 3.
Telegraph Creek, Stikine River, British Columbia, Canada.
Genus 3. NAPAEOZ.^PUS, Preble
1899. NAPAEOZAPUS, Preble, North Amer. Fauna, no. 15, p. 33.
Type Species. — Zapus insignis, Miller.
Range. — Eastern North America; forms described from New Brunswick,
Ontario, Wisconsin and North Carolina.
Number of Forms. — Four.
Characters. — Like Zapus, but M.2 appearing relatively larger, and dental
pattern differing; the teeth nearly completely flatcrowned,
the folds extremely narrow, isolating and dividing on the surface of the tooth,
so that there may be ten or more small islands on the tooth surface in adult.
Lower molars similar to the upper series. P.4 absent. Interorbital constriction
greater than Zapus (Preble).
Essential external characters as Zapus (but tail with white tip).
574 NAPAEOZAPUS— CARDIOCRANIINAE: SALPINGOTUS
Remarks. — The dental peculiarities of these three types seem sufficient to
warrant the retention of Eosapiis and Napaeosapiis as full
genera. The three types are well figured in Preble's revision o( Zapiis. A much
larger series, however, would be welcome, as comparatively few skulls are
represented in London.
Forms seen: insignis.
List of Named Forms
(Genus revised by Preble, North Amer. Faima, no. 15, p. ■;^, 1899.)
1. NAPAEOZAPU.S INSIGNIS IN.SIGNI.S, Millir
1891. .Amer. Nat., XXV, p. 743.
Restigouche River, New Brunswick, Canada.
2. NAPAi:()ZAPL'S INSIGNIS ROANENSIS, Preble
1S99. North Amer. Kauna, no. 15, p. 35.
Roan Mountain, Mitchill County, North Carolina.
3. NAPAl-;0/-APllS INSIGNIS ABIETORUM, Prchlc
1S99. North Amer. Fauna, no. 15, p. 36.
Peninsula Harbour, north shore of Lake Superior, Ontario, Canada.
4. NAPAEOZAPUS INSIGNIS 1" RITECTANUS, Jackson
1919. Proc. Biol. Soc. Washington, XXXII, p. 9.
Crescent Lake, Oneida Coimty, Wisconsin.
Subfamily CARDIOCRANIINAE
Geographical Distribution. — Central Asia: Northern Tibet, Mongolia
(Gobi, Altai), and Afghanistan.
Number of Genera. — Two.
Remarks. — The two genera inchuiei-l in this subfamilv are represented at
the British Museimi by only one badly smashed skull, type of
Salpingotiis tliuiiiasi.
The important character in this subfamily is that the three central metatarsal
bones of the hindfoot are not fused, in which the genera agree with Zapodinae
and Sicistinae, but differ from Kuchoreutinae and Dipodinae. They agree,
however, with the higher Jerboas in that the bullae and mastoids are inflated,
this inflation indeed being carried much further in this group than in any other
Jerboas, according to figures published of the skulls; and in the general form of
the zygoma, which differs from Euctioreutes but stands nearer the form found
in Dipodinae in that there are already two portions, a horizontal and a vertical;
but these portions are connected by a curvature in Cardiocraniinae, and do not
apparently form such a sharp angle with each other as they do in the Dipodinae.
The cheekteeth apparently also agree more with Dipodinae than with Zapodinae.
Genus i. SALPINGOTUS, Vinogradov
1923. SALPINGOTUS, Vinogradov, Kozlow "Mongolia & Amdo," p. 540.
Type Species. — Salpinnotiis kozlovi, Vinogradov.
SALPINGOTUS— CARDIOCRANIUS S75
Range. — Known from the Gobi desert, and Afghanistan.
Number of Forms. — Three.
Characters. — "Hindfoot with three toes and three not-ankyloscd meta-
tarsals. Bullae as in Cardiocranius. Zygomata broad in their
anterior half, a process directed obliquely downwards and backwards and rising
from the middle of each zygomatic arch. Jugal not reaching lachrymal. . . .
Infraorbital foramen relatively narrower than in true Jerboas. Bony palate
spreading far backwards, pterygoids very short. A horizontal process rising
externally between angular and articular process of mandible. . . . Upper
incisors without grooves. Cheekteeth, P. i, M. i| ; . . . Toes covered with long
curved hairs which form a thick brush" (Vinogradov).
The mandible as figured is strongly reminiscent of the formation found in
Aplodontia.
The degree of development of the downwardly projecting process on the
zygomatic arch differs, I believe, in the different species. The tail is described
as very long in the type species, and normal, but shorter and more or less of
the thickened shape in crassicauda, which has a perforation in the angular portion
of the mandible. S. thomasi belongs evidently to the thicktailed group. The
mastoids and bullae are enormous, the mastoids occupying about a third of
the upper surface of the skull.
According to Vinogradov, the "tubercular structure of molars is visible
only in young and subadult specimens."
Forms seen : thomasi.
List of Named Forms
kozlovi Group
1. SALPINGOTUS KOZLOVI. Vinogradov
1923. K02I0W "Mongolia & Amdo," p. 540.
Gobi, Mongolia (near the ruins of Khara-khoto).
crassicauda Group
2. SALPINGOTUS THOMASI, Vinogradov
1928. Ann. Mag. Nat. Hist. 10, I, p. 373.
Afghanistan.
3. SALPINGOTUS CR.'\SSICAUDA. Vinogradov
1924. Zool. Anzeiger, 61, p. 150.
Gobi Altai.
These forms are here treated as separate groups on account of the tail
formation, which has elsewhere in this family been used as a generic character.
Vinogradov suggests that 5. crassicauda may ultimately have to form a new
genus. The mammae of this species are quoted by him as p. 2 — 2 ; i. 2 — 2.
Genus 2. CARDIOCRANIUS, Satunin
1903. Annuaire Mus. St. Petersb., vii, p. 582.
Type Species. — Cardiocranius paradoxus. Satunin.
37 — Living Rodents — I
576 CARDIOCRANIUS— EUCHOREUTINAE
Range. — River Scharogol-dschin, in Nan Shan, Central Asia.
NiTMBER OF Forms. — One.
Remarks. — This genus is not represented at the British Museum. I am not
including any genus I have not seen in my keys, as to endeavour
to key an unexamined genus is always difficult, in the case of a Muroid im-
possible.
Vinogradov has keved the genera Salpingotus and Cardiocranius in cranial
characters as follows:
" Infraorbital canal complete, its external wall being in contact with wall
of maxilla. Zygoma with well-developed process rising from its
middle and directed downwards posteriorlv. Anterior ends of
nasals projecting beyond premaxillae. Palate bones considerably
longer than upper toothrow, projecting unusually backwards.
Salpingotus
Infraorbital canal incomplete, its external plate not reaching wall of
maxilla. Zygomatic arch simple, without any process rising from
its middle. Anterior ends of nasals not projecting beyond pre-
maxillae. Palate bones about as long as upper toothrows, not
considerably projecting backwards. Cardiocranius"
In addition to these differences, it may be noticed that in Cardiocranius, as
described, there are five toes to the hindfoot, the outer toes placed higher up
than the central three, the hallux considerably higher than D.5, and functionless.
The tail is described as broad and flattened, like that of Pygeretmus. Cheek-
teeth 4, considered in the original description to be similar to those of Dipus and
Al!act<ii;a, having nothing in common with those of Euchoreutes. Jugal not
reaching lachrymal. Apices of bullae in contact. Upper incisors grooved.
Mastoids enormous, as in Salpingotus, and evidently mandible strongly inflected,
in a similar manner.
Both Cardiocranius and Salpingotus are very small forms; Cardiocranius has
a head and bodv measurement of 73 mm. (type).
The genus is, I believe, exceedingly rare, and still onlv known by a very few
specimens, though first described over thirty years ago.
List of N.^med For.ms
I. CARUIOCRANIUS P.ARADOXUS, Satunin
1903. .Annuaire Mus. St. Petersb., vii, p. 584.
Nan Shan (Scharogol-dschin), Central .\sia.
Subfamily EUCHOREUTINAE
Geographical Distribution. — China: known from Yarkand, Chinese
Turkestan, and the Alashan desert (Inner
Mongolia, bordering Kansu).
Number of Genera. — One.
I
EUCHOREUTINAE : EUCHOREUTES 577
Characters. — Hindfoot with three central metatarsals fused to form a
cannonhone. Differing from the Dipodinae, with which it
shares these characters, in cranial and dental characters.
Jugal slanting gradually up towards lachrymal; rostrum much elongated;
frontals with constriction placed considerably behind the lachrymals (skull as
a whole broad, and constriction noticeable but certainly not excessive, at any
rate as compared with a typical Murine); lachrymal small; bullae considerably
inflated, their apices in contact, mastoids relatively large; mandible with
perforation in angular process. Incisive foramina large, and a large second pair
are situated between the toothrows. Ear abnormally large. Os penis present
(Vinogradov); skeleton of hindfoot as in Allactaga group (Vinogradov). Cheek-
teeth, described below, differing considerablv in pattern from Dipodinae; j\1.3
(so far as seen, and as figured by Vinogradov) vestigial.
Genus i. EUCHOREUTES, Sclater
1890. EUCHOREUTES, Sclater, Proc. Zool. Soc. London, p. 610.
Type Species. — Euchoreutes naso, Sclater.
Range. — As in the subfamily Euchoreutinae.
Number of Forms. — Two.
Ch.'\r.\cters. — As indicated above. Zygomata very narrow. Jugal in contact
with lachrymal. Nasals projecting beyond premaxillae.
Incisors white, the upper ones plain. Palate broad, projecting beyond M.3,
terminating in spinous process. Pterygoid fossae deep.
Cheekteeth I, hypsodont, narrow. M.3 extremely reduced, simple, smaller
than P.4. (Two skulls seen only). M.i slightly larger than M.2, with four main
cusps, each cusp separated from its neighbours by a deep valley. Posterior part
of teeth straight, not rounded. Lower teeth narrow, like the upper series in
pattern, but M.i with an extra shallow re-entrant fold posterior to second inner
cusp, and M.2 with this peculiarity, and with a very small extra cusp, external,
anteriorly.
Cusps of cheekteeth high.
Mammae 8 (Sclater). Ears extremely elongated, appearing about half the
length of head and body. Snout elongated. Tail considerably longer than head
and body, well haired, with a black and white brush at the end. Forelimbs
short, foot with five digits, claws well developed. Hindfoot extremely elongated,
narrow; five digits present, but only three reaching the ground. Fur long and
soft.
Remarks. — Although agreeing with Dipodinae in the highly specialized
character of the skeleton of the hindfoot, this genus differs so
markedly from them in dental characters, and also in the important character
of the zvgoma (in which it is transitionarv towards Sicista and Zapodinae), that
I think the subfamily Euchoreutinae must be retained.
Forms seen : naso.
Fig. 150, EuCHOREUTES naso naso, Sclater.
B.M. No. y().ii.5.6, V; ■ 3.
Fic. 151. EucHOREUTES NASO NASO, Sclater.
B.M. No. 99.1 1.5.6, V; X 3.
EUCHOREUTES— DIPODINAE
S79
List of Named Forms
1. EUCHOREUTES NASO NASO, Sclater
i8go. Proc. Zool. Soc. London, p. 6io.
Yarkand, Chinese Turkestan.
2. EUCHOKKUTI^S NASO ALASCHANICUS, Howell
igzS. Proc. Biol. Soc. Washington, XLL P- 42-
Inner Mongolia, Alashan desert, lOO miles north-west of Ningsia,
Kansu.
Fig. 152. EUCHOREUTES naso naso, Sclater.
Cheekteeth: B.M. No. 99.11.5.6, ?; X 11.
It may be noted that according to Vinogradov's latest work on the famiiv,
the three metatarsals of Eiiclioreiites are less completely fused than in the
Allactaga group or in the Dipus group (Faune de L'URSS, Inst. Zool. Ac.
Sci. URSS, 1937, III, no. 4, p. 49, fig. 5).
Subfamily DIPODINAE
Geographical Distribution. — .\frica: Senegambia and Morocco to Egypt
and Somaliland; Europe, across southern
Russia nearly to Roumanian border (Dnieper). .\sia I\Iinor, Persia, Arabia,
Afghanistan, large portions of Russian .\sia; Baluchistan, Kashmir; China east
to Mongolia and Chihli.
Number of Genera. — As here understood, there are eight genera in two
generic groups, the equivalent of the Allactaginae
and Dipodinae of \'inogradov.
sSo DIPODINAE: ALLACTAGA
Characters. — Jugal in two portions, a horizontal and a vertical, the portions
forming a sharp angle with each other, and not connected by
a curvature (compare Cardiocraniinae). Lachrymal enlarged. Frontals broad,
very rarely showing anv constriction; if this is present, it is very slight, and
situated immediately behind the lachrymals. Functional digits of hindfoot 3.
Three central metatarsals fused to form a cannonbone. Bullae moderately or
greatly inflated. Externally highly specialized for bipedal saltatorial life.
The subfamily is here divided into tw'o generic groups; a key to these groups
has already been given (pp. 563, 564).
The Allactaga Group
Anterior vertical branch of the jugal not greatly broadened. Bullae feebly
inflated, their apices not in contact. Anterior ends of nasals not reaching
alveoli of upper incisors. Upper incisors not grooved, pro-odont. Ears large
(larger than in Dipiis group, but smaller than in Euchoreutinae). Digits of
hindfoot five (four in one species of Allactaga), three only functional. Os penis
absent.
Kf.y to the Genera of the Allactaga Group
Cheekteeth relatively complex; upper main teeth with three external
folds, lower main teeth with three internal folds. Vertical branch
of zygoma about as broad as horizontal branch. Allactaga
Cheekteeth simplified; in adult upper main teeth with only two external
folds, M.2 lower with only two external folds. Vertical branch of
zygoma narrower than horizontal branch.
Tail longer than head and body, narrow, evenly round, tufted ter-
minally. Margins of supraorbital less angular. Alactagulus
Tail shortened, flattened and thick throughout most of its length, not
tufted terminally. Margins of supraorbital more angular.
Pygeretmus
Genus i. ALLACTAGA, Cuvier
1836. Allactaga, Cuvier, Proc. Zool. Soc. London, p. 141.
1841. Scarturus, Gloger, Gemeinn. Naturgesch, i, p. 106. Dipiis tetradactylus, Lichten-
stein.
1844. SciRTOMYS, Brandt, Bull, phys.-niath. Ac. Sci. St. Petersb., II, p. 220. DIpus
tetradactylus, Lichtenstein.
1841. Scirtetes, Wagner, Gelehrte Anz. k. bay. .Ak. Wiss. MOnchen, XII, p. 413.
New name for Allactaga, Cuvier.
Type Species. — Dipus alactaga, Olivier.
Range. — North Egypt; Mesopotamia; North Arabia; Asia Minor; the
Caucasus, and southern European Russia (quoted by Vinogradov
from former Chernigov, Kursk, southern part of Tula, Riasan, Tambor, Pensa,
southern part of Kazan, Samara, LIfa govts., westwards to Dnieper; also Astra-
khan and Kalmuck districts, and the Crimea); North Persia; throughout
Fig. 153. Allactaca euphratica, Thomas.
B.M. No. 5.7.2.12, cJ; X zj.
Fig. 154. .\ll.\ct.^ga elphr.\tica, Thomas.
B.M. No. 5.7.2.12, a; x 2j.
ss-
ALLACTAGA
Russian Turkestan and South-west Siberia to Semipalatinsk and the Ahai ;
Afghanistan, probably Kashmir; the Altai Mountains; Persian Baluchistan;
Kansu, Chinese Turkestan, Mongolia, Shansi, North Chihli, and Trans-
baikalia.
Number of Forms. — About twenty-nine.
Ch.^r.^cters. — Frontals broad, braincase very broad. Lachrymal large.
Bullae feebly inflated except in biillata (not seen), and
slightly less than usual in hotsoni. Mastoids not appearing in superior aspect
of skull, so far as seen. Incisive foramina relatively large; usually a well-
marked second pair present between the toothrows. Skull without supraorbital
ridges. Infraorbital foramen very large indeed. Mandible with angular process
perforated, and root of incisor forms process below and beside condyle.
Fig. 155. Allactaca euphratica, Thomas.
Cheekteeth: B.M. No. 5.7.2.12, 0'; 8.
Jugal in two portions, a horizontal and a vertical, as in all higher Jerboas.
Cheekteeth |, semihypsodont, very complex; P. 4 normally minute; M.3
considerably reduced, but larger than the premolar except in the sibirica group.
M.I and M.2 each with three external re-entrant folds, the middle one of which
is small, and each tooth with one internal fold. Sometimes the small middle
external fold may wear out, and the whole pattern ultimately becomes lost.
Occasionally traces may be seen of a very small fourth extra outer fold in the
posterior part of tooth. M.3 with one inner, three outer folds when cut,
apparently.
In the lower teeth, M.i has one small fnmt told, three inner folds, and two
outer ones, the middle inner one being small; M.2 is like M.i but without the
anterior fold. M.3 with one outer fold, and one or two inner ones, which
wear out.
\
ALLACTAGA 583
Size moderately small to largest of family. Ears very large, though not
comparing with those of Kiichoreiites. Forefoot with five digits, the pollex
short ; claws well developed ; hindfoot perissodactyle with three functional digits,
and with two subcqual outer digits placed high on the leg, not reaching the
ground. Tail longer than head and body, with a black and white brush
terminally, round and thin (normal) throughout most of its length. The tail
well haired. Hindlimb enormously elongated, the claws of the three central
digits with large pad present under each claw.
Lyon states: " Allactaga has the cannonbone of Dipus, but on either side of
it is a small non-functional toe, consisting of a metatarsal and a digit; . . . the
cervical vertebrae show a tendency towards consolidation, but not that complete
fusion found in Dipus." The outer digits appear from Ljon's figure to be
situated where the cannonbone finallv breaks into three for the central digits.
In A. letradactyla. North Egjpt, the hallux is suppressed, but D.5 remains.
On this account it has long been regarded as type of a genus " Scarturus."
But the presence or absence of a functionless digit is scarcely to be considered
a character of generic importance, as is seen when dealing with the genus
Dipodomys. Neither Pocock nor Vinogradov were able to find any differences
between the skulls of "Scarturus" and Allactaga; Vinogradov remarks, "The
skull of Scarturus is very similar to that in Allactaga, especially the members of
the A. euphratica group, the resemblance may be seen not only in general
outlines but also in dimensions; the interorbital constriction is, however, con-
siderably broader than in A. euphratica . . . the frontoparietal border of the
squamosal has no incisure . . . the incisive foramina in Scarturus are somewhat
broader and more opened than in Allactaga; it is impossible to find, however,
more important cranial differences between the genera."
Compared, however, with Dipodomys it will be seen that both D.i and D.5
in Allactaga are equal in size; whereas in Dipodomys the hallux is much shorter
than D.5, and higher on the leg. The complete disappearance of D.5 in
A. tetradactyla in this case may, therefore, I think be regarded as a specific
group character, in the present state of our knowledge, and it is curious that in
the several specific groups of Allactaga only this one rare species has for no
apparent reason entirely lost the digit on one side of the foot, but retains the
other functionless digit normally and unreduced.
As remarked above (p. 561), a skull of A. elater in the British Museum,
no. 13. 12. 1. 3, appears to have a very small extra molar situated at the back of
the series on one side of the jaw.
The A. sibirica group differ rather noticeably from the other species in that
M.3 has become more reduced, and apparently the premolar less so than in
others, so that the premolar is only slightly smaller than M.3, or subequal with
it, or even sometimes, according to Vinogradov, a little larger than it. The bodily
size is larger than in the elater group. Vinogradov has keyed the species occurring
in the U.S.S.R. The measurements here used are based on his measurements,
and also on those of the British Museum specimens.
I am inclined provisionally to recognize five specific groups of Allactaga, as
follows:
584 ALLACTAOA
1. The sihirica group, with dental characters indicated above.
2. The tctradactvla group, with characters as indicated above.
3. The elater group. Small forms; hindfoot less than 65 mm. (usually about
48-55 according to B.M. material) in elater. slightly larger on average,
52-58 in euphratica. Including hotsoni (hindfoot 58 in type specimen;
bullae more inflated than others).
4. The major group. Usually giant forms; typically hindfoot over 85; in
scTertzovi about 70-80.
5. The idUiamsi group. Like the last, but smaller, hindfoot 65-70, or smaller
than major group, larger than elater group, and differing from the major
group in the characters of the penis, according to Vinogradov.
I am unable to allocate A. hullata, Allen, as I have not seen it. It was de-
scribed as a species with audita! bullae about three times the size of ^. mongolica
(sibirica group), which may indicate that it should form a group by itself.
The proportions of the cheekteeth are, according to Vinogradov's key,
normal, i.e. not agreeing with sibirica group (hindfoot 70 mm.).
It should be noted that in tetradactyla, as far as known, the size (head and
body about no, hindfoot about 57) agrees with the elater group, and it is
probablv a close ally of euphratica.
The nomenclature of the species of the genus here differs from that of
Vinogradov and follows that of Chaworth-Musters, who has published several
papers on this point. My thanks are due to Mr. Chawortli-Musters for much
information regarding this genus.
Forms seen: aralychcnsis. caucasica. decumana. elater, euphratica, hotsoni,
indica, "jaculus" (=major), laticeps, mongolica. rikkheili. salicns, seiertzovi,
salt at or. tetradactyla. uilliamsi.
List of X.^med Forms
incertae sedis
ALL.-\CTAGA .\RUND1NIS, F. C
u\ ler
1836. Trans. Zool. Soc. London, II. p. 134.
"Barban.'" (? error).
Not allocated to group
2. ALLACTAGA BULLATA, G. Allen
1925. .Amer. Mas. Nov. 161, p. 2.
Tsagan Nor, Mongolia.
tctradactvla Group
3. ALLACTAGA TETRADACTYLA. Lichtenstein
1823. Verz. Doublet, Mus. Berlin, p. 2.
Eg>'pt, near Alexandria.
Synonym: brucei. Lesson, Man. Manini., p. 253, 1S27. Barca.
abyssiiiicus, Illiger. 1804, Abh. ph. Kl. k. Akad. Wiss., Berlin,
p. 77. (?)Eg>pt.
ALLACTAGA 585
elater Group
4. ALLACTAGA EUPHRATICA, Thomas
1881. Ann. Mag. Nat. Hist. 5, XVIII, p. 14.
Mesopotamia.
5. ALLACTAGA ELATER I;L.\TER, Lichtenstcin
1825. Abh. k. Akad. Wiss. Berlin, p. 55.
Turkestan ; desert region.
6. ALLACTAGA ELATER STRANDL Heptner
1934. Folia Zool. Hydrob. 6, p. ig.
Transcaspia, "in der Nahe von Mer\\% Transkaspien."
7. ALLACTAGA EL.VFER CAUCASL\, Brandt
1855. Mi\. Biol. Ac. St. Petersb., II, p. 303.
Transcaucasia (Saljany, Mugan-Steppe).
8. ALLACTAGA ELATER KIZLJARICUS, Satunin
1907. Mitt. Kaukas. Mus. 3, p. 45.
N.-E. Caucasus.
q. ALLACTAGA ELATER ARALYCHENSIS, Satunin
1901. Zool. .Anz. XXIV, p. 461.
Transcaucasia.
10. ALLACTAGA ELATER DZUNGARIAE, Thomas
1912. .\nn. Mag. Nat. Hist. 8, IX, p. 406.
Zungaria, Central .Asia.
11. ALLACT.AGA ELATER INDICA, Gray
1842. Ann. Nat. Hist. X, p. 262.
Simkoh Hills, .Afghanistan.
SynonvTn: bactriana, Blyth, 1863, Cat. Mamm., p. no. Afghanistan.
12. ALLACTAG.A HOTSONI, Thomas
1920. Joum. Bombay Nat. Hist. Soc. XX\'I, p. 936.
Kant, Sib, Persian Baluchistan.
tcilliamsi Group
13. ALLACTAGA WILLIAMSI WILLIAMSI, Thomas
1897. Ann. Mag. Nat. Hist. 6, XX, p. 309.
Van, Kurdistan, .Asia Minor.
14. ALL.ACTAGA WILLIAMSI L.ATICEPS, Nehring
1903. Sitz. Ber. Ges. Naturf. Berlin, p. 357.
N.-W. Asia Minor.
15. ALLACTAGA WILLLAMSI SCHMIDTI, Satunin
1907. Mit. Kaukas Mus. 3, p. 239.
Caucasus, Kasimabad, Kr. Geokcai.
major Group
16. ALLACTAGA SEVERTZOVI, Vinogradov
1925. Proc. Zool. Soc. London, p. 583.
Tomar-Utkul, district of Kopal, Province Semiretchensk, Russian
Turkestan.
5S6 ALLACTACiA
17. ALLACTAGA MAJOR MAJOR. Kerr
1792. Anim. Kingd., p. 274.
Between Caspian Sea and River Irtish, Siberia.
Synonym: jaculus, I'allas, 1778, Nov. Spec, trlir. Ord., p. 87 (pre-
occupied).
auhcotis, Wagner, 1S43, .Abli. .Akad. Wiss. Miinclien III.
p. 211. (?).-\rabia.
itiacrotis, Brandt, 1844, Bull. Acad. Sci. St. Petersb., XI,
p. 220.
Jltn'csfens, Brandt, 1844, sanie reference.
nigricans, Brandt, 1844, same reference.
hrncliYotis, lirandt. 1844, Bull. .-Xcad. Imp. Sci. St. Petersb.,
II, p. 220.
For use of the name "major " Kerr, instead of "jaculus," auct., see
Chaworth-Musters, Ann. Mag. Nat. Hist. 10, XIV, p. 556, 1934.
18. ALLACTAGA M.AJOR SPICULLM, Lichtcnstein
1S25. .^bh. Akad. Wiss. Berlin, p. 154.
Barnaul, N.-W. Altai.
iq. ALLACTAGA ^L\JOR CHACLOVI, Martino
1930. .\nn. Mus. Zool. .Acad. Leningrad, 31, p. 209.
Karabulak .Saissan, Russian Turkestan.
20. ALLACTAG.'^ A1AJOR DECUMAN.^, Lithtenstcin
1S25. .Abhandl. Akad. Wiss. Berlin, p. 154.
Slatoust, Ural.
21. ALLACTAGA MAJOR FUSCU.S, Ognev
1924. Rodents N. Caucasus, Rostov-on-Don, p. 8.
Tischlovsk, Kizljar, Daghestan, Caucasus.
22. ALLACTAGA MAJOR VEXILLARLS, Evcrsmann
1S40. Bull. Nat. Moscow, p. 42.
No locality.
sihiricii Group
2j. ALLACTAGA SIBIRIC.A SIBIRICA. Forster
1778. Kongl. Vet. .Akad. Handl. XXXIX, p. 112.
Transbaikalia.
Synonym: saliens, .Shaw, 1790, Nat. Misc., vol. 2,p. i. Transbaikalia.
media, Kerr, 1792, Anim. Kingd., p. 274. Transbaikalia.
brachyurus, Blainville, 1S17, Nouv. Diet., XIII, p. 12b.
halticiis, Illiger, in Lichtenstein, .\bhandl. Wiss. Berlin,
1S25, p. 154; see Chaworth-Musters, .Ann. Mag. Nat.
Hist., 1934, 10, XIV, p. 556.
alactaga, Olivier, 1800, Bull. Soc. Philom., II, pi. iv, p. 121.
E'or use of the name " sihirica" instead of "saliens," auct., see
Chaworth-Musters, Ann. Mag. Nat. Hist. 10, XX, p. 96, 1937.
24. ALLACTAGA SIBIRICA AN.NLLATA, Milnc-Edwards
1867. .Ann. Sci. Nat. VII, p. 376.
Mongolia.
25. ALLACTAGA SIBIRICA SUSCHKIM, Satunin
1900. Zool. Anz. XXHI, p. 139.
Desert Ssasa Kopa, south of Irgis. Turgai, Kirghiz Steppe, S.-W.
Siberia.
ALLACTAGA— ALACTAGULUS 587
26. ALLACTAGA SIIilKICA MCJNGOLICA. Radde
1861. Mel. Biol. Acad. Sci. St. Pctersb., iii, p. 680.
North Gobi, Mongolia.
Synonym: (?) longior, Miller, 1911, Proc. Biol. Soc. Washington, XXIV,
p. 54. Fifteen miles north-east of Ching-ning-chow,
Kansu, China.
27. ALLACTAGA SIBIRICA UOcKI!i;n,l, Thomas
1914. .•\nn. Mag. Nat. Hist. 8, XIII, p. 571.
On banks of River Uszek, Djarkent, Semiretchensk, Central Asia.
28. ALLACT.'^GA .SIBIRICA SALT.VrOR, Eversmann
1848. Bull. Nat. Moscow, p. 188,
Tchuya Steppe, Altai.
29. ALLACTAGA SIBIRICA (?)GRISKSCENS, Hollister
1912. Smiths. Misc. Coll. LX, no. 14, p. 2.
Chuisaya Steppe, 8 miles south of Kosch Agatsch, Altai, Siberia.
Genus 2. ALACTAGULUS, Nehring
1897. Al.\ctagulus, Nehring, S.B. Ges. Nat. Berlin, p. 154.
Type Species. — Dipiis acontion, Pallas.
Range. — North Caucasus, Volgo-Ural Steppe, Kazakstan, south to Termez
region; Semirechyia ; and in Ordos desert, Mongolia.
Number of Forms. — Three or four.
Char.\cters. — Like AUactaga, but vertical branch of zygoma considerably
narrower than horizontal branch, and cheekteeth simpler,
with only two outer folds in ALi and M.2 in the upper series, the folds straighter
than in AUactaga; dentition generally appearing simpler, crowns flatter; in the
lower teeth M.i is similar to AUactaga, but M.2 lacks the small extra central
inner fold, having only two folds each side. Essential external characters as in
AUactaga. P. 4 absent.
Vinogradov states: " Alactagulus differs from AUactaga not only in the num-
ber of the cheekteeth and their structure, but also by some peculiarities of the
penis." The genus does not seem very distinct from Pygeretmus; on this point
Vinogradov states: " Pvgeretmus platviirus is very closely related to Alactagulus,
such resemblance exists also in the characters of the external genitals."
Forms seen: "acontion" {—pumiliu), dinniki.
List of Named Forms
I. AL.^CTAGULUS PUMILIO PLiMILIO, Kerr
1792. .\nim. Kingd., p. 275.
_ Betiveen Caspian Sea and River Irtish, Siberia.
Synonym: acontion, Pallas, 181 1, Zoograph. Rosso-.Asiatica, p. 182.
Kirghiz Steppes.
minor, Pallas, 1778, Nov. Spec. Quad. Glires, p. 295.
pygmaeus, Illiger, 181 1, Abh. ."Vkad. Berlin, p. 62. nom. nud.
minimis, Blainvillc. i8i7,Nouv. Diet. Nat. Hist. XI 1 1, p. 127.
For use of name piimilio in place of acontion, Pallas, auct., sec
Chaworth-Mustcrs, .Ann. Mag. Nat. Hist. 10, XIV, p. 556, 1934.
SSS ALACTAGULUS— PYGERETMUS
2. ALACTAGULUS PUMILIO DINNIKI, Satunin
1920. Trav. Mus. Georgie Tiflis, no. 2, p. ig6.
Prikumsk Steppe, Caueasus.
3. ALACTAGULUS PUMILIO POTANINI, Vinogradov
igaf). C.R. Acad. Leningrad, p. 233.
Ordos Desert, near Ulan Morin River, Mongolia.
In N'inogradov, 19:5 •^, List of Rodents of the U.S.S.R., there is quoted a race
Alactaguhis piiinilio piillithis. The reference to this has not been found.
Genus 3. PYGERETMUS, Gloger
1841. PYGERETMUS, Gloger, Gemeinn. Hand- u. Hilfsbuch d. Naturgesch., i, p. 106.
1S44. Platycercomys, Brandt, Bull, phys.-math. Acad. Sci. St. Petersb., ii, p. 225.
(Dipiis platyiirtis. Lichtenstein.)
Type Species. — Dipus platyurus, Lichtenstein.
Range. — S.-W. Siberia: Semirechia, and parts of valley of Ural, and
adjoining plain, and Kuvan-Daria (Aral region).
NrMBER OK Forms. — Two.
Ch.xR-ACTErs. — Like Aliict<i<;iiliis; margins of interorbital region more
angular; interorbital constriction more marked; cheekteeth f,
essentially similar to those of Alactai^ulus in two skulls available for examination.
Externally diflering from Alactoiiuhis and AUactaga in the structure of the
tail, which is relatively shorter, broad, flattened throughout its length, and
not tufted terminally.
Two species are known, evidently considerably distinct from each other;
Vinogradov gives measurements as follows:
platviinis : tail 80-90; hindfoot 32-35.
shitkovi: tail 95-107; hindfoot 40-43.
Forms seen: plat y urns, shitkoin.
List of Named Forms
platyurus Group
1. PYGERLTMUS PLATYURUS, Lichtenstein
1823. In Eversmann's Reise, p. 121.
Aral Sea region.
shitkovi Group
2. PYGERETMUS SHITKOVI, Kuznecov
1930. C.R. Acad. Sci. U.S.S.R., p. 623.
Kirghiz Steppes of Semipalatinsk, U.S.S.R.
The Dipus Group
Infraorbital foramen relatively smaller than in Allacttiga group, and anterior
vertical portion of zvgoma greatly broadened. Bullae larger; mastoids con-
siderably inflated, "their internal cavity consisting of two chambers separated
I'ARADIPUS 589
only liy one very low septum; this chamber is communicated with the cavity of
the tympanic bullae, as it can be observed even in Dipus, with its relatively
feebly inflated mastoids" (Vinogradov). Nasals reach alveoli of upper incisors.
Incisors not pro-odont, the upper ones usually grooved. The ears smaller than
in the Allactaaa group (possibly excepting Paradipus). Outer functionless digits
of hindfoot entirely suppressed. Os penis present (Vinogradov).
Key to the Genera oe the Dipus Group
(not including the genus Eremodipus, which is unrepresented in British Museum)
Palate terminating on level with hinder part of third molars. Apices of
bullae not in contact. Ear relatively larger. Upper incisors plain.
Mandible lacks process formed by root of lower incisor. Paradipus
Palate terminating behind level of third molars. Bullae with apices in
contact. Upper incisors grooved. Ear relatively smaller. Mandible
with process formed by root of lower incisor.
Mastoids not projecting on lateral sides of posterior part of braincase;
cheekteeth normally with more complex pattern. DiPus
Mastoids projecting on lateral sides of posterior part of braincase;
cheekteeth normally relatively simpler.
Mastoids not greatly inflated. Tail not heavily tufted, gradually
increasing in width from about halfway along its length to
the end. Squamosal with no ridge formed by lateral process
of parietal. Scirtopoda
Mastoids relatively enormously inflated. Tail heavily tufted
terminally, long; thin and round throughout most of its
length. Squamosal with ridge formed by lateral process of
parietal. ' Jaculus
Genus 4. PARADIPUS, Vinogradov
1930. Paradipus, Vinogradov, Bull. Acad. Sci. Leningrad, p. 333.
Type Species. — Scirtopoda ctcnodactyla, Vinogradov.
Range. — Described from Repetck, Turkmcnia, U.S.S.R.
Number of Forms. — ^One.
Characters. — Posterior edge of palate terminating on level with third
molars, instead of considerably behind them. Anterointernal
apices of bullae not in contact with each other. Mastoids rather inflated,
appearing in superior aspect of skull, hut not so large as in Jaculus. Postglenoid
fenestrae very small. .Mandible without process formed by root of lower
incisor. Cheekteeth ;j. Upper incisors plain.
One skull of this interesting Jerboa has recently been acquired by the
British Museum. It is evidently old; the cheekteeth appear to me to be quite
5<jo PARADIPUS— DIPUS
ditTerent from those of other Jerboas examined; their erowns are completely
flat, and with isolated enamel islands, these straight, surrounded by rather broad
enamel, two on M.i, two on M.z, one on M.3, upper and lower series.
Externally large; differing from other members of the Dipiis group in the
relatively large ears, "llindfoot with three long subequa! toes; imder surface
of lateral toes covered internallv with brusli consisting of long hairs and extern-
ally it is furnished with a comb consi.sting of a row of thickened horny bristles
about twice shorter than the long hairs." The ear is given as about 30 mm. by
\inogradov.
This genus, with its long ears, plain incisors, short palate, bullae with apices
not in contact, and, if constant, rather different appearance of worn cheekteeth,
(simpler than others), stands isolated in the Dipiis group.
Forms seen: ctetiodactvhis.
List of Named Forms
I. P.AR.'^DIPU.S CTENOn.^CTYLUS, Vinogradov
192Q. C.R. Acad. Sci. Leningrad, p. 248.
Rcpetek, Turkmenia, U..S.S.R.
Genus 5. DIPUS, Zimmermann
17S0. Dipus, Zimmermann, Geog. Geschichte Menschen und vierfiiss. Thiere, ii, p. 354.
I'lio. DiPODiPUS, Trouessart, Faune Mamm. Europe, p. 207. (Miis sagiltn, Pallas.)
Type Species. — Miis sagitta, Pallas.
R.^NGE. — U.S.S.R. and China; Kisljar district and North Caucasus; Volgo-
Ural steppe; Kazakstan to south Semipalatinsk; Altai; Semi-
rechie; Turkmenia, Usbekistan; Chinese Turkestan, Mongolia, to Shensi and
Chihli.
Number of Forms. — Eight.
Characters. — V,'\ke Juculus, to be subsequently described (Genus no. 7), in
cranial characters, except: mastoids much less inflated, less
so than in other members of the Dipus group; not appearing in superior aspect
of the skull. Postglenoid fenestrae "open into cavity of braincase"; "partly or
entirely closed by portion of petromastoideum in Scirtopoda, Jticiiliis, Para-
ilipiis." Squamosal region without the ridge characteristic of Jik 11 1 us.
Cheekteeth :',, semihypsodont. M.i in the upper series with a deep outer
told, placed far backwards, an anterior fold, and an inner fold. M.2 with two
outer, one inner fokls, the anterior outer fold normally, so far as seen, retained
(this fold becoming suppressed in allied genera). M.3 smaller than M.2, but
with the same elements originally. P. 4 minute.
The folds are deep, the cusps moderately high; four main cusps at corner
of each tooth.
Lower cheekteetii with two outer and two inner folds in M.i and M.2;
sometimes the folds nearly meet across the teeth; and M.3 with two outer, one
DIPUS— SCIRTOPODA S9i
inner folds. In all these teeth, the front outer fold is considerably smaller than
the second one, which is more persistent. Some of the folds wear out in old
age. Essential external characters as Jaculus.
All described forms are evidently regarded now as races of the type, by
\'inogradov.
Forms seen: halli, lagopus, sowerbyi, deaysi.
List of Named Forms
1. DIPUS SAGITTA SAGITTA, Pallas
>773- Reisc, ii, p. 706.
Siberia.
2. DIPUS SAGITTA NOGAI, Sauinin
igoy. Tiflis Mitt. Kaukas. Mus. 3, p. 34.
N.-E. Caucasus.
3. DIPUS SAGITTA INNAE, Ognev
1930. Zool. Anzeiger, 91, p. 207.
Astrachan Gouv., S.-E. Russia.
4. DIPUS SAGITTA LAGOPUS, Lichtenstein
1823. In Eversmann's Reise, p. 121.
Transcaspia.
5. DIPUS SAGITTA ZAISS.^^NENSIS, Selewin
1934. Bull. Univ. Tachkent, ig, p. 76.
Saissan-nor, Central Asia.
6. DIPUS SAGITTA DEASYI, Barrett-Hamilton
1900. Proc. Zool. Soc. London, p. 196.
Nura, S. Chinese Turkestan.
7. DIPUS SAGITTA HALLI, Sowerby
1920. Ann. Mag. Nat. Hist. 9, V, p. 279.
N.-E. Chihli, N. China.
S. DIPUS SAGITTA SOWERBYI, Thomas
igoS. Ann. Mag. Nat. Hist. 8, II, p. 307.
Yu-Iin-fu, Shensi, China.
Genus 6. SCIRTOPODA, Brandt
1844. SciRTOPODA, Brandt, Bull, phys.-math. Acad. Sci. St. Petersbourg, ii, p. 212.
1844. H.ALTicis, Brandt, Bull, phys.-math. Acad. Sci. St. Petersbourg, ii, p. 213.
(Dipus halticus, Illiger.)
1925. Stylodipus, Allen, Anier. Mus. Nov., no. 161, p. 4. Stylodipus aiidrezisi. .Allen:
not seen ; status fide Vinogradov.
Type Species. — According to Vinogradov the type is now taken to be
Dipus teltini, Lichtenstein.
R.\nge. — Russia (Crimea, Ciscaucasia, Lower Volga, Kazakstan east to
Saissan, Aral Sea and Lake Balkash regions, Semirechie, Kara-
kum) (Vinogradov); also known from Mongolia.
38 — Living Rodents — I
502 SCIRTOPODA
Number of Forms. — Seven.
Ch.^r.'vcters. — Like Jaciiliis (next to be described) in cranial characters, but
squamosal with no ridge formed by re-entrant lateral process
of parietal, and mastoids much less inflated, though in this genus they are more
advanced than in Dipiis in that they show in the superior aspect of skull each
side of the braincase. Cheekteeth normally a; * in the type of " Stylodipus"
andreusi; but according to Vinogradov a minute upper premolar may be present
in the young, but disappearing with age, in the other species. The pattern, as
far as seen, not essentially difl'erent from Jaciiliis, but the teeth appearing rather
flatter, less angular, and simpler than in Dipiis. Upper incisors, as normal for
this section, grooved.
Externally like Jaculus except that the tail is relatively shorter, with the tuft
weaker, less terminal, less developed, the tail gradually becoming wider from
about halfway up its length; the terminal portion not black and white. The
tail in fact seems to be somewhat intermediate between the normal AHactaga or
Jaculus type, and the Pygeretmus type.
S. andrewsi, the type of Allen's genus Stylodipus. is regarded as a Sciriupodu
by Vinogradov. It is not represented at the British Museum; but it appears
from the figures published that it is a closely allied form to 5. telum, and rightly
placed in this genus.
Forms seen : leluiii, proximus.
List of Named Forms
I. SCIRTOPOD.^ TELUM TKLUM, Lichttnstcin
1823. In Eversmann's Reise, p. 120.
Aral Sea region.
;. SCIRTOPODA TELUM FALZFEIXI, Ognev
1916. Bull. Soc. Nat. Crimee, 5, p. loi.
Taurida district, Crimea, S. Russia.
3. SCIRTOPODA TELUM TUROVI, Heptncr
1Q34. Folia Zool. Hydrob. 6, p. 19.
Don Steppe, S.-E. Russia.
4. SCIRTOPODA TELUM KARELINI, Sclcwin
1934. Bull. Univ. Tachkent, 19, pp. 76-78.
Kazakstan, Russia. Mountains of Semej-Tau, near Semipalatinsk.
5. SCIRTOPODA TELUM AMANKARAGAI, Sekuin
1934. Bull. Univ. Tachkent, 19, p. 76.
Kazakstan, Russia: Aman-Karagai, N. Kazakstan.
'.. SCIRTOPODA TELUM PROXIMUS. Fairmaire
1853. Rev. et. Mag. Zool., p. 145.
Janiankala, Ural.
7. SCIRTOPODA ANDREWSI, Allen
1925. Amer. Mus. Nov. no. 161, p. 4.
Ussuk, Mongolia.
According to Vinogradov, S. andrnosi retains the vestigial premolar in adult
specimens; the hindfoot is larger (about 55) than telum (46-51).
JACULUS 593
Genus 7. JACULUS, Erxleben
>777- Jaculus, Erxleben, Syst. Regn. Anim., p. 404.
1844. Haltomys, Brandt, Bull, phys.-math. Acad. .Sci. St. Petcrsb., ii, p. 215. {Dipus
aegyptiiis, Hasselquist).
(1922. Scirtcipoda, Pocock, based on ^ orientalis; not as now accepted. (For note on
the type of the genus Scirtopoda see Vinogradov, Bull. Acad. Sci. L'URSS, iy3o,
P- 332-))
Type Species. — Jaculus orientalis, Erxleben (see St. Leger, Proc. Zool. Soc.
London, 193 1 : Genera of African Rodentia).
Range.- — Northern Africa, Senegambia, Morocco, across the Sahara to
Egypt and Somaiiland; extending into Arabia, Palestine, Syria,
Iraq and Persia.
Number of Forms. — About twenty are named.
Characters. — Skull with extremely broad frontals, and even broader
braincase, becoming gradually narrower from behind
forwards; rostrum narrow; lachrymal large. Jugal as usual in the group, with
anterior vertical portion much broadened. Superior margin of canal in infra-
orbital foramen for nerve-transmission fused to wall of maxilla in adult. Bullae
greatly inflated, mastoids appearing prominently in superior aspect of skull,
much more than in other 3-toed Jerboas examined. Squamosal region with
well-marked downwardly directed ridge formed by lateral process of parietal.
Supraorbital region rather angular. Palate extending behind level of i\L3 ;
palatal foramina well open, relatively large, and a second pair present as pits
between the toothrows. Mandible with perforation in the angular process, and
a process formed by the lower incisor root.
Upper incisors one-grooved. Cheekteeth 5, the upper teeth with one wide
inner and one wide outer re-entrant fold which is placed further backwards
than the inner one; M.i also with anterior notch which tends to wear out;
cusps lower than usual as a rule. Al.i the largest tooth, M.3 the smallest.
Lower cheekteeth with two outer folds in M.2, and one inner one; one outer
fold persistent in M.3 only; M.i with a fold each side as a rule, and an anterior
notch.
Fur soft. Ear large, but considerably smaller than in members of the
Allactaga group. Tail considerably longer than head and body, normal in
shape (narrow, round, not thick, flat), and well-haired throughout, with a
conspicuous black and white tuft terminally. Forefoot with five digits, D.5
moderate but shorter than the central three, the pollex less reduced than is
usual among non-tossorial Rodents; claws thin but strong. Hindfoot immenselv
elongated, very narrow, with three digits; soles heavilv hairv.
The forelimbs are proportionately extremely shortened, and can only be
seen in the living animal on the rare occasions when the animal is still. Jerboas
of this type (J. jaculus) are very fond of scratching and rolling in their sand;
they have a curious habit of lying down and stretching the hindleg as far back-
wards as it will go, then bringing it round and stretching it forwards, so quickly
Fig. 156. Jacilus jaculus jaculus, Linnaeus.
B.M. No. .S.4.4.52. .. ; • ai.
Fig. 157. J.ACULUS jaculus jaculus, Linnaeus.
B.M. No. 8.4.4.52, V; • 2*.
JACULUS
S9S
that the eye can scarcely follow the procedure ; it is at such times that one gets
a good idea of the elongation of the limb. Their leaping powers are prodigious,
and they often walk along on their hindlimbs; they appear quite unable to go
on all fours. The eye is very large.
According to Lyon, the axis and four succeeding vertebrae are completely
fused into one large compound "axis." On the skeleton of the hindfoot all
traces of the outer digits have disappeared. The femur is short, the tibia and
cannonbone long. The upper caudal vertebrae are thick and powerful.
Fig. 158. Jacllus jaculus jaculus, Linnaeus.
Cheekteeth: B.M. No. 8.4.4.52, $; X 11.
The genus name Scirtopoda was revived bv Pocock for the Greater Egv'ptian
Jerboa, J. orienlalis, on the grounds that the structure of the penis differs from
that of the jaculus group. \ inogradov does not retain the division, and shows
that Scirtopoda is not available for the species (Thomas having previously and
apparently erroneously chosen the type of Scirtopoda for a different animal
from S. telum which Mnogradov states should be considered the type).
As I have endeavoured to show, when dealing with Sciuridae, names based
solely on the structure of the baculum are not to he considered valid as full genera.
Two clearly marked groups occur, the smaller jaculus (hindfoot about 60,
only one in British Museum (blaufordi, Persia) e-xceeding 63), and the orientalis
group, differing, as indicated above, in baculum structure, and larger size
(hindfoot 70-81, exceeding any member of jaculus group, so far as seen).
All members of the jaculus group arc regarded as of one species except
blanfordi, which appears to be rather larger than the others. I think that many
of the races of jaculus will ultimately have to be placed in synonymy.
Forms seen : airensis, blanfordi, butleri, centralis, deserti, faionicus, florentiae,
gordoni, jaculus, loftusi, orientalis, schlueteri, sefrius, syrius, locator, tulturnus.
5y6 JACULUS
List of Named Forms
incertae sedis
1. JACULUS MICROTIS, Ri-ichenow
18S7. Zool. Anz. X, p. 369.
Samar. N.-E. Africa.
2. JACULUS MACROTARSUS, Wagner
1S40. Abh. Akad. Wiss. Munchen, III, p. 214.
Arabia Petraea, Mount Sinai.
uricntalts Group
3. J.^CULUS ORIENTALIS ORIENTALIS, Er.xleben
1777- Syst. Regn. Anim., p. 404.
Egypt.
Synonym: gerboa, Olivier, Bull. Soc. Philom., i, 2, No. 40, p. 121,
iSoo. Egypt.
locusta, Illiger, 1804-1811, Abh. Akad. Berlin, p. 77.
Egypt.
bipes, Lichtenstein, 1823, Verz. Doublet. Mus. Berlin, p. 5.
Egypt.
('>)acgyptiiis, Hasselquist, 1744, Acta. Soc. R. Sci. Upsala,
p. 17. Egypt.
4. JACULUS ORIENTALIS MAURITANICUS, Duvernoy
1842. Mem. Soc. Hist. Nat. Strasb., iii, p. 30.
Oran, Algeria.
jaculus Group
5. JACULUS JACULUS JACULUS, Linnaeus
1758. Syst. Nat., loth Ed., p. 63.
N. Egypt.
Svnonym: (})hirtipes, Lichtenstein, 1823, Verz. Doubl. Mus. Berlin,
p. 5. Near Assuan, Upper Egypt.
6. JACULUS JACULUS BUTLERI, Thomas
1922. Ann. Mag. Nat. Hist. 9, IX, p. 296.
Khartoum.
7. JACULUS JACULUS GORDONI, Thomas
1903. Proc. Zool. Soc. London, i, p. 299.
Kaga Hills, W. Kordofan, Sudan.
8. JACULLS JACULUS VULTURNUS, Thomas
191 3. Ann. Mag. Nat. Hist. S, XI, p. 485.
Berbera, Somaliland.
.). JACL'LUS JACULUS AIRENSIS, Thomas & Hinton
1921. Nov. Zool. XXVIII, p. 10.
Aderbissinat, north of Damergou, Sudan.
10. JACULUS J.\CULUS CENTRALIS, Thomas S; Hinton
1921. Nov. Zool. XXVIII, p. II.
Oued el .Abiad, In-Salah, Air, .Sahara.
JACULUS— EREMODIPUS 597
11. JACULUS JACULUS DESERTI, Loche
1867. Explor. Alger., p. 100.
Ouargla district, N. Algerian Sahara.
Synonym: darricarrerei, Lataste, 1883, Ann. Mus. Civ. Genova, XVIII,
p. 661. Bou-Saada, .Algeria.
12. JACULUS JACULUS SEFRIUS, Thomas & Hinton
1921. Nov. Zool. XXVIII, p. 10.
Ain-Sefra, Algeria.
13. JACULUS JACULUS F.'WONICUS, Thomas
1913. Ann. Mag. Nat. Hist. 8, XI, p. 483.
Trarza country, S.-W. Mauritania.
14. JACULUS JACULUS SCHLUETERI, Nehring
1901. S.B. Ges. Nat. Fr. Berlin, p. 163.
Palestine.
15. JACULUS JACULUS SYRIUS, Thomas
1922. Ann. Mag. Nat. Hist. 9, IX, p. 296.
Karyatein, Syrian Desert.
16. JACULUS JACULUS FLORENTIAE, Chetsman & Hinton
1924. Ann. Mag. Nat. Hist. 9, XIV, p. 556.
Jabal Aqula, Jabrin, Central Arabia.
17. JACULUS JACULUS ORALIS, Cheesman & Hinton
1924. Ann. Mag. Nat. Hist. 9, XIV, p. 557.
Koweit, N.-E. Arabia.
18. JACULUS JACULUS VOCATOR, Thomas
1921. Ann. Mag. Nat. Hist. 9, VIII, p. 441.
Sohar, Muscat, Arabia.
iq. JACULUS JACULUS LOFTUSI, Blanford
1875. Ann. Mag. Nat. Hist. 4, XVI, p. 312.
Mohumrah, Mesopotamia.
20. JACULUS BLANFORDI, Murray
1884. Ann. Mag. Nat. Hist. 5, XIV, p. 98.
Bushire, Persia.
Genus 8. EREMODIPUS, Vinogradov
1930. Hull. .Acad. Sci. Leningrad, p. 334.
Type Species.- — Scirtopoda Uchtensteini, Vinogradov.
Range. — Vicinity of Merv, Turkmenia.
Number of Forms. — One.
Rem.arks. — This genus is not represented at the British Museum. It
evidently stands nearest Jactilus.
Vinogradov keys this genus against Jiiciiliis as follows:
" Lateral process of each parietal bone is furnished with a sharply developed
conical prong directed externally and downwards. Postglcnoid
jqS eremodipus
fencstrae arc very small, usually somewhat elongated. Superior
margin of external wall of infraorbital canal not ankylosed to wall
of maxilla. Root of lower incisor forms a feebly developed alveolar '
process. Eremodipus
Lateral process of each parietal bone is furnished with a sharply developed
crista, its base being formed by surrounding parts ot the squa-
mosal. Postglenoid fenestrae are greatly enlarged and form nearly
equilateral triangles. Superior margin of external wall ot intra-
orbital canal is completely fused to wall of maxilla in adult speci-
mens. Root of lower incisor forms a prominent alveolar process.
J.\CULUS "
List of N.\med Forms
I. EREMODIPUS LICHTENSTEINI, Vinogradov
1927. Zeitschr. f. Siiugetierk. 2, p. 92.
Vicinity of Merv, Turlvnienia.
The family is known fossil from the Pleistocene at least, from both the Old
and the New Worlds.
Miller & Gidley refer the European Eocene-Miocene family Theridomyidae
to this group; Winge places them in the neighbourhood of the Anomaluridae.
DIPODIDAE:
GEXERAL WORKS OF REFERENCE
Vinogradov, 1925, Proc. Zool. Soc. London, p. 577. Structure of external genitalia of
Zapodidae and Dipodidae.
Vinogradov. Bull. Acad. Sci. LeninRrad 1930. p. 331. Cranial Characters of genera of
Family Dipodidae.
Lyon, Proc. U.S. Nat. Mus. 7, XXIII, p. 659, igoi. Comparison of the osteology of
Jerboas and Jumping-mice.
Vinogradov, 1933, Tab. Anal, de la Fauna de L'URSS, Inst. Zoot.Acad. Sci. 10. p. 11.
Rodents occurring in the LT.S.S.R. (Sicistinae, Dipodmae.)
Vinogradov, 1923, Kozlow Mongolia & Amdo, p. 540. (Salpmgotus kozloii.)
Vinogradov, 1928, Zool. Anz. 61, p. 150. (Salpiiigotus crassicauda.)
Preble, North Amer. Fauna, no. 15. p. 13, iSgg. Revision of Jumping-mice (Zapus,
Eozapiis, Napcicozapiis).
COUES, Monogr. North .American Rodentia, p. 461, 1S77. Zapodidae.
Miller, Cat. Mamm. W. Europe, 1912, p. 535. Zapodidae (Sicista).
PococK, External characters of Scarlurus and other Jerboas, compared with Zapus and
Pedetcs, Proc. Zool. Soc. London, 1922, p. 659.
SCL.ATER, Proc. Zool. Soc. London, 1890, p. 610. (Euchoreutes.)
DoBSON, Proc. Zool. Soc. London, 1882, p. 640. (Dipodidae transferred to the "Hystri-
comorpha.")
Tlllberg, Nova Acta Reg. Soc. Sci. Upsalicnsis, XVIII, 3, no. i, 1899. (Sicista,
Zapus. AUactaga, " Dipus aegyptius" =Jaculus.)
Ch.aworth-Mtsters, .Ann. Mag. Nat. Hist. 10, XIV, p. 556, 1934: Nomenclature of
Alactagulus and certain species oi AUaclaga; .\nn. Mag. Nat. Hist. 10. XX, p. 96,
1937: Nomenclature of AUactagn sihirica; .Ann. Mag. Nat. Hist. 10, XIV, p. 554,
1934: Nomenclature of Sicista suhtilis group.
MUROIDAE 599
Vinogradov, 1937, Inst. Zool. Acad. Sci. de L'URSS, no. 13, vol. II, no. 4. Family
Dipodidae. (A Monograph of the Family, with figures of cranial, dental, skeletal
and reproductive characters of all leading species, and distribution maps ; Russian
with English resume.)
Superfamily MUROIDAE
1896. Thomas: Myomorph.a, part. Families Gliridae (Glirinae and Platacanthomyinae);
Muridae (Hydromyinae. Rhynchomyinae, Phloeomyinae, Gerbillinae, Otomyinae,
Dendromyinae, Murinac, Lophiomyinae, Sigmodontinae, Neotominae, Microtinae,
"Siphneinae" ( =Myospalacinae)) ; Spalacidae (Rhizomyinae, Spalacinae).
1899. Tullberg: Scil'ROgnathi, Myomorpha, Myoidei, part, Myoxiformes (Myoxidae =
Muscardinidae), and Muriformes; Spalacidae, Nesomyinae, Cricetidae, Lophiomyi-
dae, Ar\icolidae, Hesperomyidae, Muridae (Murini, Phloeomyini, Otomyini),
Gcrbillidae.
1918. Miller & Gidley: Superfamily MuROlD.\E, part. Families Muscardinidae; Cricet-
idae (Cricetinae, Gerbillinae, Microtinae, Lophiomyinae) ; Platacanthomyidae ;
Rhizomyidae (Tachyor>ctinae, Rhizomyinae); .Spalacidae (Myospalacinae, Spalac-
inae); Muridae (Dendromyinae, Murinae, Phloeomyinae, Otomyinae, Hydromy-
inae). Superfamily Dipodoidae, part. Family Graphiuridae.
1924. Winge: Family Myoxidae ( =Muscardinidae) ; (Graphiurini, Myoxini). Family
Muridae (Rhizomyini, Cricetini, Murini). Family Dipodidae, part, Spalacini
(Spalax only).
1928. Weber: Myoxoidea, Myoxidae (=Muscardinidae) and Platacanthomyidae.
MuROlDEA, Spalacidae, Nesomyidae, Muridae (Cricetinae, Lophiomyinae, Microt-
inae, Murinae, Gerbillinae, Hydromyinae).
Geographical Distribution. — Cosmopolitan, including Madagascar and
the Australasian region.
Characters. — I have already written at some length on the characters of
the superfamily Muroidae as here understood, on pp. 35, 36.
The infraorbital foramen always transmits muscle, but is never vers- much
enlarged, at any rate as compared with Dipodoid, Anomaluroid, Pedetoid
or Hystricoid types; except in the two genera Graphiurus and Deomys the
zygomatic plate is tilted upwards to a greater or lesser degree. The fibula is, so
far as known, always fused w ith the tibia high on the leg. In the whole of the
family Muridae, containing well over half the entire Order, there are, except
in abnormalities, never more than I cheekteeth present.
In this group I include the Muscardinidae, which, though typically very
distinct from ^Iuridae, contains annectant forms such as Typhlomvs which
make it not possible to keep them separate as a distinct superfamily; the Muri-
dae, and a few highly specialized or aberrant genera which it has seemed desir-
able to make types of distinct families, as Rhisomys, Spalax, and Lophiomys.
Key to the F.\.milies of Muroid.^e
Upper and lower cheekteeth with a pattern of many transverse crossridges
extending across crown ; in primitive forms more or less basin-
shaped, as in Sciuridae, and with well-marked corner cusps, in
progressive forms becoming flatcrowned, with the ridges separated
boo MUROIDAE— MUSCARDINIDAE
by depressions. Premolars usually, not always, present. Caecum
usually, not always, absent. Jugal bone usually relatively long.
Family Muscardinid.^e
Upper and lower cheekteeth various, but never with pattern as just
described. Premolars invariably absent (or cheekteeth formula not
exceeding fj, e.xcept in abnormalities). Caecum, so far as known,
present. Jugal bone usually, not always, strongly shortened.
Temporal fossae roofed in by bony plates rising from jugals, frontals,
and parietals. Family Lopiiiomyid.\e
Temporal fossae never roofed in by bony plates.
Infraorbital foramen much reduced, its lower border nearly
straight; zygomatic plate tilted very strongly upwards, and
masseter muscle extending line of attachment on inside of
infraorbital foramen (TuUberg). Family Rhizomyid.^e
Infraorbital foramen not much reduced, its lower border usually
V-shaped; zygomatic plate tilted upwards less strongly;
massetei muscle so far as known never extending line of
attachment on inside of infraorbital foramen.
External form and skull extremely specialized for underground
life; eyes suppressed; zygomatic plate much narrowed,
and nearly completely below infraorbital foramen.
Family Sp.'vl.^cid.^e
External form and skull less extremely specialized for under-
ground life ; eyes always retained ; in sub-fossorial genera,
zygomatic plate not narrowed, and well tilted upwards.
Family Murid.\e
Family MUSCARDINIDAE
For use of the family name "Muscardinidae" instead of "Gliridae" see
Palmer, Science, n.s., vol. X, no. 247, p. 412, 1S99.
1896. Thomas: Mvomorpha, Family (ilindae: .Subfamily Glirinae (including
Graphiiiriis) ; Subfamily Platacanthomyinae.
1899. Tullbcrg: Mvomorpha: Myoxiforincs. Family Myoxidae.
igi8. Milltr &: Clidlcy: Superfaniily MuRomAE, part: F'aniily Muscardinidae (Elioinys,
Dvromys, GIis. Miisaudinus); Family Platacanthiimyidae (Piatacantliomys, Typh-
lomys). .Supcrfamily Dn>ODom.^F,, part: Family Graphiuridac {Gniphitinis).
1924. VVinge: Family Myoxidae. Subfamilies Graphiurini .ind Myoxini (the latter
including Platacanlhomys).
1928. Weber; MvoxomEA. Family Myoxidae (including Graphiurus); Family Flata-
canthomyidae.
Geogr.xphical Distribution. — Africa; Palaearctic region; parts of the
Indo-Malayan region (Peninsular India and
South China).
Number of Gener-A. — Nine.
MUSCARDINIDAE 6oi
Characters. — Zygomasseteric structure in progressive genera (all but
Graphiurinac), approaching or agreeing with that of the
iMuriilae; infraorbital foramen transmitting muscle, though little enlarged, and
comparatively unspecialized; zygomatic plate tilted upwards to a certain degree;
mandible with angular portion usually pulled inwards, and sometimes with a
perforation. In (iraphiurinae, the zygomatic plate remains beneath the small
infraorbital foramen, and the masseter muscle does not extend attachment on
its forepart ; masseter lateralis superficialis has its anterior head not distinct
(according to Miller & Gidley and as figured by TuUberg), whereas in Mus-
cardininae and Platacanthomyinae this portion of the muscle, as in Muridae, is
distinct from the zygoma. The jugal is generally long.
Dental formula i. {, c. ^, p. j, m. :| =20 in Muscardininae and Graphiurinae;
or i. }, c. H, p. H> Ti- t = i6, in Platacanthomyinae. Cheekteeth brachyodont,
always with pattern of a series of ridges extending across the crown. In more
primitive genera, as Graphiuriis, F.Uom\s, the cusps are well marked, and the
pattern and arrangement ot cusps and ridges is strongly reminiscent of that of
the Sciuridae. In progressive types, Glis, and to a greater degree Muscardinus,
the crowns become nearly flat, with obsolete cusps, and well-marked narrow
ridges surrounding moderately wide depressions. In Platacanthomyinae, which
are slightly more hypsodont, the ridges become widened, and the depressions
take on a more clear and definite pattern, and sometimes tend to isolate as islands.
Normally the bullae are large and well inflated, but are small in Plata-
canthomyinae, and flattened and rather reduced in Glirulus.
'I'he external form is slightly modified as a rule for arboreal life. The tail
normally is bushy.
The caecum most often is suppressed ; but this is not the case in Tvphlomys.
The systematic position of these animals is by no means clear, and has been
one of the major problems of the present classification of the Order.
Winge recognized only eight families of Rodents as here understood, this
group one of them (distinct from Muridae).
Weber has regarded the group as a superfamily.
Tullberg evidently regards the group as a natural one, separate from the
Muridae (in his Muroidei he has three equal groups, the "Myoxiformes"
( = Muscardinidae), Dipodiformes ( = Dipodidae), and Muriformes (Muridae,
Spalacidae, Lophiomyidae and Rhizomyidae as here understood).
-Miller & (iidley refer Graphiurus to a separate superfamily, the Dipodoidae,
from other .Muscardinidae, which are placed in the Muroidae. This arrange-
ment is based entirely on zygoma.sseteric structure, but is in my opinion rather
an unnatural division, in that Graphiurus seems to share very many essential
characters with Eliomys (Muscardininae). I'laiacanthomys and Typhlornys these
authors refer to a family Platacanthomyidae in the "quadrituberculate series"
of Muroidae; whereas their .Muscardinidae are referred to the "trituberculate
series" of Muroidae.
If wc take Graphiurus, and compare it with Eliomys, as regards arrangement
of zygomatic plate and infraorbital foramen, it is noticeable that if the infra-
orbital foramen of, say, Graphiurus hueti were slightly narrowed below and
6o2 MUSCARDINIDAE
considerably above, the result would be as is now in Eliomys. On looking
through the skulls at the British Museum I was struck by the fact that there
seems a slight variation in form in the infraorbital foramen of various specimens
of Graphiurus. In G. surdiis, for instance, it is not far from the primitive Muroid
type as characterizes Miiscardhiiis, Glis, Eliomys and others.
In the Muridae, the African genus Deomvs would certainly have to be
referred to the "Dipodoidae" of Miller & Gidley it their classification were
followed to the letter.
Although Miller & (jidley were not of opinion that any of their super-
families were derived one from another, I am inclined to suspect that in the
present case, one zygomasseteric structure, say that of Muscardininae, has been
derived from the other as typified by Graphiurus. In zygomasseteric structure,
as in many other characters, the present group seems to be one of the most
primitive groups of Rodents, not very far removed, at any rate as regards
arrangement of infraorbital foramen, from the type of Rodent {? Aplodontoid)
that probablv gave rise to most or all of the modern families.
We have now to consider whether the Muscardinidae (all Dormice being
referred to one familv) are distinguishable from the Muroidae as a superfamiiy.
If we take Dyroinvs and compare it with, say, Graiinnoiiixs representing the
Muridae, we find the following differences.
The cheekteeth in Dvromxs are basin-shaped, the upper and lower series
being characterized by many narrow transverse ridges; in Grammomvs the
molars are cuspidate, the upper series bearing three longitudinal rows, the
lower molars two longitudinal rows.
The fourth premolar is present in Dvromxs; in Gramiiionixs it is absent (or
at any rate only ij cheekteeth are present in the latter). The mandible of
Dxrotiixs has the angular portion pulled inwards, after the manner of Dipodidae,
Sciuridae, Aplodontiidae, etc. In Grainmoinxs this is not the case. The jugal
in Dxromxs is long; whereas in Gnimiuomxs it is becoming shortened; in very
many other Muridae it is strongly shortened. In Dyroinvs the caecum is sup-
pressed; in Graniiiiomxs, presumablv, this is not the case. The tail is thickly
bushy in Dxromxs; mostly naked and scaly in Grainmoinxs. And in Dxromxs the
zvgomatic plate is relatively weak and narrow, in Grainmoinxs it is broader and
stronglv tilted upwards, as is often the case in Muridae. The bullae are large,
inflated in Dxromys, rather small in Grammomys.
Between these two therefore there are clear distinctions. But there are
intermediate genera which appear to break down all these characters. In
Platacantlwmxs (Muscardinidae), the premolars are suppressed, and the dental
formula is as in Muridae. The mandible in Txphlomxs (Muscardinidae) has
no perforation, and is reduced, and not noticeably inflected. The jugal in
Tachxorxctes, Braclixuroinxs and others (Muridae), is long, forming the greater
part of the zygoma. The caecum is not suppressed in Typhloinys (Muscardin-
idae;), but becoming very reduced, according to Thomas, in Ichthyomys
(Muridae). The tail is nearly naked in Txphlomxs (Muscardinidae), thickly
bushy in Cratcromys (Muridae). The zygomatic plate is very narrow in
Hvdromys (Muridae), very much as in Platacanthoinys representing the
MUSCARDIMDAE; GRAPHIURINAE 603
Muscardinidae. The bullae are small in Glirulus (Muscardinidae); very
large in many Gerbillinae (Muridae). The cheekteeth alone remain. I can
call to mind no members of the vast group referred to Muridae which bear
any close resemblance to Muscardinidae. Perhaps Gymnuromys of Madagascar
stands nearest Platacanthomys in this respect. But pattern of cheekteeth seems
scarcely a valid character on which to base superfamilies. Compare, for
instance, the teeth of Kattus, Cricettis, Alicrotus, Otomys, Sigmodon. All
appear widely distinct in pattern; yet all belong to the one family. Compare
Ctenodactvlus with Ctenomys (essentially similar but belonging to different
superfamilies); or Phloeomys (Muridae) with /)//)/o/H3i (Hystricoidae), which are
also similar in general arrangement.
This being the case the Muscardinidae are regarded provisionally as primi-
tive and aberrant members of the superfamily Muroidae.
Three subfamilies are here retained.
Key to the Subf.a.milies of Muscardinidae
Zygomatic plate very narrow, completely beneath infraorbital foramen.
Subfamily GR.ypHiURiNAE
{Graphiurus)
Zygomatic plate broadened to a certain degree, always tilted upwards.
4
Cheekteeth -, with transverse ridges on crown moderately or well
developed, always narrow, the depressions not tending to
become isolated on crown surface, and not clearly marked as a
rule. Palate without a series of foramina between the tooth-
rows. Bullae usually large, well inflated. Subfamily Ml'sc.ardinin.ie
{Mxomimus (not seen), Eliomys, Dyromys, Glirulus, Glis,
Muscardinus)
Cheekteeth-, with transverse ridges clearly marked, broadened, and
3
the depressions tending to become isolated on crown surface,
always well marked. Palate with a series of foramina or a single
very large pair between the toothrows. Bullae small, reduced.
Subfamily Platac.anthomyixae
{Platacanthomys, Typhlomys)
Subfamily GRAPHIURIN.\E
Geographical Distribution. — .-Vfrica, south of the Sahara.
Number of Genera. — One.
CHAR.ACTERS. — As indicated in the key. Cheekteeth *, basin-shaped, the
ridges weak, the pattern as a rule not clear.
Though currently referred to three or four genera, it seems most convenient
to regard all members of the present subfamily as belonging to one genus only.
6o4 GRAPHIURUS
Genus i. GR.\PHIURUS, Smuts
1S32. Graphuris, Smuts. Enum. Mamm. Cap., pp. 32, 33.
1936. Aethoglis, Allen. Journ. Mamm. 17, p. 292. Graphiurus nagtglasi, Jentink.
1888. Cl.wiglis, Jentink, Notes Leyden Mus., p. 41. Claviglis crassicaudatus,
Jentink. Valid as a subgenus.
1925. Glirisci's, Thomas & Hinton, Proc. Zool. Soc. London, p. 232. Graphiurus
platyops, Thomas. Valid as a subgenus.
Type Species. — Sciurus ocularis, Smith. {Grapliiuriis capensis. Smuts.)
Range. — Africa: Sudan, Sahara, Abyssinia, SomaHhind, Kenya, Uganda,
Tanganyika; Senegal, Liberia, Gold Coast, Nigeria, Cameroons,
Congo; Angola, Rhodesia, Nyasaland, Mozambique, South-west Africa,
Bechuanaland, Transvaal, Cape.
Number of FomMS. — About fifty-three.
CH.AR.ACTERS. — Zygomatic plate not tilted upwards and completely beneath
the infraorbital foramen. Skull considerably constricted
between the frontals; braincase smooth and round; nasals projecting forwards.
Palate relatively broad; palatal foramina situated considerably in front of
toothrows. Bullae large and inflated as a rule; mandible normally without
perforation in the angular process.
Crowns of cheekteeth concave, with two low main external cusps; the
general arrangement in pattern evidently near Elioinvs (below), but the ridges
in most indistinct, and a general tendency towards simplification. The pre-
molars are usually only moderately reduced ; but in the type species are strongly
reduced. G. rupicola has a rather reduced lower P. 4.
Externally with thickly bushy tail (normally); hindfoot with five digits, the
fifth relatively long, the hallux short; the feet are of arboreal type. Caecum
(said to be) absent.
Thomas & Hinton divided the genus into three main groups, which they
keyed as follows :
Premolar minute, simple; surface of teeth with scarcely perceptible
ridges. Graphiurus
Premolar little smaller than the molars, its outer edge, when unworn,
notched as in the molars; surface of teetii with distinct ridges.
Skull of normal height, braincase strongly convex upwards. Claviglis
Skull flattened, muzzle low, braincase scarcely convex upwards. Gliriscus
In the first case, it must be pointed out that Gliriscus cannot be used in a
generic sense if only on the grounds of consistency. Hinton (Monograph of
N'oles and Lemmings, 1926, p. 44) writes, of the genus Alticola, "some remark-
able species inhabiting the bare talus slopes of Central Asia, have acquired
remarkably flattened skulls fitting them for life in rock crevices; these have
been referred to a special subgenus Platycranius by Kascenko, but apart from
the peculiar flattening of the skull there is nothing to distinguish them from
Fig. 159. Gr-^phiurus hueti hueti, Rochebrunc.
B.M. No. 25.10.24.1; X 2j.
Fig. 160. Graphiurus hueti hlieti, Rochebrune.
B.M. No. 25.10.24.1; X 2J.
6o6 GRAPHIURUS
the more specialized forms ot Alliculii" ; and (on p. 325), "'l"he subgenus Phitv-
craitiiis seems to be an offshoot of the genus Alticola which has become special-
ized for life in the crevices of bare rocks; and in this habit and the correlated
cranial characters it affords a parallel to Glirisciis, a similar offshoot from
Giaphiiiriis, the great African genus of Dormice."
No authors dealing with Palaearctic mammals have ever given Plalycranius
full generic rank so far as I know. But when exactly the same specialization
occurs in Africa, it seems it must be generic ! I have often failed to see why
animals must have full generic rank just because they inhabit the African
Continent, but in many cases this seems to be the sole reason.
Fui. 161. Graphiurus hueti hueti, Rochebrune.
Cheekteeth: B.M. No. 25.10.24.21; 10.
It may also be added that Grafiliiunis s.s. has the skull just as flattened as
in Glirisciis.
Between Graphiurus (ucidtiris) and other subgenera there is a wide distinction
in the cheekteeth ; in the former the premolars are extremely reduced. But
contra to the statement of the above key I have been quite unable to detect any
difference in the vagueness of pattern of Graphiurus and many forms referred
to Claviglis. On the reduction of the premolars alone I do not care to give
this species full generic rank, bearing in mind that there is a very general
tendency for certain reduction to take place throughout the genus in these
teeth. Though G. ocularis has reached a stage rather sharply distinct from the
others, it is rather at the end of a long series than so sharply distinct from all
others that it must be considered as a full genus.
At the other end of the series stands the giant West African G. hueti. In
1936 Allen gave this species generic rank under the name .-icthoglis, on a
number of characters which do not prove to distinguish it clearly trom all
other members of the genus.
One of the main characters was the animal's larger size. It seems unnecessary
for me to have to point out that size cannot possibly be used in a generic sense
GRAPHIURUS 607
unless \vc have a genus name for almost ever)' known species. Compared with
Claviglis, the bullae are relatively smaller (the difference not great); the zygoma
is said to be less bowed ; the nasals narrowed throughout instead of being broad
anteriorly. The vomer continuing to the posterior edge of bonv palate, and
dividing the posterior nares; the incisors face anteriorly normally, not turned
inwards as in Claviglis; and P. 4 is farther forward. But the species has evi-
dently not been compared with the much smaller G. crassicaudatus, from the
same area. In this species the nasals are precisely as in " Aethoglis" and the
incisors are as in " Aethoglis." On the remaining characters it is difficult to
regard the hueti group as more than a well-defined specific group of the sub-
genus Claviglis. Even the "large size" character is covered by the Angolan
species monardi.
The genus, which is in much need of revision, is extremely difficult to
arrange in any natural order, .\part from the subgenera, Graphiurus, for
ocularis, and Gliriscus for platyops group, there remains a large assemblage
of more normal .\frican Dormice. Both ^Ir. Hayman and myself have tried to
arrange these into groups, but without much success. Mr. Hayman reports:
"Subgenus Claviglis: this contains the remainder and by far the largest
number of African dormice. .Attempts have been made to divide them into
groups of related forms, but apart from perhaps four easily distinguished
forms the remainder do not appear separable into definite groups. Variation in
size is considerable in some forms, so that overlapping invalidates any arrange-
ment based on size, while although extremes of colour in the subgenus are
wide (from pale grey to brown), division into groups based on colour breaks
down when it is seen that in a very large series from northern Rhodesia nearly
all the colour shades found in forms from elsewhere in the range of the genus
are represented.
The following forms are easily distinguished.
crassicaudatus, Jentink. Nasals and incisors as "Aethoglis," but anterior end of
toothrow behind zygoma root (not so in hueti group, but at base of it),
and size small. (The frontals unusually broad (J.R.E.).)
surdus, Dollman. Nasals as "Aethoglis'' ; size small. (Infraorbital foramen
foramen apparently not as usual in the genus (J.R.E.).)
uoosnami, Dollman. Very pale grey form.
monardi, St. Leger. Size large, head and body 150-160. Skull normal, in no
characters resembling hueti, as suggested by author."
The remainder have a normal skull, with nasals expanded anteriorly to
form part of sides of rostrum. 1 lead and body from 70 to 116.
Forms seen: ansorgei, angolensis, brockmani, butleri, christyi, dorotheae, foxi,
griselda, haedulus, hueti, internus, johnstoiii, lorraineus, microtis, monardi, mon-
tosus, murinus. nanus, ocularis, olga, orubinus, parvus, platyops, raptor, rupicola,
saturatus, smithi, spurrelli, soleatus, surdus, woosnami.
Certainly monardi and crassicaudatus and, I think, probablv woosnami and
surdus are sufficiently distinct to be regarded as types of specific groups. The
39 — Living Kodcnts — I
6oS GRAPHIURUS
remainder will have to be referred to a single group, in which it appears that
there are far too many outstanding "distinct species" at the present day.
List of Named Forms
(The references and type localities for all Muscardinidae are the work of
Mr. R. W. Hayman.)
Subgenus Graphiiirus, Smuts
1. GRAPHIURUS OCULARIS, Smith
1829. Zool. Journ. IV, p. 439.
Plattcnberg Ray, Cape Province.
Synonym: copensis, Cuvier, 1S29, Mamm., pi. 254.
typicus. Smith, .\fr. Zool., 1834, p. 65.
elegam, Ogilby, Proc. Zool. Soc, London, 1S38, p. 5.
(Damaraland).
cattoiri, Fisch. .Synops. Mamm., p. 310, 1829 (mdeter
niinate according to G. M. .Alien, I939)-
Subgenus Glhisciis, Thomas & Hinton
z. GRAPHIURUS PLATYOPS, Thomas
1897. Ann. Mag. Nat. Hist. 6, XIX, p. 388.
Enkeldoorn, Mashonaland.
J. GRAPHIURUS EASTWOODAE, Roberts
1913. Ann. Transv. Mus. IV, p. 80.
Woodbush, Transvaal. (Stated to be related to oculnris, but description
does not bear this out; measurements suggest Glirisnis (R.W.H.).)
4. GRAPHIURUS RUPICOLA RUPICOLA, Thom.is & Hinton
1925. Proc. Zool. Soc. London, p. 232.
Kanbib, S.-W. Africa.
5. GRAPHIURUS RUPICOLA MONTOSUS, Thomas & Hinton
1925. Proc. Zool. Soc. London, p. 233.
Cireat Brukaros Mountain, S.-W. .'\frica.
Subgenus Clari(;lis, Jentink
hueti Group
li. GRAPHIURUS HUETI HUETI, Rochebrune
1S83. Faune Seneg., p. 109, pi. vi, fig. i.
St. Louis, Senegal.
7. GRAPHIURUS HUETI ARGENTEUS, Allen
1936. Journ. Mamm. Baltimore, 17, p. 293.
Lolodorf, Cameroons.
S. GR.APHIURUS HUETI N.AGTGLASI, Jentink
1888. Notes Leyden Mus., X, p. 38.
Dii Queah Ri\er, Liberia.
cnissicaudalus (jroup
>,. GRAPHIURUS CR,-\SSICAUD.'\TUS CR.-\SSICAUD.\TUS, Jentink
18SS. Notes Leyden Mus., p. 41.
Du Queah Ri\'er, L^iberia,
GRAPHIURUS 609
10. GI<.\PH1URUS CRASSICAUDATUS DOROTHEAE, Dollman
1912. Ann. Ma;;. Nat. Hist. 8, IX, p. 312.
Oban district, S.-E. Nigeria.
surdus Group
11. GRAPHIURUS SURDUS, Dollman
1912. Ann. Mag. Nat. Hist. 8, IX, p. 314.
Benito River, French Congo.
woosnami Group
12. GRAPHIURUS WOOSNAMI, Dollman
1910. Ann. Mag. Nat. Hist. 8, VI, p. 393.
North of Okwa, Kalahari Desert.
monardi Group
13. GR-i^PHIURUS MONARDI, St. Legcr
1936. Ann. Mag. Nat. Hist. 10, XVII, p. 465.
Chiumbe River, Angola.
murinus Group
14. GRAPHIURUS OLGA, Thomas
1925. Ann. Mag. Nat. Hist, g, XVI, p. 191.
Asben, South Sahara.
15. GR.APHIURUS OROBINUS, Wagner
1845. Arch. f. Naturgesch., XI, i, p. 149.
Senaar, Sudan.
16. GRAPHIURUS BUTLERI, Dollman
1912. Ann. Mag. Nat. Hist. 8, IX, p. 319.
Jebel .Ahmed .Aga, Sudan.
17. GRAPHIURUS BROCKMANl BROCKMANI, Dollman
1910. Ann. Mag. Nat. Hist. 8, V, p. 287.
Burao, Somaliland.
18. GRAPHIURUS BROCKM.A.NI INTERNUS, Dollman
1912. Ann. Mag. Nat. Hist. 8, IX, p. 318.
Northern Guaso Nyiro, Kenya.
19. GR.APHIURUS FOXI, Dollman
1914. Ann. Mag. Nat. Hist. 8, XIII, p. 196.
Kabwir, Bauchi Province, N. Nigeria.
20. GRAPHIURUS PARVUS PARVUS, True
1893. Proc. U.S. Nat. Mus., XVI, no. 95A, p. 601.
Tana River, Kenya.
21. GRAPHIURUS PARVUS DOLLMANI, OsEOod
1910. Field Mus. Nat. Hist. Zool. ser., X, no. 3, p. 15.
Ulukenya Hills, Kenya.
22. GRAPHIURUS SOLE.VrUS SOLE.ATUS, Thomas & Wroughton
1910. Trans. Zool. Soc. London, XIX, p. 499.
Ruwenzori, Uganda.
6io GR.A.PHIURUS
23. GI<.\PHIURUS SOl.KATUS COLLARIS, Alkn & Loveridsc
1933. Bull. Mus. Comp. Zool. Harvard Coll., LXXV, no. 2, p. 122.
UUinsa Mountains, north of Lake Nyasa. Tanganyika.
24. GRAPHICRU8 PKRSONATUS, HilU-r
1911. Smiths. Misc. Coll. LVI, no. 17, p. 2.
Rhino Camp, Lado Enclave, X. Uganda.
25. CiR.APHIURLS MCRINUS MURINUS, Desmarest
1S22. Mamm. Suppl., p. 54a.
Cape Colony.
Syntmym: coupei, Cuvicr, Mamm. 1S22, pi. 251.
fiythrohromhus. Smith, Zool. Journ. IV, 1829, p. 43S.
ciiH-raceus, Ruppcll, Mus. Scnck. 3, 1842, p. 136 {fide
Trouessart).
lalamiianus, Schinz, 1825, Cuvier's Thicrreich, 4, p. 393.
2h. GRAPHIURUS MURINUS TZANEP:NENS1S, Roberts
1913. .Ann. Transv. Mus. IV, p. 79.
Traiis\aal.
27. GRAPHIURUS MURINUS ISOL.ATUS, Heller
1912. Smiths. Misc. Coll. LIX, no. 16, p. 3.
Taita Hills, Kenya.
2S. GR.A.PHIURUS MURINUS GRISEUS, .Allen
191 2. Bull. Mus. Comp. Zool. Harvard Coll., LIV, p. 440.
Northern Guaso Nyiro, Kenya.
Synonym :jo/;hs/o«/. Heller, 1912, Smiths. Misc. Coll. LIX, 16, p. 2, not
of Thomas.
2y. GR.APHIURUS MURINUS SATURATUS, Dollman
1910. .Ann. Mag. Nat. Hist. 8, V, p. 204.
Mount Elgon, Kenya.
30. GRAPHILRL:S MURINUS RAPTOR, Dollman
1910. .Ann. Mag. Nat. Hist. 8, V, p. 96.
.Mount Kenya.
31. GRAPHIURUS MICROTIS, Noack
18S7. Zool. Jahrh., II, p. 248, pi. ix.
Marungu, S.-E. Congo.
A synonym of G. m. muriniis.fide G. M. .Allen, 1939.
32. GRAPHIURUS SMITHI, Thomas
1S93. Arm. Mag. Nat. Hist. 6, XII, p. 267.
Speke Gulf, Victoria Nvanza.
Synonym: ('f)suhriifus, Neumann, 1900, Zool. Jahrb. Syst. XIII,
p. 547. Tanga, Tanganyika.
33. GR.APHIURUS ANSORGEI, Dollman
1912. .Ann. Mag. Nat. Hist. 8, IX, p. 317.
Mossamedes, S. .Angola.
34. GRAPHIURUS LORRAINEUS, Dollman
1910. Ann. Mag. Nat. Hist. 8, V, p. 2S6.
Molegbwe, south of Setema Rapids, Uele River, Belgian Congo.
GRAi'HIURUS 6ii
35. GRAPHIURUS SPURRKLLI, Dollman
1912. Ann. Mag. Nat. Hist. 8, IX, p. 315.
Bibianaha, Gold Coast.
36. GR.APHIURUS H.\I;DULL'S, Dollman
1912. Ann. Mag. Nat. Hist. 8, IX, p. 316.
Bumba River, Camuroons.
37. GRAPHIURUS CHRISTYI, Dollman
1914. Extr. Rev. Zool. Ah., IV, fasc. i, p. 80.
Maniboka, E. Congo.
38. GRAPHIURUS ANGOLENSIS ANGOLENSIS, de Winton
1897. .Ann. Mag. Nat. Hist. 6, XX, p. 320.
Caconda, Angola.
39. GRAPHIURU.S ANGOLENSIS JORDANI, Roberts
1929. Ann. Transv. Mus. XIII, p. 95.
Isoka, N. Rhodesia.
40. GRAPHIURUS GRISELDA GRISELDA, Schwann
1906. Proc. Zool. .Soc. London, p. 105.
Kuruman, Bechuanaland.
41. GRAPHIURUS GRISELDA PRETORIAE, Roberts
1913. Ann. Transv. Mus. IV, p. 79.
Wonderboom, Pretoria, Transvaal.
(A race of muriniis according to G. M. Allen, 1939.)
42. GRAPHIURUS KELLENI, Reuvens
1890. Die Myoxidae oder .Schlaefer, p. 35, pi. i, fig. i, pi. iii, fig. 3.
Damaraland.
43. GRAPHIURUS NANUS, de Winton
1896. Proc. Zool. Soc. London, p. 799.
Mazoe, Mashonaland.
44. GRAPHIURUS JOHNSTONI, Thomas
1897. Proc. Zool. Soc. London, p. 934.
Zoniba, Nyasaland.
Not seen and not allocated to group
45. GR.'VPHIURUS ALTICOLA, Roberts
1929. Ann. Transv. Mus., XIII, p. 98.
Wakkerstroom, Transvaal.
46. GRAPHIURUS LITTORALIS, Roberts
1929. Ann. Transv. Mus. XIII, p. 97.
\Iasiene, coast of Portuguese E. .\frica.
47. GRAPHIURUS STREETERI, Roberts
1913. .\nn. Transv. Mus. IV, p. 80.
Transvaal.
4.S. GR-APHIURUS TASAUNI, Roberts
1929. Ann. Transv. Mus., XIII, p. 95.
Gwelo, .S. Rhodesia.
6i2 MUSCARDININAE
4y. GRArHILRUS VANDAMI, Roberts
lyzQ. Ann. Transv. Mus.. XIII, p. 97.
Lower Olifants River, Portuguese E. Africa.
50. GRAPHIURUS ZULUENSIS, Roberts
1Q31. Ann. Transv. Mus., XIV, p. 229.
Ubonibo Bush, North Zululand.
51. GRAPHIURUS SCHWABI, G. M. Allen
IQ12. Bull. Mus. Comp. Zool. Harvard Coll. LIV, p. 441.
Kribi, Cameroons;
(A synonym of G. haedtihis, according to CJ. M. .-Mien, 1939.)
5;. GRAPHIURUS PARVULUS, Monard
1932. Bull. Soc. Neuch. Sci. Nat. 57, p. 54.
Rio Mbale, Mossamedes, S. Angola.
53. GRAPHIURUS VULCANICUS, Lonnberg & Gyldenstolpc
1925. Arkiv. Zool. 17B, no. 9, p. 2.
Mount Karisinibi, Birunga Volcanoes, Kivu, E. Congo.
Subfamily MUSCARDININAE
Geogr.'^phic.^l Distribution. — Palaearctic region: Europe from southern
Scandinavia to the Mediterranean, and
England eastwards; Asia Minor; Sinai; North Africa, south to Rio de Oro;
across Russian Asia to Tianshan, Zungaria, and North-west Frontier (specimens
of Dvroinvs from last locality in British Museum); Japan.
NinVIBER OF GENER.'i. — Six.
Ch.'\r.\cters. — DilTering from the Graphiurinae in the more Murine
zygomatic plate, which is tilted upwards to a certain extent,
the muscle attaching line of attachment on its forepart; masseter lateralis
superficialis with anterior head distinct. No caecum (so far as known). Cheek-
teeth i.
In Eliomys, the cheekteeth are hasin-shaped and cuspidate much as in
Graphiuriis; the premolar is not reduced, and there are on all main upper teeth
two high main outer cusps, and one long main inner cusp (as in normal Sciuri-
dae); the cross-ridges are arranged much as in Sciuridae. In Dyromys, the
premolars are reduced, and the cheekteeth are less concave, but the general
dental effect is near Eliviiivs. In Glis, the cheekteeth are more nearly flat-
crowned, with five or six low cusps on outer margin of upper main teeth; the
skull is more strongly ridged than in the other genera; but M.i is not con-
spicuously different in size from M.2; in Muscardinus, a more specialized
dental effect is present, the premolar being vestigial, the first molar much
larger than the second, the ridges arranged differently, and the crowns ot the
teeth are flat.
These four genera have been thorouglily dealt with in Miller, Catalogue of
Mammals of Western Europe, p. 549, 1912. The remaining genera, Glirulus
and Myomimi'.s are very little known; the latter is not represented at the
British Museum.
MUSCARDININAE: ELIOMYS 613
Key to the Genera of Muscardimnae
(not including the genus Myomimus which has not been examined)
Crowns of cheekteeth flat; M.i much larger than M.2, the ridges of this
tooth arranged differently, the depressions between them unusually
wide (angular portion of mandible with perforation; tail not
distichous). Muscardinl's
Crowns of cheekteeth not completely flat; M.i not conspicuously larger
than M.2, the ridges of this tooth not arranged differently, the
depressions between them not unusually wide.
Bullae low, relatively small, and flat, scarcely rising above general
level of base of skull. (Mandible without perforation in angular
process.) Glirulus
Bullae large, well inflated, rising clearly above general level of base
of skull.
Outer side of upper main cheekteeth with five or six low cusps;
crowns nearly flat, .\ngular portion of mandible not per-
forated. (Tail conspicuously distichous; skull with rather
well-marked supraorbital ridges.) Glis
Outer side of upper main cheekteeth with two high cusps; crowns
concave. Angular portion of mandible perforated. (Skull
without clear supraorbital ridges.)
Cheekteeth markedly concave; premolars clearly cuspidate;
tail not uniformly haired. Eliomys
Cheekteeth less markedly concave; premolars not clearly
cuspidate; tail uniformly haired. Dyromys
The position of the genus Glirulus must be regarded as provisional owing
to the scarcity of material available. The unrepresented genus Mvomimus
differs from all the above in the character of its tail, which is stated to be scantily
haired, like that of a Mouse.
Genus I. ELIOMYS, Wagner
1843. ELio.vn'S, Wagner, Abb. Bayer. .Akad. Wiss. Miincben, math.-phys. Ill, p. 176.
1885. BiFA, Lataste, Le Naturaliste, no. 8, pp. 61-63. Bifa lerotina, Lataste.
Type Species. — Eliomys melanurus, Wagner.
Range. — Continental Europe, from Iberian Peninsula, France and Italy,
north to Baltic coast of Germany; Dalmatia; Balearic Isles;
Corsica, Sardinia; Sicily; Russia (former Smolensk, Leningrad, Novgorod,
Tver, Orel, Kiev, Ulianov, Orenberg governments) (Vinogradov). Asia Minor
(Miller). Sinai, Syria. North-western Africa, from Tunis, Cyrenaica, and
Algeria to Morocco, south to Cape Blanco.
NiiMBER of For-MS. — Thirteen.
Fig. 162. Eliomys quercinus quercinus, Linnaeus.
B.M. No. 8.8.4.64, V; :• 2i.
Fig. 163. Eliomys qiercinus qliercinus, Linnaeus.
B.M. No. 8. 8.4.64, i; .• 2.!.
ELIOMYS
615
Characters. — Skull strongly constricted between the frontals; rostrum
relatively long; superior portion of skull not or scarcely
ridged. Jugal relatively long. Palate broad, the palatal foramina situated
considerably in front of toothrow, and narrowed anteriorly. Bullae large and
inflated. The palate, as in allies, usually has a small pair of foramina present at
posterior border. Infraorbital foramen narrow; zygomatic plate clearly tilted
upwards, though relatively narrow compared
with average IVluridae. Angular portion of
mandible perforated. Cheekteeth with crowns
conca\e; in upper series, there are two high
main cusps on the outer side, and one on the
inner side; and four main transverse ridges
are present, separating three depressions, the
general effect reminiscent of that of Sciuridae.
P.4 slightly smaller than the molars, well
cusped. M.3 slightly smaller than M.2.
Lower molars with three outer and two inner
cusps; more basin-shaped than the upper
teeth; four main ridges present, the anterior
and posterior of which form the terminal
margins of the teeth. Premolars with three
cusps, one each side, one anteriorly, and
with one ridge.
Mammae 8 [qiiercimis). Tail rather narrow,
w'ell haired but not conspicuously bushy, the hairs on the lower portion longer
and more thick than those of the upper portion. Hindfoot with D.5 nearly as
long as three central digits, and hallux short. Forefoot with four well-marked
digits.
I do not think that there is more than one valid species of this genus in
Europe. In tact I think with adequate material the North African species would
mostly be referable to quercinus as races as well. A few skins seen of melanurus
seem di.stinct by their paler coloration; and the bullae seem larger than in other
forms except cvrenakus, which also seems distinct externallv in the imiformly
dark and more bushy tail. But whether " nuinbyamis" is distinct from quercinus
is at the moment not clear.
Forms seen: " amori." cyrenaiais, gymnesicus, lerotinus, lusitanicus, melanurus,
munbyanus, occidentalis, ophinsae, pallidus, quercinus, sardus.
Fio. 164.
Eliomys QVERCINT'S
Cheekteeth; x lo.
List of N.\med Forms
t. ELIOMYS QUERCINUS QUERCINUS, Linnaeus
1766. Syst. Nat. i, 12th Ed., p. 84.
Germany.
Synonym: horlualis, Cabrera, 1904, Bol. Real. Soc. Espan. Hist. Nat.
IV. p. I S3. Valencia, Spain.
hamiltoni, Cabrera, 1907, Bol. Real. Soc. Espan. Hist. N'at.
VII, p. 226. El Pardo, near Madrid, Spain.
6i6 ELIOMVS— DYROMV.S
2. ELIOMYS QUERCINUS SL'PERANS, Ognev & Stroganov
1936. Abs. Works. Zool. Inst. Moscow State Univ. 3, p. S4.
Kalinin district, Penorsk region; River Jukopa, the right tributary of
the Volga (former Ostashov subdistrict of the Tver government)
(.Russia).
3. ELIOMYS QUERCINUS GYMNESICUS, Thomas
1903. Ann. Mag. Nat. Hist. 7. XI. p. 494.
San Cristobal, Minorca.
4. ELIOMYS Ql'liRCINUS PALLIDUS, Barrett-Hamilton
1S99. Ann. Mag. Nat. Hist. 7, III, p. 226.
Palermo, Sicily.
Synonym; cincticauda. Miller, 1901, Proc. Biol. Soc. Washington, XIV,
p. 39. Sorrento, Italy.
5. ELIOMYS QUERCINUS SARDUS, Barrett-Hamilton
1901. Ann. Mag. Nat. Hist. 7, VII, p. 340.
Tricoli, Sardinia.
6. ELIOMYS QUERCINUS LUSITANICUS, Reuvens
1890. Die Myoxidae oder Schlaefer, p. 28, footnote.
Lisbon, Portugal.
Synonym: aniori, CJraels. 1897, Mem. Real. Acad. Sci. Madrid, XVII,
p. 48 1. Cordova, Spain.
7. ELIOMYS QUERCINUS OPHIUSAE, Thomas
1925. .-Vnn. Mag. Nat. Hist. 9, XVI, p. 3S9.
Fonnentera, Balearic Islands.
8. ELIOMYS MUNBYANUS MUNBYANUS, Pomel
1856. C.R. Ac. Sci. Paris, XLII, p. 653.
Tunis.
9. ELIOMYS ML NBYANUS LEROTINUS, Lataste
18S5. Le Xaturaliste. p. 61.
Ghardaia, Mzab, Algerian Sahara.
10. ELIOMYS MUNBYANUS TUNETAE, Thomas
1903. Ann. Mag. Nat. Hist. 7, XI, p. 495.
Karouana, Tunis.
(A synonym of E. in. munbyanus, according to G. M. Allen, 1939.)
11. ELIOMYS MUNBYANUS OCCIDENTALIS, Thomas
1903. Nov. Zool., X, p. 300. Rio de Oro, W. Sahara.
12. ELIOMYS CYRENAICUS, Fcsta
1922. Boll. Mus. Zool. Anat. Comp. Torino, 740, p. 4.
Gheminez, Cyrenaica, N. Africa.
13. ELIOMYS MELANURUS, Wagner
1843. Abh. Bayer. Akad. Wiss. Munchen, p. 176. pi. 3. fig. i.
Sinai.
Genus 2. DYROMYS, Thomas
1906. Dryomys, Thomas, Proc. Zool. Soc. London, 1905, p. 34S. Not of Philippi.
1907. Dy-romys, Thomas, Ann. Mag. Nat. Hist. 7, XX, p. 406.
Typk Species. — Mus iiitcdula, Pallas.
R.i^NGE. — From SwitZfrland, North Italy, Silesia, and S.-E. Europe (Greece,
Bulgaria, Yugoslavia), across Russia (Ukraine, former Tver, Orel,
Fig. 165. Dyromys nitedila nitedlla, Pallas.
B.M. No. 12.12.17.12; .< 3i.
FiC. 166. D'YROMVS NITEDL'LA NITEDLLA, Pallas.
B.M. No. 12. 12. 17.12; X 3J.
6i8
DVRO.MYS
district, Fergana, Semircehia,
\'oroncj go\-ts., Bessarabia, Astrakan, Lower Volga, former Kasan govt.)
(Vinogradov), Caucasus and Asia Minor to Russian Turkestan (Tashkent
former Scmipalatinsk go\1.), Persia, I'ianshan,
Dzungaria, and North-West Frontier (North
India).
Number of Forms. — Eighteen.
Ch.'\r.'\cters. — Very closely related to
Eliomys; upper cheekteeth
less concave; the main cusps arranged as in
Eliomys; five main transverse ridges in upper
teeth, the main central depression with quite
a well-marked ridge, this vestigial in Elioiiivs;
premolar more reduced, not strongly cuspi-
date. Lower molars with four main ridges,
and three rudimentary ones between them.
Premolar reduced, and simple.
Parietals not narrowed to a point anteriorly,
differing in this character from EUomys.
Size smaller, and tail more uniformly haired, flattened and moderately bushy.
Mammae 8 (type species).
Remarks. — This genus is not very widely separated from Eliomys. The
cheekteeth are tending to be a little less complex.
Fig. 167
d'vromvs nitedila
Cheekteeth; X lo.
I77S.
1902.
4.
1927.
1907.
1920.
List of Named Forms
DYROMYS MTF.DULA NITEDUL.A, Pallas
Nov. Spec. Quadr. Glir. Ord., p. 88.
Region of Lower Volga, Russia.
Synonym: dryiis, Schreber, 17S2, Saugth., pi. CCXX\', B.
DYROMYS NIT!-:DI"L.A INTERMEDIUS, Nehring
Sitz. Ber. Ges. Nat. Fr. Berlin, p. 155.
Near Lienz, Tirol, Austria.
DYROMYS NITEDULA WINGEI, Nehring
Sitz. Ber. Ges. Nat. Fr. Berlin, p. 5.
Parnassus, Greece.
DYROMYS NITEDULA CARPATHICUS, Brohmer
Die Tienv. Mitt. Europ. 7, lief. 3, p. 32.
Upper Silesia.
DYROMYS NITEDULA PHRYGIUS, Thomas
Ann. Mag. Nat. Hist. 7, XX. p. 407.
.Murad Dagh, Ushak Province, .\sia Minor.
DYROMYS NIT1:DULA TICHOMIROWI, Satiinin
Trav. Mus. Georg. Tiflis, no. 2, p. 161.
Tiflis, Caucasus.
DYROMYS 619
7. DYROMYS N1T1;DUI,A OBOLEiNSKII, Ognev & Worobiev
1923. Fauna Woronesh, p. 129.
Voroncj government district, Russia.
S. DYROMYS NITEDULA OGNEVI, Heptner & Formozoff
1928. Zool. Anz. 77, p. 27S.
Daghestan, E. Caucasus.
9. DYROMYS NITEDULA BILKJEWICZI, Ognev & Heptner
1928. Zool. Anz. 75, p. 265.
Mikhailovsky, Kopct-DaKh, 46 miles west-south-west of Askhabad,
Russian Turkestan.
.0. DYROMYS NITEDULA ANGELUS, Thomas
1906. .A.nn. Mag. Xat. Hist. 7, XVII, p. 424.
Tian Shan, Central .Asia ; near Przewalsk.
11. DYROM\-S NITEDULA CAUCASICUS, Ognev & Turov
1935- Wiss. Ber. Moskauer Staats. Univ. 4, p. 98.
Environs of Tarskaja station. Northern Caucasus (former Tersk
Province).
12. DYROMYS NITEDULA DAGESTANICUS, Ognev & Turov
1935. Wiss. Ber. Moskauer Staats. Univ. 4, p. 98.
Khasav-Jurt, Daghestan, Caucasus.
13. DYROMYS NITEDULA KURDIST.'\MCUS, Ognev & Turov
1935. Wiss. Ber. Moskauer Staats. Univ. 4, p. loi.
Riv. Terter, Kurdistan.
14. DYROM\'S NITEDULA P.ALLIDUS, Ognev & Turov
1935. Wiss. Ber. Moskauer Staats. Univ. 4, p. 102.
Vail. Riv. Bosturgay, Karatau Mountains, former Province of Syr-
Darya, Turkestan.
15. DYROMYS NITEDULA T.\NAITICUS, Ognev & Turov
1935. Wiss. Ber. Moskauer Staats. Univ. 4, p. gS.
Atamano\sky Khutor, Tarasovsky district (former Don Province),
S. Russia.
ih. DYROMYS NITEDULA PICTUS, Blanford
1875. Ann. Mag. Nat. Hist., 4, XVII, p. 311.
Kohrud, south of Caspian, Persia.
17. DYROMYS MILLERI, Thomas
1912. .■\nn. Mag. Nat. Hist. 8, IX, p. 394.
Bogdo-Ola Mountains, S.E. Dzungaria, Central Asia.
.8. DYROMYS ROBUSTUS, Miller
1910. .\nn. Mag. Nat. Hist. 8, VI, p. 459.
Rustschuk, Bulgaria.
Forms seen: angelus, milleri, nitedula, obolenskii, pictus, phrygius, robtistus,
witigei.
620 GLIRULUS— GLIS
Genus 3. GLIRULUS, Thomas
1906. Glirl'lis, Thomas, Proc. Zool. Soc. London, 1905, p. 347.
Type Species. — Graphiurus elegans, 'remminc\<.= Myoxiis japoiiiciis, Schinz.
Range. — Japan.
Number of Forms. — One.
Ch.^racters. — "With regard to the generic position of this Dormouse I
think it cannot be assigned to any of the existing groups, and
must have a special name of its own. It is no doubt most nearly allied to Ellomys
(Drvomvs, subgen. 11.) nitediihis, Pallas, but may be readily distinguished by the
rather more complicated pattern of its teeth, its small bullae, the absence of the
angular foramen in its mandible, and by its peculiar colour pattern" (Thomas).
There are only two skulls in the British Museum of this genus, and two
specimens in spirit. The teeth are too worn in the skulls for any detailed notes,
though apparently nearer Dyromys than Glis. The size is small, the tail bushy;
according to Thomas the mammae are 8. The main genus distinction is the
possession of the small low flattened bullae, conspicuously different from those of
other Muscardininae examined. The premolars are smaller than the molars, as in
Dvromvs.
Forms seen : japonicus.
List of N.amed Forms
I. GLIRULUS J.APONICL^.S, Schinz (emended by Thomas from "jaTam'cus^')
1845. Syst. Verz. Saug., II, p. 530.
Japan.
Synonym: elegans. Temminck, 1845, Faun. Japon. Mamm., p. 53.
lasiotis, Thomas, iSSo, Proc. Zool. Soc. London, p. 40.
Genus 4. GLIS, Brisson
1762. Glis, Brisson, Regn. Anim. Class, IX, 2nd ed., p. 13.
1780. Myoxus. Zimmermann, Geogr. Ges. II, p. 351. (Sclunis glis, Linn.)
^I'y'PE Species. — Glis, Br\sson = Sciiirus glis, Linnaeus.
Range. — Continental Europe from Atlantic coast of France eastwards,
north to North Germany, south through Switzerland and Italy to
Sicilv; Northern Spain; Sardinia; Yugoslavia, Roumania; Asia Minor, Persia;
Russia (former Minsk, Podol, Volvn, Kiev, Harkov, Astrakan, Samara, Saratov,
Pensa, Ulianov gov'ts.) (Vinogradov). Bessarabia, Caucasus, and South Turk-
menia. Introduced in England.
Number qf P'orms. — Eleven.
Ch.\r.'vcters. — Interorbital region of skull well ridged, the ridges tending to
unite in old age. Jugal approaching the laclirymal. Rostrurh
less pointed than in Eliomys. Bullae prominent. Zygomatic spread relatively
F'f- i6S. Gus
GLis CLis, Linnaeus.
BM. No. 6.8.4.1, J; 'x
Fig. 169. Glis clis rr is i ■
622
GLIS
great. The zygoma is in some ways reminiscent ot that of Anomalurtis, though
in the latter genus the infraorbital foramen has become much more widely open
^_^ for muscle-transmission whereas in
Glis the zygomatic plate has become
more broadened. Mandible without
perloration in angular process;
coronoid noticeably powerful.
Cheekteeth simpler than in Elioiitvs,
more flat; the outer side of upper
series with fiye low cusps, the inner
side with four. M.i and M.2 with
seven transverse ridges of which
four are well developed, the three
alternating between them weaker.
P. 4 considerably smaller than the
other teeth, and with its elements
reduced. Lower teeth like the upper
series in general arrangement.
Fur thick and soft; tail densely
bushy; feet broad; general appear-
ance of animal Squirrel-like. Size
rather large for family (head and
body up to 190 mm.). Mammae
12. D.5 hindtoot long, about equal
to D.2."
Forms seen: glis, insidaris, itidicus, inclonii, " postns" spoliatus.
Fif;. 170. Glis glis
Cheekteeth: :• 10.
1766.
List of N.wied Forms
GLIS GLIS GLIS, Linnaeus
Syst. Nat. I, i2th Ed., p. S7.
1S9S.
1920.
Germany.
Synonym :
csciileiittis, Blumenbach, 1779, Handb. Nat., p. 79.
vulgaris, Oken, 1816, Lehrbuch. Nature. Ill, pt. 2, p. 868.
(Gemiany.)
aiellainis. Owen, 1840, Odontography, II, p. 25, pi. 105.
(^)giglis, F. C\nier, 1S32, Nouv.
Paris, I, p. 444, num. nud.
.^nn. Mus. d'Hist. Nat.
GLIS GLIS ITALICUS, Barrett-Hamilton
.Ann. Mag. Nat. Hist. 7, II, p. 424.
Siena, Italy.
Synonym: iiisulniis, Barrett-Hamilton, iSqg
III, p. 228. Palermo. Sicily.
postiis, iMontauu, 1923, Pmc Zi
"^ 'ugo-Sln\*ia.
GLIS GLIS INTER.MEDIUS, Altohello
Fauna deli' Abruzzo e del Molise. Mamniiferi, III (Rodentia), p. 22
Abruzzi, Italy.
.Ann. Mae. Nat. Hist. 7,
il. Sue. London, p. 866.
GLIS— MUSCARDINUS 623
4. GLIS GLIS ABRUTTI, Mtobello
1924. Rend. Union. Zool., p. 30; fig. in Monitore Zool. Ital. 35.
S. Italy.
5. GLIS GLIS MINUTU.S. Martino
1930. Proc. Russ. Sci. Inst. Belgr. 2, p. 60.
Serbia : Predejane, 30 km. south of Leskovac.
6. GLIS GLIS PYRENAICUS, Cabrera
1905. .Ann. Mag. Nat. Hist. 8, I, p. 193.
Alio, Navarra, Spain.
7. GLIS GLIS MELONII, Thomas
1907. Ann. Mag. Nat. Hist. 7, XIX, p. 445.
Marcurighe, Ogliastra, Sardinia.
8. GLIS GLIS ORIENTALIS, Nehring
1903. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 1S7.
Near Scutari, .Asia Minor.
g. GLIS GLIS SPOLI.ATUS, Thomas
1906. Ann. Mag. Nat. Hist. 7, XVIII, p. 220.
Khotz, near Trebizond, Asia Minor.
10. GLIS GLIS TSCHETSHENICUS, Satunin
1920. Trav. Mus. Georg. Tiflis, no. 2, p. 150.
Caucasus.
11. GLIS GLIS CASPICUS, Satumn
1905. Mitt. Kaukas. Mus., II, p. 55.
Aschabad, Transcaspia.
There is reason to believe that the name persicus, Er.xleben, Syst. Regn.
Anim. i, p. 417, 1777 (which has been used for Sciurus anomaltis), is based on
a form of this species.
Genus 5. MUSCARDINUS, Kaup
1829. MUSCARDINUS, Kaup, Enr«-. Ges. Nat. Europ. Thierwelt, I, p. 134.
Type Species. — Mtis avellanarius, Linnaeus.
Range. — England; France; Southern Germany; Central Sweden; Italy,
Sicily; Austria; Greece; Asia JSIinor; Russia (former Vitebsk,
Minsk, Smolensk, Moscow, Vladimir, Kostroma, Harkov, Kasan, Ulianov,
Kiev, Poltava, Volyn, Podol, Odessa governments, Bessarabia) (Vinogradov).
Number of Forms. — Five.
Characters. — Zygomatic plate rather broader than in allied genera, its
superior border ridged; infraorbital foramen small. Incisive
foramina rather longer than usual, and not widened posteriorly. Zygomata
widely spreading anteriorly. Palate very short, not extending back to M.3.
-A small perforation in angular portion of mandible usually present. Bullae
moderately inflated.
40 — Living Rodents — I
Flo. 171. MliCiRDINU; A-.-ELLAN-IRirS A\-EU^-^
B.M. No. 2.M.3'.', 2 ; ' 3*.
i^jrmaeus.
172. .M:;^', ijtULvui .'i\'£LL.t.x.«iKiL's .i%TLL«.'»'.4iiiv s. Linnaeus.
B.M. No. 2.is.4.3fc, -; ^ 35.
MUSCARDINUS 625
P.4 very small indeed. M.i conspicuously lai^r than M.2, with five well-
developed ridges, the depressions between them very broad; the ridges oblique,
the tooth lengthened; M.2 with seven transverse
ridges, the depressions between them narrow;
M.3 like M.i but smaller, with ridges less
developed. P.4 usually with two ridges. In the
lower cheekteeth there are six ridges extending
across each tooth except the much reduced
premolar; the lower first molar is less enlarged
than the upper first molar, and its ridges less
oblique.
Size very small, head and body under 100.
Forefoot with "digits relatively longer than in
the other European genera, and closing obliquely
inward so as to come into opposition with the
much enlarged inner tubercle, the unusual size of
which . . . enables it to function as a low, broad
thumb" (Miller). Hindfoot with four long digits,
and hallux more rudimentary than usual. Tail
uniformly haired, said to be partly prehensile, more narrowed than is usual
in the group. Stomach extremely complex (Thomas). Mammae (type species) S.
Re.m.\rks. — This genus is very distinct from the other members of the sub-
family, and in several wavs its type of dentition seems to be
leading towards that of the Platacanthomvinae.
Forms seen: "anglicus" (izrllaniiriiis. ptikher, trapezius.
Fig. 17,^
mlsc.\rdinvs .\vki,u\narus
Cheekteeth; \ 10.
17S8.
1920.
1932-
4-
1898.
List of Named Forms
MUSCARDINUS AVELL.\NARIUS AVFLLAXARIUS. l.innacus
Sj'st. Nat. I, loth Ed., p. 62.
Central Sweden.
S\Tion\nii: muscardimis, Schreber, 1782, Siiugth., pi. CCXWll.
Cjeniiany.
ttvellanarius anglicus, Barrett-Hamilton, 1900, Proc. Zool.
Soc. London, p. 86. Northampton, England.
corilimim, Fatio, 1S69, Faune Vert. Suisse, i, p. 183.
MUSCARDINUS AVELLANARIUS NIVEUS, Aliobello
Fauna dell' Abruzzo e del Molisc. Mamniiferi, III, Rodcnti;i, p. 27.
Abruzzi, Italy.
MUSCARDINUS AVELLANARIUS ZEUS. Chaworth-Musters
.\nn, Mac. Nat. Hist. 10, IX. p. 170.
East slope Mount Olympus, Thcssaly, Greece.
MUSCARDINUS I'ULCHER, Barrett-Hamilton
Aiui. Mag. Nat. Hist. 7, II, p. 423.
Perugia, Italy.
Synonym: (})spfdosiis, Dehnc, 1S55, .-\llgem. Deutsche Naturhist.
Zeitung, i, p. iSo. Tursi, Basilicata, Italy.
626 AnOMIMUS-^l'LATACANTHOMVlNAE
5. MrSCARDINUS TRAPEZIUS. Miller
igoS. Ann. Mag. Xat. Hist. S, I, p. 6q.
Trchizond, Asia Minor.
Genus 6. MYOMIMUS, Ognev
1924. MvoMiMi's, Ogncv, Nature and Spurt in Ukraine, Kharkov, p. i.
Type Species. — Myumimus f^crsuiiattis, Ognev.
R.'iNGE. — Described from Transcaspia, near the Persian frontier.
Number of Forms. — One.
This genus is not represented in the British Museum. The tail is described
as being thinly haired, as in Mice. The cheekteeth arc described as near those
of Dyromys. In the skull as figured by Ognev & Heptner, it may be noted that
the bullae are well inflated, the angular portion of the mandible perforated, and
the palate normal (not as in Platacanthomyinae).
List of Named Forms
I. MYO.Ml.MUS PERSONATUS, Ognev
1924. Nature &: Sport in Ukraine, Kharkov, p. i.
Transcaspia : near Persian frontier.
Subfamily PLATACANTHOMYINAE
Geogiwphical Distribution. — Peninsular India; South China.
Number of Genera. — Two.
Characters. — Differing from Muscardininae in the suppression of the
premolars, the more definite and more specialized pattern
of the cheekteeth, the presence ot a large series of foramina or a single pair of
very large foramina between the toothrows, the much smaller bullae, and in the
genus Typhlomys the presence of a small caecum.
There may be some doubt as to whether this group is rightly placed in the
Muscardinidae. Peters came to the conclusion that Platacaiithomvs was a
member of the Muridae showing signs of affinity with Phloeomvs and Meriones.
Thomas in his classification of the Order in 1896 stated: "Dr. Winge has placed
Platacanthomys in the Gliridae, from which it was removed to the Muridae by
Dr. Peters, and in this he has been followed by Dr. TuUberg, and I am informed
by Dr. Forsyth Major . . . that he holds a similar view. On the whole though
I think there is enough evidence of Murine affinity in Platacanthomys and its ally
Typhlomys to make the question rather doubtful, I am inclined to agree to the
reference of these genera to the family Gliridae, on account of the structure of
the teeth and interorbital region, the peculiar Glirine twisting of their mandibular
angles, and of their (at least the former's) want of a caecum, a character found
in the Gliridae alone of Rodents." But some years later Thomas discovered
that in Typhlomys a small caecum is present.
PLATACANTHOMYINAE : PLATACANTHOMYS 627
Miller & Gidley, 1918, and Weber, 1928, separate the two genera as a family
the Platacanthomyidae, which the former define as "Like the Cricetidae but
zygomasseteric structure unusual; infraorbital foramen of normal Cricetine
form, but zygomatic plate much narrowed, and masseter lateralis profundus
extending its line of attachment along upper zygomatic border to side of rostrum
above foramen. Cheekteeth subhypsodont, enamel pattern a modified hcp-
tamerous, with tendency to form oblique parallel cross-ridges (parallel Mus-
cardinidae)."
Hut the zygomatic plate is no more narrowed than in Hydromys, which
these authors refer to their Muridae; something similar judging from TuUberg's
figure seems to occur in that genus, in the zygomasseteric structure.
It has been suggested to the present author that Typhlomys is a Dipodoid.
But in dental structure, and in the much more important zygomasseteric struc-
ture, it is very clear that this is not the case.
I do not think that there is very much doubt that these two genera should
form a well-marked subfamily of the present family. Mtiscardinus, as stated
above, seems to be leading towards Platacanthomys in dental characters, though
considerably less modified than in that genus. On the pattern of the cheekteeth
I do not think the present group could be referable to the Muridae; and there is
very little doubt that Typhlomys is a close ally of Platacanthomys.
Key to the Gener.\ of Pl.\t.\canthomyin.'ve
Caecum absent (Thomas). Tail thickly bushy, shorter than head and
body. Fur densely spiny. Skull with well-marked supraorbital
ridges. Palate with one very large pair of foramina between the
toothrows. Pl.\t.\c.\nthomys
Caecum present (Thomas). Tail longer than head and bodv, poorlv
haired except terminally. Fur soft. Skull with supraorbital
ridges feeble or absent. Palate with a series of foramina between
the toothrows. Typhlomys
Genus i. PLATACANTHOMYS, Blyth
1859. Plat.^canthomys, Bljth, Joum. Asiat. Soc. Bengal, XXVIII, p. 288.
Type Species. — Platacanthomys lasiunis, Blyth.
Range. — Southern India (Malabar, Coorg, Travancore).
Number of Forms. — One.
Characters. — Skull of the "arboreal" type seen in many Muridae, with
broad frontals, even broader braincase, and very large inter-
parietal. Hea\T supraorbital ridges present, extending backwards on to brain-
case. Zygomatic plate very narrow, but tilted strongly upwards.
Jugal not extending so far forwards as in most Muscardinidae. Palate
broad; a large pair of foramina between the toothrows take up most of this
Fig. 174. Platac.anthomys lasiurus, Blytli.
B.M. No. 13.8.22.57, 3; /, 2i
Fig. 175. Pl.^tai'anthomvs i.asU'rl's, Blyth.
B.M. Nn. 13.8.22.57, o"; ■ 2i.
PI.ATACANTHOMYS— TYPHLOMYS 629
space. Incisive foramina very far forwards. Bullae small, reduced. Mandible
with low coronoid, and angular portion without perforation, and pulled inwards
as is usual in the family.
Upper cheekteeth with four depressions appearing as re-entrant folds, the
second one cutting right across the tooth; in .M.i and I\1.2 the folds curving
obliquclv backwards from outer side. The transverse ridges (between the
depressions) much broader than in Muscardininae. Lower cheekteeth with
Fig. 176. Platacanthomys lasiurus, Blyth.
Cheekteeth: 13. M. No. 13.8.22.57, cj; X 10.
four long re-entrant folds, the front one usually isolated on crown surface, the
others placed externally. The pattern is clearer and more definite than in
Muscardininae.
Tail thickly bushy terminally, less so on joining the body. Hindfoot very
broad, hallux more reduced than in other members of the family, Muscardinus
perhaps excepted, and with no claw (from its appearance on dried skins, perhaps
opposable). D. 5 about as long as D.2. Back and sides covered with flattened
spines.
Forms seen : lasiurus.
List of N.\mf.d Forms
t. PLATACANTHOMYS LASIUKUS. Blyth
1859. Journ. .\siat. Soc. Bengal. XXVIII, p. 289.
.^lipi, Malabar, India.
Genus 2. TYPHLOMYS, Milne-Edwards
1877. Typhlomvs, Milne-Edwards, Bull. -Soc. Philom. Paris, XII, for 1876, pt. 2, p. 9.
Type Species. — Typhlomys cinereus, Milne-Edwards.
R.\NGE. — Known from Fokien (South China), and Tongking.
Fig. 177. Typhlo.mys cinereus cinerels, Milne-Edwards.
B.j\l. No. 8. 8. II. Ill, i; ■ 4.
Fig. 17S. T'iTHLOMYS cinereus cinereus, Milne-Edwards.
B.M. No. 8. 8. II. Ill, 9; X 4.
TYPHLOMYS 631
Number of Forms. — Two.
Characters. — Skull with rounded braincase and almost unconstricted
interorbital rtfjion. Infraorbital foramen and zygomatic
plate much like Platacaiithomys. Huiiac- .small. Palate with a series of foramina
extending from palatal foramina, which are as usual far in front of toothrows,
to back of palate, the number of these foramina varying in different specimens.
Coronoid low; angular process reduced, and scarcely pulled inwards. Cheek-
teeth narrow. Originally evidently there are five folds cutting obliquely across
Fig. 179. Typhlomys cinereus cinerecs, Milne-Edwards.
Cheekteeth: B.M. No. 8.8.11.111, ?; x 14.
each tooth (often isolated on surface), but sometimes some of these wear out.
Lower teeth with five folds or depressions, less isolated than in the upper series,
particularly the third and fourth, which are long. Only a small series of skulls
at present available for examination.
Size very small. Tail considerably longer than head and body, scalv, naked
or with a faint growth of rather long hairs through the lower part of its length,
which at the end are produced into a brush. Fur without spines. Hindfoot
evidently not specially broadened, but D.5 long. Eyes much reduced. A small
caecum present (Thomas).
Forms seen : cinereus, chapensis.
List of Named For.ms
1. TYPHLOMYS CINEREUS CINEREUS, Milnc-Edwards
1877. Bull. Soc. Philom. Paris, XII, p. 9.
Fokien, China.
2. TYPHLOMYS CINEREUS CH.APENSIS, Osgood
1932. Field iMus. Nat. Hist. Zool. Scr. XVIII, no. 10, p. 29S.
Chapa, Tongking.
632 LOPHIOMYIDAE: LOPHIOMYS
The family Muscardinidae is known fossil in the Palaearctic region from the
Middle Miocene.
Before passing to the other Muroid families, all of which are very closely
allied to each other, it must be noticed that according to TuUberg all other
Muroidae as here understood examined by him have a hornv layer present in
the cardiac portion of the stomach; but this is not the case in Muscardinidae
examined bv him; I have no notes on the character in Platacanthomxs and
Txj^hlomvs.
MUSCARDINIDAE:
SPECIAL WORKS OF REFEREXCE
Miller. Catalogue of Mammals of Western Europe, 1912, p. 549. Muscardinidae.
Tl'LLBERG, Nova Acta Res. !3oc. Sci. Upsaiiensis, iSgg.
Cl'vier, Du genre Graphiure: Nouv. .Ann. Mus. d'Hist. Nat. Paris, I, I'A'iz.
Peters, On the systematic position of Platacanthomys lasiiirus, Proc. Zool. Soc. London,
1865, p. 3Q7.
Reuvens, Die Myoxidae oder Schlaefer, iSyo, p. i.
The remaining families in the Order are all closely allied to each other;
the majority of the genera are referred to the Family Muridae which contains
well over half the Order. Three groups, one containing two genera, the others
a single genus each, appear to be too aberrantly specialized to be referred to the
Muridae.
Family LOPHIOMYIDAE
1896. Thomas: Myomorph.'\, part: Family Muridae, part. Subfamily Lophiomyinae.
1899. TuUberg: Myomorph.\; Muriformes, part: Family Lophiomyidae.
1918. Miller & Gidley: MuROtD.\E, part: Family Cricetidae, part. Subfamily Lophio-
myinae.
1924. Winge: Family Muridae, Cricetini, part, Criceti, part.
1928. Weber: Mt'ROlDE.'k, part: Family Muridae, part. Subfamily Lophiomyinae.
Geographical Dlstribution. — Eastern Africa : Abyssinia, Somaliland,
Kenya, and Sudan (Kassala district).
Number of Gener.>\. — One.
C'har.\cters. — Zj-gomasseteric structure as in typical specialized Muridae,
but skull highly abnormal, the temporal fossae roofed in by
plates of bone rising from the frontals, parietals and jugals, a structure not
known elsewhere in the Order. Dental formula i. \, c. f;, p. |^, m. i| = 16;
cheekteeth rooted laminate, each lamina separated by wide valleys, and
bearing two cusps (parallel — cuspidate Cricetinae). Fur with an erectile crest
on the centre of the back. Tail densely bushy. Bullae reduced, small. Feet
considerablv specialized for arboreal life, with hallux partly opposable.
Genus I. I.OPIIIOMYS, Milne-Edwards
1S67. LoPHiOMVS, Milne-Edwards, L'InstitiU, vol. 35, p. 46.
Type Species. — Lophioinvs iinJiausi, Milne-tldwards.
LOPHIOMYS
Range. — As in the Family Lophiomyidae.
Number of Forms. — Six.
633
Characters. — Temporal fossae of skull completely roofed by bone, the
surface of which is granulated, rising from the frontals and
parietals and joining the centre of the jugals; extending backwards from jugal
independently of and above the posterior portion of zygoma. Orbit thus ren-
dered very small. Under these bony plates it may be seen that the skull is
Fig. 180. LoPHio.MYs i.mhalsi imhavsi, Milne-Edwards.
B.M. No. 26. 5. 12. 100, 9; X ij.
constricted between the orbits, as in many Muridae. Paroccipital process rela-
tively long; occipital region prominently ridged. The granulations of the skull
usually extend on to the occipital region, and on the nasals and sides of rostrum.
Bullae very small. Incisive foramina long, extending to anterior molars. Hamu-
lar processes high and raised; palate with rather large lateral pits between
posterior molars, and mesopterygoid space u.sually wide. Zygomatic plate of
specialized Murine type; tilted prominently upwards, well ridged on its superior
Fig. I is I. LoPHlOMYs i.mhalsi imhausi, Milne-Eduards.
B-M. No. 26.5.12.100, -; ,-. i^.
Fill. 182. LoPHiOMYS IMHAUSI IMHAI'SI, Milne-Edwards.
Cheekteeth: B.M. No. 26.5.12.100; 5.
LOPHIOMYS 635
border. Infraorbital foramen wider above than below, its outer side usually
ridged. Mandible with hinder portion of angular process rather flat. Coronoid
process well developed.
Incisors moderately broad. Cheekteeth decreasing in size from M.i to
M.3; three laminae on M.i, two on the other teeth, each lamina formed by an
outer and inner cusp, the cusps approximately equal in size to each other;
each lamina separated from the one behind it by a broad outer and inner re-
entrant fold, these folds separated from each other by a narrow ridge running
down the centre of the tooth, the general effect much as in the genus Cricetus;
the cusps of each lamina separated from each other by a furrow-. Teeth, ex-
cepting the raised cusps and longitudinal ridges, coloured black. Anterior
portion of M.2 w4th a small inner and outer fold in front of the two anterior
cusps. Lower teeth much like those of the upper series; the two anterior cusps
on M.I rather small; the furrow separating the two posterior cusps tending to
take the form of an inner re-entrant fold.
Size rather large for a .Muroid Rodent, head and body to 360 mm. Forefoot
with four well-developed digits, of which D.3 is slightly the longest, and a rudi-
mentary pollex. Hindfoot rather broad, with six plantar pads, three central
digits slightly longer than the hallux and D.5, which are more or less subequal.
The hallux is opposable, though clawed, and in appearance much less specialized
than certain Indo-Malayan climbing Rats as Chiropodomys, Hapalomys, etc.
Tail very thickly haired throughout its length; shorter than head and body,
and terminating evidently in a small knob, which I have seen in captivity
specimens used as an aid to climbing wire-netting. Ear relatively small. Fur
excessively thick, the central portion forming an erectile crest ; when this is lifted
up, a long more or less bare patch of skin running along the sides is to be seen in
dried skins, and in the living animal. In the young animal, it is much more
evident than in the adult, perhaps owing to the fact that the coat, even at ten
weeks old, is scanty. Clavicles imperfect. Dorsal vertebrae more numerous
than is usual; fifteen pairs in a skeleton in my possession.
Some time ago I was fortunate enough to obtain some of these interesting
animals, some of which I gave to the London Zoological Gardens, and some of
which were kept by a friend of mine. They lived quite well for a time, and,
moreover, my friend bred and reared a single young one which lived for fourteen
months. The diet was the chief difficulty in keeping these animals alive, and
the problem was not solved. They seemed to thrive best in a temperature of
about 60 degrees. They were rather strictly nocturnal, and at first not easv to
tame; but when placed in a large cage and given a hollow log to sleep in and
plenty of climbing facilities they soon got to know us, and would come to call
and feed from hand in the evening. I have seen it stated that these animals
cannot climb; my impression is that they are the most perfect and natural
climbers. When provided with a cage about six feet high they lost no time in
climbing to the top of it, and they always climb down vertical wire-netting head
downwards and front feet first; even the young one which was bred doing
this rather astonishing feat quite naturally; moreover, they seemed able
to swarm up concrete for a short distance. On the whole they were ver\-
636 LOPHIOMYS— SPALACIDAE
slow-movinj; animals, and it is rather a mystery how they have managed
to survive.
The remarkable skull structure seems to make it desirable to refer these
animals to a distinct family; though perhaps derived from Cricetine Muridae
I think that sometimes there are more important features to be taken into
account in classification than similarity of cheekteeth alone.
Forms seen : bosasi, hiiidei, ibeamts, iui/niiisi, Icstudo.
Thomas evidentlv came to the conclusion that all the East African "species"
were one, as there is a note in his tracts to this effect. I am inclined to go
further and think that until more material comes to hand all forms must be
treated as races of the earliest name imhaiisi. I believe there would be many
differences which could he regarded as individual or sexual, in a large series.
Though I have seen very few alive, I must note that in every case the females
seem to be larger than the males.
List of Named Forms
(References and tvpe localities of the Lophiomyidae have been collected by
Mr. R. W. Hayman.j
1. LOPHIOMYS IMHAISI IMHAUSI, Milne-Edwards
1867. L'Institut, vol. 35. p. 46.
Somaliland.
Syncjnym: siiiitlii. Khoads, 1896, Proc. Acad. Nat. Sci. Philadelphia,
p. 524. Somaliland.
huzasi, Oustalet, 1Q02, Bull. Mus. Hist. Nat. Paris, p. 400.
Goba. S. Abyssinia.
2. LOPHIOMYS IMHAUSI AKTHIOPICUS, Peters
1867. Zeitschr. Ges. Nat\ir. XXIX, p. 195.
Near Kassala, Sudan.
(A synonym of L. ;'. imhausi according to G. M. .^llen, 1939.)
3. LOPHIOMYS IMHAUSI IBEANUS, Thomas
1910. Ann. Mas. Nat. Hist. 8, VI, p. 223.
Mile 513 on Uganda Railway, between Londiani and Lumbwa Stations,
Mau region, Kenya.
4. LOPHIOMYS IMHAUSI HINDEI. Thomas
1910. Ann. Mag- Nat. Hist. 8, VI, p. 223.
Mutaragwa, Aberdare Mountains, Kenya.
5. LOPHIOMYS IMHAUSI TESTUDO, Thomas
1905. Ann. Mag. Nat. Hist. 7, XV, p. 80.
Ravine Station, Kenya.
h. LOPHIOMYS IMH.AUSI THOMASI, Heller
1912. Smiths. Misc. Coll. LIX, no. 16, p. 4.
Mount Gargucs, Matthews Range, Kenya.
Family SPALACIDAE
1S96. Thomas: Mvomorpha, part: Family Spalacidae, part. Subfamily Spalacinae.
1899. Tullberg: Mvomorpha, part: Family Spalacidae, part, included Myospalax
{" Siphneus"). Rliizowys. Tacliynryctes.
SPALACIDAE 637
1Q18. Miller & Gidley : Muroidae, part: Family Spalacidat, part, Subfamily Spalacinae.
:924. Winge: Family Dipodidae, part, Spalacini.
1928. Weber: Muroiuea: Family Spalaeidae, part, included IViizomys, Myospalax,
Tachyoryctes.
Geographical Distribution. — Palaearctic : Eastern Mediterranean region:
Galicia, Hungary, Roumania, Yugoslavia,
Turkey, Greece, Southern Russia to the Caspian Sea (Poltava, Harkov, Dnepro-
petrovsk, Voronej, Saratov, Don, Stalingrad districts, Ciscaucasia, Trans-
caucasia) (Vinogradov); Asia Minor, Syria, Palestine, North Egypt, into Libya.
Number of Genera. — One.
Characters. — (As here understood the family contains the genus Spalax
only.) Skull and external form extremely modified for sub-
fossorial life. E.xternal eyes suppressed (in this character unique among
Rodents). Tail absent; ear vestigial. Claws not specially enlarged. Skull
with supraoccipital region sloping forwards to level of the posterior zygomatic
root, this region occupying a third or more of whole length of skull. Zygomatic
plate relatively narrow, and nearly completely beneath the infraorbital foramen,
which is large for a Muroid Rodent; masseter lateralis superficialis with anterior
head distinct from zygoma (as figured by Tullberg), as in normal Muroid
Rodents. Dental formula i. \, c. g, p. g, m.| = 16. Cheekteeth rooted, the inner
and outer re-entrant folds forming the pattern of the young animal soon becom-
ing isolated on crown surface. Lower incisor forming powerful process on
mandible beside the condyle.
Remarks. — As will be seen above, the family has in most previous classi-
fications contained several other genera, such as Myospalax,
Rhizomys and Tachyoryctes. Miller & Gidley show in their classification of the
Order that the zygomasseteric structure of Rhizomys is totally different from
that of Spalax, indeed the two groups being at the opposite end of two extremes ;
Rhizomys and Tacltyoryctes were accordingly referred to a family Rhizomyidae
by these authors, while Spalax and Myospalax were retained in the Spalaeidae.
But the genus Myospalax is very much less highly modified as regards the
arrangement of zygomatic plate and infraorbital foramen than Spalax, if the two
are compared with a normal member of the Muridae. Their cheekteeth also
are very different; and most authors are agreed in placing Mvospalax in the
Muridae, where its position seems to be in the neighbourhood of the Cricetinae
(as indicated bv Winge), or the Microtinae. Such superficial resemblance as
exists between Spalax and Myospalax cranially, such as the occipital region
which is abnormally sloped forwards in both, is probably brought about by the
similar mode of life which the two genera lead.
Winge regarded Rhizomys, Tachyoryctes and Myospalax as Muridae, but
transferred Spalax to the Dipodidae. But according to Tullberg the stomach of
Spalax agrees with the Muridae rather than the Dipodidae; and the infra-
orbital foramen and zygomatic plate of Spalax are certainly not Dipodoid.
It should be mentioned that the family Spalaeidae of earlier authors appeared
to contain all the Old World Rodents which live underground, and even formerly
638 SPALAX
included the Bathyergidae ! I do not think that there is much doubt that the
grouping together of Spalax, Mvospalnx, Tachvorvctes and Rliizonivs is a \-ery
unnatural arrangement.
Genus I. SPALAX, Guldenstaedt
1770. Spalax, Guldenstaedt, Nov. Com. Acad. Sci. Imp. Petrop. XIV, pt. i, p. 410.
IQOQ. Macrospalax, Mehely, A Foldi Kutyak Fajai, Budapest, p. 23. (New name for
Spalax s.s.)
1909. Mesospalax, Mehely. A Foldi Kutyak Fajai, Budapest, p. 22. (Based on Spalax
moNticola, Nehring, and S. luingaricus. No type designated. If the type has not
already been chosen I here choose the fonner as type species.)
1898. Microspal.ax, Nehring, S.B. Ges. Nat. Fr. Berlin, p. 168, for December 1897.
" Smaller species of Spalax." Not of Trouessart.
1903. Nannospal.w, Palmer, Science, new ser., XVII, p. 873. (To replace Uliaospalax,
Nehring. If the type has not heretofore been designated I choose Spalax kirgisorum,
Nehring.)
1922. UjHELVi.'iNA, Strand, Arch. Naturg. Berlin, 88, .Abt. A. Hft. 4, p. 142. To replace
Microspalax, Nehring.
Type Species. — Spalax iiiiciophthahinis, Guldenstaedt.
Range. — As in the Family Spalacidae.
Number of For.ms. — About twenty-seven.
Char.^cters. — Skull extremely fossorial; occiput high and broad, slanting
forwards to level of the posterior zygomatic root. This region
of skull roofs over the space between the auditory meatus and the posterior
zygomatic root. Frontals extremely constricted. Rostrum long and heavy.
Jugal short, reduced. Incisive foramina small, far in front of toothrow. Bullae
of medium size. Zygomatic breadth considerable, greatest near the posterior
zygomatic root. Foramen magnum less than half the height of occipital shield.
Palate narrow. Infraorbital foramen relatively large, higher than wide. Zygo-
matic plate narrow, nearly completely beneath it. The ascending branch of the
zygoma outside the infraorbital foramen is in some cases broader than the
zygomatic plate. But compared, for instance, with Jaciiliis representing the
Dipodidae, to w'hich the genus was transferred by Winge, the infraorbital
foramen is in this case very much less enlarged, and the zygomatic plate is rather
more IVlurine in general appearance. A small protuberance on the lower part
of the inner side of the infraorbital foramen is present, and might mark the
division between the muscle-transmitting portion and the nerve-transmitting
portion. Mandible with the lower incisor root forming a large process between
the condylar and the top of the angular portion, and projecting considerably
outwards and upwards above these bones; coronoid process high and slender;
angular portion low, rather spread sideways.
Incisors broad and heavy, the root of the upper teeth forming a slight
knob in the side of the palate in front of M.i. Molars small, semihypsodont.
M.I not much larger than M.2 and M.3 little smaller than this tooth. Good
figures of the unworn teeth are supplied by Mehely; these show that as usual
in the Order the very young teeth are extremely complex. At a later stage,
inner and outer re-entrant folds form the tooth pattern, in which m M.i and
Fig. 183. Spalax monticola dolbrogeae, Miller.
B.M. No. 5.10.25.2, ¥; x ij.
41 — Living Rodents — I
640
SPALAX
M.2 there are usually two baekwardly curved outer and one lorwardly curved
inner folds. The folds isolate as enamel islands, the second outer fold being the
most persistent. I\1.3 varies in pattern; in the subgenus Nannospalax there are
ahvavs two isolated more or less parallel enamel islands; in the subgenus Mcso-
spalax there is one, the centre of which is joined as a rule by a short island at
right angles to it. Typical Spalax appears to agree with Mesospalax. M.3 in
voung XiDinospalax is S-shaped, formed by one inner and one outer re-entrant
fold, and M.2 is more or less similar, when very young, but later seems to take
on a pattern more like M.i or as described above.
Fig. 184. Spalax monticola dolbrogeae, Miller.
Cheekteeth showing enamel pattern at various stages of wear. Upper row,
maxillan,' teeth; lower row, mandibular teeth (semi-diagrammatic) ; ■ 5.
(From Miller's Catalogue of the Manimals of Western Europe.)
In each ot the lower teeth there are in Mesospalax usually one inner and one
outer fold in moderately worn specimens. In Nannospalax there appear to be
generally two inner folds on the two front teeth; the islands generally in this
subgenus are three on M.i, two on the other teeth; the other subgenera often
have only two islands on M.i.
Form Mole-like. Head round. Hat. External eyes absent; they are said to
be more or less developed but quite functionless beneath the skin. Ears vesti-
gial. Tail absent. Forefoot with five digits, the pollex very small, otherwise
proportions of digits not unlike that of a human hand. Hindfoot digit propor-
tions much as in forefoot. Claws medium, not specially developed, the digging
being done apparently with the incisors. A line of stiff bristles running across face
from the nostrils about half-way to the ear present, otherwise fur very short, soft.
SPALAX
641
The genus was thoroughly monographed by Mehely, Species Generis
Spalax, 1909, A Foldi Kutyak I'ajai, Budapest.* This author divided the genus
into three subgenera, Mesospalax, Macrospalax, and Microspalax. Macruspalax
is a "new name for Spalax s. str," and is therefore unnecessary and must be
placed in synonymy. Microspalax is preoccupied, and has been renamed twice;
consequently the nomenclature of the genus is somewhat complicated.
'l"he three valid subgenera are at the present time Spalax (synonym Macro-
spalax, Mehely); Mesospalax; and Nannospalax (synonyms Microspalax,
Ljhelyiana).
Their main differences pointed out by IMehely are below.
Subgenus
Nannospalax
Subgenus
Mesospalax
Subgenus
Spalax
Length of Skull .
42-45 mm.
47-54 mm.
53-74 mm.
Supraoccipital
Shorter
Shorter
Very long
Foramen
supracondyleum
Present
Present
Absent
Petromastoideum .
Short and broad
Short and broad
Longer
External auditory
Wider and
Wider and
Narrow
meatus
longer
longer
Pterygoid fossae .
More open
More open
More closed
Condyle
Little lower than
Considerably
Considerably
incisor knob
lower than in-
lower than in-
cisor knob
cisor knob
Angular process .
Spread out from
body of lower
Spread out from
body of lower
Xtry slightly re-
moved from
jaw
jaw
body of lower
jaw
M-3 • • •
With two ena-
One enamel
One enamel
mel islands
island
island
M.I and M.2,
Two lingual
Two lingual
One lingual
chewing surface
folds
folds, the
posterior one
early replaced
by an island
enamel fold
^ German edition in Math.
text and Atlas.
Naturwiss. Ber. Ungarn. 88 (1910). Leipzig 1913. pp. i-390.
642 SPALAX
This table briefly summarizes the main characteristics pointed out by Mehely.
The material examined does not show much difference in the characters of
pterygoid fossae, condyle and angular process; but very few skulls are available
of Spiilax s.s.
Forms seen: (legvptiaciis, anatoUcus, captoniin, coiyhaiitiiim, dolbrogetie,
threnbergi, hellenicus, hungaricus, insularis, microphthabmis, nehringi, serbiciis,
thermaicus, transsvhonicus.
S. giganteiis, not seen, is said to differ considerably from other members of
subgenus Sptilax in cranial characters (broadened rostrum, etc.).
List of Named Forms
(References and type localities for Spalacidae are the work of Mr. R. W.
Ilayman.)
Subgenus Natmospalax, Palmer
(All members of this group are regarded as one species by Mehely under the
name ehrenbcrgi \ but kirgisorum has page priority, and so must be used.)
1. SPAI.AX KIRGISORUM KIRGISORUM, Nehnng
1898 (for 1897). Sitz. Ber. Ges. Nat. Fr. Berlin, p. 176.
Kirshiz Steppes. (? no Spalax of this type quoted by Vinogradov in
Rodents of U.S.S.R.; according to Mehely comes from North
Syria.)
Synonym: inlermedius, Nehring, i8(>8, Sitz. Ber. Ges. Nat. Fr. Berlin,
Dec. 1897, p. 181. Syria.
berytensis. Miller, 1903, Proc. Biol. Soc. Washington, XVI,
p. 162. Beyrout, Syria.
2. SPALAX KIRGISORUM EHRENBERGI, Nehnng
1898 (for 1897). Sitz. Ber. Ges. Nat. Fr. Berlin, p. 17S.
Jaffa, Palestine.
3. SPALAX KIRGISORUM AEGYPTIACUS. Nehnng
1898. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 180.
Ramleh, North Egypt.
Subgenus Mesospalax, Mehely
4. SPALAX MONTICOLA MONTICOLA, Nehrini;
1898. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 6.
Kupres, Bosnia, Yugoslavia.
5. SPAL.AX MONTICOLA NEHRINGI, Satunin
1898. Zool. Anz. XXI, p. 314.
Kasikoporan, Armenia.
6. SPALAX MONTICOLA ARMENIACUS, Mehely
1909. A Foldi Kutyak Fajai, Budapest, p. 79.
Kura-Quellan, .Armenia.
7. SPALAX MONTICOL.A CILICICUS, Mehely
1909. A Foldi Kutyak Fajai Budapest, p. 84.
Cilician Taurus, .Asia Minor.
SPALAX 643
8. SPALAX MONTICOLA ANATOLICUS, Mfihcly
IQ09. A Kcildi Kutyak Kajai, Budapest, p. 88.
liurnabad, near Smyrna, Asia Minor.
9. SPALAX MONTICOLA IIKLLENICUS, Mdhely
iQog. A Foldi Kutyik Fajai, Budapest, p. 100.
Lamia. Thessaly, Greece.
10. SPALAX MONTICOLA TURCICUS, M^hely
1909. A Foldi Kutyak Fajai, Budapest, p. 105.
Makri-Koi, Constantinople, Turkey.
11. SPALAX MONTICOLA DOLBROGEAK, Miller
1903. Proc. Biol. Soc. Washington, XVI, p. 161.
Dobrudscha, Roumania.
12. SPALAX MONTICOLA HERCEGOVINENSIS, M^hely
1909. A Foldi Kutyak Fajai, Budapest, p. 129.
Ulog-Obruga, Herzegovina, Yugo-Slavia.
13. SPAL.\X MONTICOLA SVRMIENSIS, M^hely
1909. A Foldi Kutydk Fajai, Budapest, p. 133.
Szerem, Slavonia, Yugo-Slavia.
14. SPAL.AX MONTICOLA SERBICUS, M^hely
1909. A Foldi Kutyak Fajai, Budapest, p. 140.
Serbia.
15. SPALAX MONTICOLA INSULARIS, Thomas
1917. .Ann. Mag. Nat. Hist. 8, XX, p. 315.
Isle of Lemnos, Greece.
16. SPALAX MONTICOLA THERMAICUS, Hinton
1920. Ann. Mag. Nat. Hist. 9, V, p. 313.
Salonica, Greece.
17. SPAL.-VX MONTICOLA CAPTORUM, Hinton
1920. .Ann. Mag. Nat. Hist. 9, V, p. 318.
Changria, Asia Minor.
18. SPALAX MONTICOLA CORYBANTIUM, Hinton
1920. .\nn. Mag. Nat. Hist. 9, V, p. 316.
One hundred and fifty miles east of Smyrna, .Asia Minor.
19. SP.AL.AX MONTICOLA LAB.AUMEI, Matschie
1919. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 35.
Eskischehir, Asia Minor.
20. SP.ALAX HUNGARICUS HUNGARICUS, Nehring
1898. Sitz. Ber. Ges. Nat. Fr. Berlin, p. 173.
Hungary.
21. SPAL.AX HUNG.ARICUS TRANSSVLVANICUS, M^hely
1909. A Foldi Kutyak Fajai, Budapest, p. 159.
TranssyKania.
Subgenus Spalax, Guldenstaedt
22. SPALAX GRAECUS GRAECUS, Nehring
1S98. Zool. Anz. XXI, p. 228.
? Neighbourhood of .Athens, Greece.
644 SPALAX— RHIZOMYIDAE
li. SPALAX GRAECLS ANTIQUUS, Mihcly
irioq. A Foldi KutN'ak Fajai, Budapest, p. 175.
Roumania.
24. SPALAX ISTRICCS. Mihely
iqog. A Foldi KuUak Fajai, Budapest, p. 186.
Barza, Roumania.
25. SPALAX POLONICUS, M^hily
iQog. A Foldi Kut\'ak Fajai, Budapest, p. 194.
Lemberg, Galicia.
26. SPALAX MICROPHTHALMUS, Guldenstaedt
1770. N. Conim. Ac. Sci. Imp. Petrop. XIV, pt i, p. 411.
Steppes of Nobochopersk, S. Russia.
Synonym: typhhis, Pallas, i77iS, Nov. Sp. Quadr. Glir. Ord., pp. 76,
154, pi. S. S. Russia.
podoUca, Pennant, 1771, Synop. Quadr. p. 277.
xanthodon, Nordmann, 1S40, DemidofF Voy. I, p. 35, pi. II.
leiicodou, Nordmann, same reference, p. 34.
pallasii, Nordmann, Bull. Ac. St. Petersb. 1835, p. 200.
27. SPALAX GIGANTF.US, Nehring
1898. Sitz. Ber. Ges. Nat. Fr. Berlin (for 1897), p. 169.
Petrovsk, Caspian Sea, Russia.
The family is known fossil from the Upper Oligocene, according to Miller
& Gidley, hut only from the range given above.
SPALACIDAE:
GENERAL WORKS OF REFERENCE
Mehelv, 1909, Species Generis Spalax: A Foldi Kutyak Fajai, Budapest.
Miller, 1912, Catalogue Mammals Western Europe: Spalacidae, p. S87. (The "Spalax
graecus" of this work is not graeciis as understood by Mehely, but monticola
hellenicus; see Thomas, 1917, Ann. Mag. Nat. Hist. 8. XX, p. 317.)
Family RHIZOMYIDAE
1 8g6. Thomas : Myo,\iorph.\, part : Family Spalacidae, part, subfamily Rhizomyinae,
part included Tachyoryctes.
1 899. Tullberg: MvoMORPH.^: Family Spalacidae, part.
1918. Miller & Ciidley: MvROiD.\E : Family Rhizomyidac, part, subfamily Rhizomyinae.
1924. Winge: Family Muridae, part: Rhizomyini, part, included Tachyoryctes and all
genera of Muridae from Madagascar.
1928. Weber: Mlroide.\: Family Spalacidae, part.
Gf-OGRAPHICVl Distribution. — Indo-Malayan region from China south
of the Yangtsekiang (ranging north to
Szechuan), and from the Eastern Himalayas (Nepal) south through Siam and the
Malay Peninsula to Sumatra.
Number of Gkner.'\. — Two are here retained.
RHIZOMYIDAE 645
Characters. — Zygomasseteric structure more specialized than in the
Muridae; zygomatic plate tilted verj- strongly upwards;
infraorbital foramen much reduced, its lower border nearly straight (instead of
V-shaped as is very general in Muridae), and situated on upper border of zygo-
matic plate, its neutral portion obliterated, the foramen usually broader than
high. Skull modified for fossorial life; dental formula i. |, c. g, p. g, m. J = 16;
cheekteeth semihypsodont, rooted, flatcrowned, characterized by a pattern of
inner and outer re-entrant folds which become isolated on crown surface as
islands; external form moderately specialized for fossorial life.
The systematic position of the Rhizomyidae is not clear ; by many authors the
group has been referred to the Spalacidae, but I have already pointed out when
dealing with that family that the zygomasseteric structure as regards arrange-
ment of zygomatic plate and infraorbital foramen is very distinct in the two
groups. Thomas, 1896, remarked, " It is doubtful whether Spalax and Rfiizomys
. . . are rightly put even in one family, their resemblances being perhaps more
adaptive than generic." Forsjth Major regarded Rhizomys, also Spalax and
Tachyoryctes (the former family Spalacidae), as primitive .Muridae; though in
many respects these genera seem to me to be more highly specialized than
is usual in the Muridae. Winge transferred Rhisomys to the Muridae, and
formed a subfamily or group Rhizomyini including Rhizomys, Tachyoryctes,
Brachytarsomys, Brachyuromys, Eliurus, Nesomys, and Gymmiromys, based
fundamentally on the character that M.i is not or scarcely larger than
M.2, whereas in all other Muridae he states that M.i has become larger
than M.2. But I have not found this the case, as I shall show when dealing
with the Muridae. Miller & Gidley, 1918, formed a family Rhizomyidae,
defined as "Like the Cricetidae, but zygomasseteric structure unusual, the
infraorbital foramen with neural portion reduced or obliterated by partial
or entire fusion of zygomatic plate with side of rostrum." This family
included Tachvoryctes, and also Bramus, a Pleistocene North African fossil
genus which has since been shown by Hinton to be a synonym of the Microtine
genus Ellobius.
Tachyoryctes has a much less specialized and abnormal infraorbital foramen
than Rhizomys and Cannomxs, and differs from them conspicuously in dental
characters. I do not think Tachyoryctes is so closely allied to the RJiizotnvidae
as maintained bv most authors. The genus is here referred to the Muridae, as
its infraorbital foramen is clearly not highly abnormal, and as figured bv Tullberg
differs widely from the Rhizomyidae ; whereas its cheekteeth are similar in pattern
to one of the Rats of Madagascar {Brachyuromys),\\h\ch. has alwavs been regarded
as a member of the Muridae.
The Rhizomyidae are here kept apart from the Muridae solely on account of
their peculiar zvgomasseteric structure, the masseter muscle as figured by Tull-
berg rising up the zygomatic plate inside the infraorbital foramen, a condition
so far as I know without parallel elsewhere among Muroid Rodents.
Thomas divided Rhizomys into three genera, Xyctocleptes, Rhizomys and Can-
nomys. But intermediate forms are now known between XyctocleptesAnil Rhizomys,
and the two groups are probably more correctly referred to a single genus.
646 RHIZOMYIDAE: RHIZOMYS
Key to the Genera of Rhizomvidae
Upper incisors strongly pro-odont; sole pads normal, not granulated. M.i
never worn below level of M.2, and the largest tooth in the upper
series. C.-vnnomys
Upper incisors not strongly pro-odont; sole pads granulated; M.i often
smaller than M.2, and often worn below the level of the latter.
Rhizomys
Genus i. RHIZOMYS, Gray
1 83 1. Rhizomys, Gray. Proc. Zool. Soc. London, p. 95.
1832. NvcTOCLEPTES. Temminck, Bijdragen Nat. Wetensch. .Amsterdam, VII, p. 7,
pi. I. (Mils sumatremis, Raffles.) Valid as a subgenus.
Type Species. — Rhizomys sinensis. Gray.
R.\nge. — Moupin, Assam, Burma, Tenasserim, Yunnan, Kwantung, Fukien,
Siam, Indo-China, southern part of Malay Peninsula, Sumatra.
NfMBER OE Forms. — Thirteen.
Characters. — Frontals much constricted; behind them in larger species a
strong sagittal ridge is formed which extends to lambdoid crest.
Occipital region upstanding and prominent, rather sloped forwards, though not
excessively so. Paroccipital process relativelv long; zvgomata widely spreading;
rostrum hea\T. Jugal thick and prominent, but most of anterior part of zygoma
formed by zygomatic process of maxillae. Bullae large, the meatus directed
outwards and upwards; the bullae appear conspicuously in back view of skull
between the paroccipital process and the upper process of supraoccipital. Palate
narrow; posterior nares typically compressed, higher than wide, but more open
in R. senex and in subgenus Nyctocleptes. Zygomatic plate tilted very strongly
upwards; infraorbital foramen reduced to a small orifice situated on its upper
border, broader than high, its lower border as a rule nearly straight. Incisive
foramina small, considerably in front of toothrows. Mandible with prominent
knob caused by lower incisor root, nearly as high as the condylar process.
Coronoid process high.
Incisors very broad, orthodont, or slightlv pro-odont in Nyctocleptes. Cheek-
teeth semihypsodont; M.i in subgenus Rhizom\s smaller than M.2, consider-
ably so in old individuals, and worn considerably below its level. M.i with
one inner fold (more persistent), and three outer folds which usually isolate as
islands. M.2 when cut appears to have one inner fold which cuts the tooth into
two lobes; the inner fold breaks in two and forms a fold in the anterior lobe;
a deep outer fold cuts across the posterior lobe; with wear this simplifies to a
pattern of three external, one deep internal enamel islands. M.3 appears to vary
to a degree individually, but is not very ditferent from M.2, and simplifies in
a similar manner with wear. Lower series: M.i not very much smaller than
M.2; usually three enamel islands present in the first two molars; M.3 appears
to consist of two lobes, the anterior of which is the larger and has in the centre
Fig. 185. Rhizomys pruinosis pruinosus, Bl>-th.
B.M. No. 20.1 1. 1. 44, ?; X li.
Fig. 186. Rhizomys pri inosis pruinosus, Blyth.
B.M. No. 20. 1 1. 1. 44, t; :< ij.
Fig. 187. Rhizomys pruinosus pruinosus, Blyth.
B.M. No. 20.11. 1.44, +; X ij.
Fig. 1S8. Rhizomys pruinosus pruinosus, Blyth.
Front view of skull: B.M. No. 20.11. 1.44, V; >'• i*-
RHIZOMYS
649
a large isolated island. In the subgenus Nyctocleptes, M.i upper is not smaller
than .M.2, but may be worn down to a lower level, as in typical RJtizomys.
Form more or less fossorial; fur very thick in the northern forms, becoming
harsh and scanty in the tropical species. Eyes and ears small. Tail varying
from about a third or more length head and body to little longer than hindfoot,
not well haired. Claws rather prominent. Forefoot with D.3 longest, D.2 a
little shorter but slightly longer than D.4; D.5 shortest. Proportions of digits
of hindfoot nearly as in toretoot, but D.2 may be relatively longer, and hallu.x
nearly as long as D.5. Mammae normally i — 3 = 8, though occasionally a minute
Fig. 189. Rhizomys pkuinosus pruinosus, Blyth.
Cheekteeth: B.M. No. 20.11.1.44, ?» ^ 5.
anterior pectoral pair may be present as well as these, and in subgenus .V3C/0-
cleptes, 2 — 3 = 10 (Thomas).
Nyctocleptes was revived as a fullgenusby Thomas, the differences being the
largersize (nota genericcharacter), the infraorbital foramen oval or circular instead
of subtriangular, but some specimens seen of R. senex appear in this formation
indistinguishable from Nyctocleptes; the posterior nares well open, often nearly
as wide as high (but R. senex is intermediate between the two groups in this
character) ; the set of the incisors more pro-odont (but not approaching Cannomys
in this character), and M.i not smaller than M.2, and rarely worn to a lower
level. The two posterior solepads are joined (separate in Rhizomys). Also
there is a strong ridge each side of the front of the palate which extends forwards
to the prema.\illae, which is more developed than in Rhizomys. But these
650 RHIZOMYS
ditferences seem suhgeneric rather than generic. The fur is harsh and short,
and the size becomes very large, up to 450 mm. head and body, or more. The
tail is nearly naked, and longest of genus. Hindfoot S3-(>5 mm. All named
forms appear to belong to one species only.
Subgenus RmzoMYS appears to me on the material examined to divide into
two groups, and six species, as follows:
Sagittal ridge always strongly developed; occipital region of skull appearing
higher; general colour grey; fur excessively thick; size at full development largest
of subgenus. R. rt'stiliis group. From Szechuan, Fokien, and North Burma.
Includes vestitiis (hindfoot 48-53), and the much smaller r/(H7(// (hindfoot 38-44).
Sagittal ridge weak, or maybe not developed in adult (i.e. the ridges may not
fuse); occipital region appears lower; fur not or less excessively thick. K.
sinensis group. R. sinensis, a little-known form from South China (hindfoot 46),
has thicker fur than the remainder referred to the group, and is greyer in colour;
the supraorbital ridges join. The remainder are browner in general coloration,
and often the supraorbital ridges do not join; R. senex has the posterior nares
well open, as in Nxctocleptes (hindfoot 44; Yunnan); R. pruinosus has the pos-
terior nares narrow (normal) (hindfoot 46-50; Assam, Yunnan). These two
species have moderate fur; in R. pannosus the fur is short and harsh, almost
exactly like that of Nxctocleptes (hindfoot about 46; Perak, Siam).
Forms seen : cinereus, davidi, insidaris, hitouchei, pannosus, pruinosus, senex,
sinensis, sunicitrensis, unibriceps, restitus, u'ardi.
List of Named Forms
(References and type localities for all Rhizomyidae are the work of Mr. R. W.
Hayman.)
Subgenus Rhizomys, Gray
vestitus Group
1. RHIZO.MYS DAVIDI, Thomas
191 1. ,\bstr. Proc. Zool. Soc. London, 90, p. 5; Proc. Zool. Soc. London, 191 1, p. 179.
Kuatun, N.-W. Fukien, S. China.
2. RHIZOMYS VESTITUS VESTITUS, Milne-Edwards
1871. Nouv. Arch. Mus. Nat. Hist., VH, Bull. p. 92.
Moupin, Szechuan.
,-;. RHIZOMYS VESTITUS WAKDI, Thomas
1 92 1. Journ. Bombay Xat. Hist. Soc. XXVH, p. 504.
Imaw Bum, Kachin Province, N. IJurma.
sinensis Group
4. RHIZOMYS SINENSIS, Gray
1S31. Proc. Zool. Soc. London, p. 9i.
S. China.
5. RHIZOMYS SENEX, Thomas
1915. Ann. Mag. Nat. Hist. 8, XVL p. 313.
S'unnan, S. China; probably neighbourhood of Mongtze.
6.
i85i.
7-
1915-
8.
1915-
9-
1916.
RHIZOMYS— CANNOMYS 651
KHIZOMYS PRUINOSUS PKUINOSUS, Blj-th
Journ. Asiat. Soc. Bengal, XX, p. 519.
Churrapunji, Khasia Hills, Assam.
RHIZOMYS PRUINOSUS LATOUCHEl, Thomas
Ann. Mag. Nat. Hist. 8, XVI, p. 59.
Quantung, S. China.
RHIZOMYS P.-^NNOSUS PANNOSUS, Thomas
Ann. MaR. Nat. Hist. 8, XVI, p. 60.
Chantabun, S. Siam.
RHIZOMYS PANNOSUS UMBRICEPS, Thomas
Ann. Mag. Nat. Hist. 8, XVIII,, p. 445.
Pcrak, Malay Peninsula.
Subgenus Nyctocleptes, Temminck
10. RHIZOMYS SUMATRENSIS SUMATRENSIS, Raffles
1822. Cat. Zool. Coll. Sumatra, in Trans. Linn. Soc. London, XIII, p. 258.
Malacca.
Synonym: javaniats, Cuvier, 1829, Regne Animal, I, p. 211.
dekan, Temminck, 1835, Mon. Mamm. II, p. 44, pi. 33.
11. RHIZOMYS SUMATRENSIS PADANGENSIS, Brongersma
1936. Zool. Meded. Leiden, 19, p. 154.
Koto Gadang (Singgalang), Padang Highlands, W. Sumatra.
12. RHIZOMYS SUMATRENSIS INSULARIS, Thomas
1915. Ann. Mag. Nat. Hist. 8, XVI, p. 58.
Deli, Sumatra.
13. RHIZOMYS SUMATRENSIS CINEREUS, MacClelland
1842. Calcutta Journ. Nat. Hist. II, p. 456.
Tenasserim.
Synonym: erythrogenys, Anderson, 1877, Proc. Asiat. Soc. Bengal,
p. 150. Salween Hill Tracts, Burma.
Genus 2. CANNOMYS, Thomas
1915. Cannomys, Thomas, Ann. Mag. Nat. Hist. ser. 8, vol. XVI, p. 57.
Type Species. — Rhizomvs badiiis, Hodgson.
Range. — Nepal, Burma, Siam.
Number of Forms. — Six.
Characters. — Like Rhizumxs, but incisors extremely pro-odont, and
lengthened. Other cranial characters much as Khizomys;
supraorbital ridges evidently joining, normally. Infraorbital foramen sometimes
extremely reduced; in some specimens it is slit-like, and about twice as wide as
high.
Cheekteeth decreasing in size from M.i backwards; M.2 smaller than M.i.
M.I often as Rhizomys in pattern, but the anterior island mav wear out so that
there are only two outer islands. M.2 with two isolated external islands, and
652 CANNOMYS
one inner fold; M.3 as a rule with only two small isolated islands. The enamel
islands as a rule more evident and the pattern more elear and definite than in
Kliizomys.
Lower teeth : in adult there are usually two isolated islands on each tooth,
though in voung specimens a much more complex pattern is visible.
Size as a rule smaller than Rhisomys, usually not exceeding 250 mm. head
and body (hindfoot about 29-36). Fur rather thick. Tail of moderate length,
about twice or more length hindfoot, nearly naked. Digits more or less as in
Rhizomys. Mammae 2 — 2 = 8 (Thomas).
Forms seen : castaneus, badiiis, minor, plumbescens, pater.
I do not think there is more than one valid species in this genus.
List of Named Forms
.. CANNOMYS HADIUS BADIUS, Hodgson
1S42. Calcutta Journ. Nat. Hist, ii, pp. 60, 410.
Nepal.
2. CANNOMYS BADIUR PATER, Thomas
1915. Ann. Mag. Nat. Hist. 8, XVI, p. 315.
Mount Popa, dry zone of Buniia.
3. CANNOMYS BADirS CASTANKUS, Blyth
1843. Journ. Asiat. Soc. Bengal. XII, p. 1007.
(?) Ar.ikan, Burma.
4. CANNOMYS BADIUS PLUMBESCENS, Thomas
1915. Ann. Mag. Nat. Hist. 8, XVI, p. 315.
Gokteik, N. Shan States, Burma.
5. CANNOMYS BADIUS MINOR, Gray
1S42. Ann. Mag. Nat. Hist. X, p. 266.
S. Siam.
6. CANNOMYS BADIUS LONNBERGI, Gyldt-nstolpe
1916. Stockholm Vet. Ak. Handl. 57, no. 2, p. 47.
E. Siam.
The group is known fossil from the Pliocene, from Eastern Asia only.
RHIZOMYIDAE:
SPECIAL WORKS OF REFERENCE
Forsyth Major, on the Malagasy Rodent Genus Brachyuromys, Proc. Zool. Soc.
London, p. 695, 1S97. Rhizomyidae, Spalax, Tachyoryctes and Rodents from
Madagascar fully compared.
All other genera of Rodcntia are here regarded as belonging to the Family
Muridae. A separate volume will be devoted to this tamily.'
' The first volume was compktcd for publiotion on January 27th, 1939. The second (and
last) volume was completed for publication on June 30th, I939-
Addendum to Subgenus Tamiops, Allen (p. 354 above) : —
The forms of the subgenus Tamiops are reviewed by Osgood, Field. Mus.
Nat. Hist. Zool. XVIII, pp. 286-97, '932-
He arranges these in four species, as follows :
Callosciurus swinhoei suinhoei, Milne-Edwards, 1874
C. swinhoei clarkei.
C. swinhoei vestitus.
C. moniicobis monticohis, Bonhote, 1900.
C. monticolus olivaceus.
C. monticolus spencei.
C. monticohis forresti.
C. monticohis russeobis, type locality .Atentze, Tibet.
C. mariiimus maritimus, Bonhote, 1900.
C maritimus formosanus.
C. maritimus liainanus, with synonym riudoni (No. 12 of my list above).
C. maritimus laotum.
C. maritimus moi.
C. tnacleUandi macleUandi, Horsfield, 1839, with (?)synonym manipurensis
(No. 2 of my list, above), "it stands directly between maclellundi and
barbei and seasonal variations in both seem sufficient to cover its
supposed characters."
C. machllandi pembertoni, Blyth ; listed above as a synonym of m. macleUundi.
C. macleUandi barbei.
C. macleUandi novemlineatus.
C. machllandi liantis, with synonym holti. No. 23 of my list above, new name
for Ivlei, Thomas, not of Bonhote.
C machllandi kongensis.
C. machllandi rodolphei. '
C. machllandi dolphoides.
C. macleUandi inconstans.
C. m. sauteri. No. 1 1 of my list, is regarded as of doubtful status,
based on characters which are likely to be seasonal.
References to all these forms will be found on pages 3^4 and 355, except
dolphoides, which is on page 376.
Addendum to genus Callosciurus.
looa. CALLOSCIURUS FERRUGINEUS SPLENUENS, Gray
1861. Proc. Zool. See. London, p. 137.
S. Cambodia.
(omitted from p. 361 in error).
653
INDEX
abassensis, Htliosciurus, 402
abbotti, Ca!losciurus, 370
abbotti, Thomomys, 512
abcrrans, Cryptomys, y2
abcrti, Sciurus, 324, 325, 327, 328, 337
abessinicus, Xerus, 420
abieticola, Tamiasciurus, 347
abictorum, Napacozapus, 574
ablusus, Citellus, 44S
ABROCOMA. 26. <?q. 7^, 102, 103. 152
ABROCOMINAH. 26. 101, 103, 151
abrutti, Glis, 623
absonus, Thomomys, 516
abstrusus, Thomomys, 513
abyssinicus, Allactaga, 584
ACANTHION. 209, 212, 213, 214. 217
ACOMYS, 3, 36, 38, 48, 50, 56
ACONAEMYS, 23, 26, 55, 73, 102, 155, i s6,
157, i6r, 554
acontion, Alactagulus, 587
acrirostratus, Thomomvs, 514
ACROBATES. 543
acticola, Hehosciurus, 401, 403
acouchy, IMyoprocta, 196
adamsi, Callosciurus, 372
adamsoni, Dremomys, 381, 382
adangcnsis, Callosciurus, 3(15
ADELPHOMYS. 134
ADJIDAUMIDAE, 14
adolphei, Sciurus, 336
adocetus, Citellus, 451
adsitus, Tamias, 433
adspersus, Liomys, 477, 478
aedium, Plapiodontia, 134
at-jij-ptiacus, Spalax, 642
acpv'ptius, Jaculus, 596
AEROMYS. 30, 64. 274, 275, 276, 290
acrula, Hystrix, 217, 220
aestivus, Perognathus, 485
aestuans, Sciurus, 326, 328
at'stuans proup, Sciurus, 327, 342
acthiopicus, Lophiomvs, 636
AETHOGLIS, 604, 606, 607
AETHOMYS. 38
AETHOSCIURUS, 264, 267, 399, 404
AETHURUS, 546
affinis, Callosciurus, 355
affinis, Coendou, 186. 188
affinis, Heliosciurus, 404
affinis, Paraxerus, 407
affinis, Ratufa, 384, 387
affinis, Tamias, 432
africacaustralis, Hystrix, 148, 200, 213, 216,
217, 219
africaeaustralis proup, Hvstrix, 212
AFRICAN rodents; 66, 67, 68, 69
4^ — Living Rodents — I
africanus, Athcrurus, 199, 206, 208
africanus group, Atherurus, 208
africana, BathvL-rgus, 84
africanus, Xerus, 422
agadius, Xerus, 420, 421
ayilis, Dipodomys, 496, 497
agilis group, Dipodomvs, 497, 503
ACJOUTI. 221
agrestis, Thomomys, 522
agricolaris, Thomomys, 514
aguti, Dasyprocta, 191, 193
aharonii. Hystrix, 219
airensis, Jaculus, 595, 596
akka, Funisciurus, 412, 413
AKODON. 39, 55
alacris, Callosciurus, 350, 357
alactaga, Allactaga, 586
ALACTAGULUS, 22, 35, 59, 276, 580, 587,
588
alascensis, Zapus, 572
alaschanicus, Citellus, 445
alaschanicus, Euchoreutes, 579
alba, Cuniculus, 225
alba, Hystrix, 219
alba, IVIarmota, 461
albatus, Thomomys, 515
albertae, Citellus, 447
albescens, Callosciurus, 350, 371
albescens, Perognathus, 489
albicauda, Callosciurus, 375
albicauda, Thrinacodus, 135
albicaudatus, Thomomys, 516
albiceps, Ratufa, 385
albicus. Castor, 468
albida, Dasyprocta, 193
albifrons, Heliophobius, 85
albigularis, Thomomys, 520
albina, Heliosciurus, 401
albinasus. Pappogeomys, 528
albipes, Ratufa, 385
albipes, Sciurus. 334
albispinus, Proechimys, 117, 118, 122
albiventer, Petaurista, 282, 284, 288
albiventer group, Petaurista, 284, 286
albivexilli, Callosciurus, 350, 362
albogularis. Tamias, 436
albolimbatus. Liomys, 478
albolimbatus. Tamiasciurus, 346, 347
alboniger, Hylopetes. 298, 299, 300
aiboniger group, Hylopetes, 299, 300
albonotatus, Sciurus, 330
alborufus, Petaurista, 2S2, 286
alborufus group, Petaurista, 284, 286
albovittatus, Xerus, 421
albulus, Perognathus, 488
albus, Castor, 468
656
INDEX
albus. Cryptomys, yz
albus, Sciurus. 330
alexandrae, Thomomys. 517
alexandri. Paraxerus, 406, 409
aitan. Microsciurus, 310
ALLACTAGA. 22. 3;. ^i. ><>. 4g6. =162, 580
ALLACTAGAE, 3s, sf>i. sf>3. =iSo
ALLACTAGINAE, 562
allcni, Citellus, 449
alleni, Liomys, 477, 47S
alieni, Orthogeomys, 532
allcni. Sciurus, 32S, 339
alleni, Tamias, 435
alleni. Zapus, S73
ALLOMVIDAK, 15
alophus. Hystrix. 217
alphonsei, Sciurus. 32S, 343
alpina, Marmota, 461
alpinus, Glaucomys, 295, 296
alpinus, Sciurus, 330
alpinus, Tamias, 430
alpinus group, Tamias, 430
alpinus, Thomom\s, 315
alpmus group. Thomomys, 510. 515
alstoni. Callosciurus, 376
alstoni, Sciurus, 339
altaicus, Citellus, 445
altaicus, Sciurus, 331
altaicus, Tamias, 436
ALTICOLA. 40, 50. 604. 606
alticola. Graphiurus. 611
alticola, Perognathus. 4S7
alticola, Thomomys. 510, 512
altinsularis, Callosciurus, 364
altitudinis, Callosciurus. 356
altivallis. Thomomys, 510, 515
aluco, Pteromys, 293, 294
amankaragai. Scirtopoda, 592
amargosae, Thomomys, 51 >
amatus, Cr\ptomys, 87, yo
ambigua, Hystrix, 216, 220
ambiguus, Dipodomys, 496. 500
ambiguus, Perognathus, 489
ameliae, Sciurus, 331
amblyonyx, Kannabateomys, 137
americanus. Castor, 467
americana, Lagostomus, 236
americanus, Tamias, 436
americanus. Zapus. ^72
AMMODILLUS, 40, 56
ammodytes, Perognathus, 486
amoenus. Hyloputcs, 2 ,9
ammophilus, Perognathus. 489
AMMOSPHRMOPHILUS. 266, 268, 437,
439, 441. 451
amoenus, Perognathus, 487
amoenus, Tamias, 430, 432
amoenus group. Tamias, 430, 432
amon, Kliomys. 615, 61O
amotus. Menetes, 390
amplus, Citellus, 452
amplus, Perognathus, 486
amygdalei, Pteromys. 293. 294
anambae, Ratuta. 389
anambensis, Callosciurus, 370
anatolicus, Spalax, 642, 643
andina, Cuniculus, 225
ANDINOMVS, 39
andrcwsi, Callosciurus, 350, 369
andrewsi, Scirtopoda, 592
anerythrus, Funisciurus, 412, 414
angelus, D\Tom\s, 619
anglicus, Muscardinus, 625
angolae, Pedetes, 551. 552
angolae, Thnonomys, 14S
angolensis, Graphiurus, 607, 611
angonicus, Heliophobius, 85
angularis, Thomomys, 510, 511
angusticeps, Cratogeomys, 529
angusticeps, Ratufa, 3S9
angustirostris, Perognathus, 4S3, 4S9
angustirostris, Platygeomys, 531
angustus, Paraxerus, 406, 407
animosus, Paraxerus, 406, 407
animosus, Tamias, 433
AMSOMYKS, 38
ANISOMYS, 21, 18, 47
ANISONYX,437
anitae, Thomomys, 510, 512
annalium, Sciurus, 328, 336
annamensis, Petaurista, 282, 287
annandalei, Funambulus, 379
annectens, Citellus. 449
annectens, Liomys. 477
annellatus, Callosciurus, 350. 361
annexus, Orthogeomys, 532
annularis, Heliosciurus, 401
annulata, Allactaga, 586
annulatus, Citellus, 441, 451
annulatus, Heliosciurus, 401
anolaimae, Cavia, 240, 242
ANOMALURELLA. S37
AXOMALURiDAE, 1^2, 182, 535, S48
ANOMALURIXAE, 32, 537
AXOMALUROIDAE. 32. 78, 535
ANOMALURODOX, s37
ANOMALCROPS, 33, 57, 68. 536, 537, 541
AXOMALCRUS. 23, 33, 57. 68, 536, 537
anomalus, Sciurus, 324, 327, 328, 333, 623
anomalus, Heteromys, 122, 474. 475
anomalus, Cnptomvs, 92
AXOTOMYS. 39, 55
ansorgei, Graphiurus, 607, 610
ansorgei, Heterocephalus, 95
ansorgei, Cr\ptomvs, 87, 90
AXTELIOAIYS, 40
anthonyi, Sciurus, 32S, 337
anthon>"i, l>iom>s, 478
anthoiiyi, Perognathus, 491
antiUensis, Dasyprocta, 193
antiquus, Spalax, 644
antomae, Paraxerus, 406, 409
antonii, Ctenomys, 163, 165
antricola, Cercomys, 124
aoris, Callosciurus, 350. 370
apache, Perognathus, 484
INDEX
apache, Sciurus, 339
apache, Thomomys, 516
aptrea, Cavia, 240, 241
apereoides. Cavia, 242
apercoidcs, Ccrcomys, 123, 124
aphrastus, Thomomvs, 512
APLODONTIA, ,7, 23, 29, 44, 45. 33,
254, 255
APLODONTIIDAK, 29, 253
APLODONTOIDAi;, 29, 77, 253
APCJDKMUS, 22, 38, 48, 50
apollmari, Thrinacodus, nb
APOMVS, 38
apricus, Citellus, 453
aquilo, Callosciums, 350, 358
aquilonius. Dipodomvs, =04
arahicus, Ctenodacn I'us, \^8
ARAKOSCIL'RUS; 268, 3"22, 325
aralychcnsis, Allactaga, 584, 385
archidonae, Myoprocta, 197
arcticus, Sciurus, 332
ARCTOMVS, 262, 268. 454
arctomys. Marmot 1, 460
arcus, PeroRnathus, 485
arenarius, Cnptomys, 92
arenarius, Gcomys, 527
arenarius, Ptrognathus, 489
arendsis, Callosciurus, 371
arenicola, Citellus, 448
arenicola, Perognathus, 485
arequipac, Lagidium, 230, 232
arequipae, Cavia, 241
arens, Citellus, 449
arenivagus, Dipodomys, 501
argentescens, Funambulus, 379, 380
argenteocinereus, Heliophobius, 85
argenteus, Anomalurus, 542
argenteus, Graphiurus, 608
argenteus, Sciurus, 328, 333
argentinus, Sciurus, 328, 341
argusensis, Thomomys, 517
arizonae, Citellus. 453
arizonensis, Citellus. 453
arizonensis. Cynomys. 462
arizonensis. Perognathus. 485
arizonensis. Sciurus. 339
arizonensis, Tamias, 431
armalis, Callosciurus, 366
armatus, Atherurus, 208
armatus, Citellus, 4(7
armatus, Echimys, 106, iii, 112
armatus group, Echimvs, 109. 112
armatus. Perognathus. '491
armenaicus. Spalax. 642
arrhenii. Heliosciurus. 404
arsenjevi. Pteromvs. 294
artemesiae. Citellus. 446
artus. Perognathus. 4S3, 490
arundinis, Allactaga, 584
arusccnsis. Paraxerus, 406. 407
arusinus. Ratufa. 387
ARVICANTHIS. 38. 56
ARVICOLA. 41. 50
657
60
aschantiensis. Heliosciurus 401
ASCHIZOMYS. 40
asiaticus. Tamias, 430, 435
assamensis, Atherurus, 206
assamensis, Callosciurus, 372
atacamensis, Ctcnomys, 169
atahualpae. Cavia. 242
ater. .Spalacopus. 161
ater. Perognathus. 490
athene. Pteromvs. 293. 294
ATHERLRI. iy, 202
ATHERURUS 23. 27, 42, 49, 57, 59, 66. 69,
198. 203. 205, 208
ATLANTOXERUS. 30. ,r, 58, 262, 263
264, 267. 272. 307. 422
atratus. Callosciurus, 351, 374
atricapilla, Citellus (Orlov). 4;2
atricapillus, Callosciurus, 366
atricapillus. Citellus (Br\ant). 451
atrirufus. Sciurus. 336
atristriatus. Callosciurus. 369
atristriatus. Tamias. 431
atrodorsalis. Callosciurus. ;5i, 352, 362
atrodorsalis. Thomomys. 520
atrogriseus. Thomomys. 511
atronasus, Dipodomys. 500
atrovarius, Thomomys, 510, 521
atroventris, Sciurus, 339
atrox, Callosciurus, 367
attwateri, Geomys, 527
aubinii. Myrsilus, 417
aubryi, Heliosciurus, 401, 403
auceps. Galea, 242, 243
audubonii, Sciurus. 339
AULACODUS. 144
aulacotis, Allactaga, 586
aurantiacus. Hylopetes. 298, 299
auratus, Mesocricetus, 525
aurea, Dasyprocta, 194
aurea, Marmota, 457, 460
aureiventer, Ratufa, 384, 387
aureiventris, Thomomys, 516
aureogaster, Sciurus. 328, 334
aureogaster group, Sciurus. 327. 334
aureus. Lagidium. 231
aureus. Thomomvs. 516
auriculatus. Funisciurus. 412, 414
auriventris. Paraxerus. 407
austini, Sciurus. 336
australis. Caviella. 246
australis. Ctenomys, 166
australis. Heteromys, 474. 475
australis. Zapus. 573
austrinus. Geomys. 526
auzembergeri. Anomalurus, 541
avara, Marmota, 458
avellanarius, Muscardinus, 625
avellanus, Glis, 622
averini, Citellus, 445
avicennia, Sciurus, 338
avunculus, Microsciurus. 319, 320
awahnee, Thomomys, 515
azarae, Cavia, 240, 241
65S INDEX
azarae, Ctenomys, 163, 166
azarae. Dasyprocta, igi, 104
haberi, Koglaucomys, 298
bactriana, Allactaga, 585
bactrianus. Spermophilopsis, 425
badiatus, Pctaurista. 282, 287
badging. Callosciurus. 368
badius, Cannomys, 652
badius, Citellus. 448
bacticus, Sciurus, 331
baibacina, Marmota, 457, 460
baileyi, Castor, 467
baileyi Dipodomys, 490
baileyi group, Perognathus, 483, 487
baileyi, Pt'rognathus. 4S7
baileyi, Tamiasciurus, 346
bailevi, Thomomys, 519
BAIOMYS. 39
BAIOSCIURUS, 265. 266, 268. 321, 324
balae, Callosciurus, 357
balae, Ratufa, 388
balcanicus, Sciurus, 331
baliensis, Ratufa, 384, 385
baliolus, Sciurus, 336
balstoni, Callosciurus, 351, 369
balticus. Castor, 468
baluensis, Callosciurus, 351, 367
bancana, Ratufa, 387
bancanus, Nannosciurus, 316
bancarus, Callosciurus, 336
bandarum. Funisciurus, 412, 414
BANDICOTA. 38, 48
bangkanus, Callosciurus, 366
bangsi, Cilaucomys, 296
bangsi. Perognathus. 485
bangsi, Sciurus, 336
bangueyae, Callosciurus, 351, 357
bangueyi, Ratufa, 38S
banksi, Callosciurus, 368
banksi, Pctaurista, 286
baraniensis, Ratufa, 384, 386
liarawensis, Paraxerus, 408
barbarus, Ctenom\s, 163, 167
barbei, Callosciurus, 351, 354
barberi, Sciurus, 338
barroni, Pctaurista, 282, 2S7
barrowensis, Citellus. 448
bartelsi. Aeromys, 291
bartoni, Callosciurus, 351, 360
bashkiricus, Sciurus, 332
batesi. Anomalurus, 540, 541
BATHYKRGI, 24. 81
BATHYKRGIDAH, 24, 79
BATHVKRGOIDAE. zi„ 77. 79
BATHVKRGOMGKPH SERIES, 23
BATHYERGUS, 22, 24, 57, 66, 80. Si, 82
83, 1 89
BATOMYS, 38
batuana, Pctaurista, 282, 285
batuana, Ratufa, 3S6
batus, Callosciurus, 356
baudensis, Sciurus, 342
bayonii, Heliosciurus, 405
bt-a, Protoxerus, 415, 416
BEAMYS, 38, 56
bcatus, Funisciurus, 412, 414
bec'bei, Callosciurus, 373
beechiyi, Citellus, 442, 451
beecrofti, Anomalurops, 542
beirae, Cn.ptomys, 87, 91
beirae, Heliosciurus, 402
belcheri, Tamiasciurus, 347
beldeni, Anomalurus, 540
beldingi, Citellus, 442, 447
belfieldi, Drcmomys, 381, 382
bellaricus, Funambulus, 378, 380
bellona, Callosciurus, 351, 373
BELOMYS, 30, 4s, 49, 64, 254, 263, 274, 275,
276, 279, 304
belone, Hylopetes, 299
belti, Sciurus, 336
belugae. Castor, 467
benga, Heliosciurus, 403
bengalensis, P'unambulus, 378
bengalensis, H>strix, 217
bengalensis, Ratufa, 384, 385
bennetti, Abrocoma, 154
bennetti group, Abrocoma, 154
bentincanus, Callosciurus, 363
berdmorei, Menetes, 390, 405
bergi, Ctenomys, 163, 165
benngensis, Citellus, 448
berkele\'ensis, Dipodomys, 498
bernardinus, Citellus, 442, 454
bernardinus, Perognathus, 491
ber\tensis, Spalax, 642
besuki, Callosciurus, 351, 368
betulina, Sicista, 566, 567
betulinus, Pteromys, 294
bhotia, Drcmomys, 381
bhutanensis, Callosciurus, 351, 358
bicolor, Coendou, 182, 185, 187
bicolor group, Coendou, 185, 187
bicolor, Ctenomys, 169
bicoior, Dasyprocta, 194
bicolor, Heteromys, 474, 475
bicolor, Ratufa, 384, 3S5
BIFA. 613
bigalkci, Cr\'ptomys, 92
bilimitatus, Callosciurus, 351, 36S
bilineatus, Callosciurus, 351, 368
bilkjewiczi. Dyromys, 619
billitonus, Callosciurus, 372
bilobidens. Galea, 243
bimaculatus. Perognathus, 484
binominatus, Citellus, 442 (footnote),
bipcs, Jaculus, 596
birrelli, lY-taurista, 282. 288
birulai, Castor. 468
birungensis, I-"unisciurus, 415
bistriatus, Isothnx, 114
bistriatus group, Isothrix, 114
blainei, Cryptomys, 87, 90
blainvillei, Echimys, 109, iii, 112
blainvillei group, Echimys, 109, 112
INDEX
659
blandus, Bclomys, 277
blanfordi, Callosciurus, 351, 373
blanfordi, Hystrix. 21S
blanfordi, Jaculus, 595, 597
HLANKORDIMYS, 41, 50
BLARINOMYS. 39
biythci, Callosciurus, 351, 353, 373
bobak, Marmot, I, 455, 457, 460
bobak yroup, Marmota, 457, 460
bocaRei, Cryptomys, 87, 91
bocki, Callosciurus, 351, 3^:8
bocourti, Callosciurus, 351, 352, 362
bochmi, Paraxerus, 406, 409
boehmi group, Paraxerus, 405, 406, 409
boimensis, Proechimys, 119
boliviac, Dasyprocta, 191, 195
bolivianus, Proechimys, 118, 121
boliviensis, Coendou, 18=;. 186
boliviensis, Ctenomys, 163, 164, 168
bolivicnsis, Dactylomys, 136, 137
boliviensis, Galea, 242
boliviensis, Sciurus, 328, 344
bolovensis, Callosciurus, 360
bombycinus, Perognathus, 485
bonariensis, Myocastor, 144
bondae, Sciurus, 328, 342
bongensis, Heliosciurus, 401
bonhotei, Callosciurus, 351, 359
boothiae, Sciurus, 328, 336
boquetensts, Microsciurus, 319, 320
borealis, Tamias, 430, 431
borealis, Thomomys, 521
borneanus, Callosciurus, 351
borneanus, Nannosciurus, 315, 316
borneanus, Pteromyscus, 281
borneoensis, Callosciurus, 366
borneoensis, Petinomvs, 301
BOKOMYS, 169
bottae, Thomomys, 510, 511
bottae group, Thomomys, 510, 511
boxi, Lagidium, 230, 232
boydi, Funisciurus, 412, 414
bozasi, Lophiomys, 636
brachialis, Heleromvs, 475
BRACHIONKS, 40
brachiura, Citellus, 447
brachyotis, Allactaga. 586
brachvotis, Xerus, 420
BRACHYTARSOMYS. 40. s7, 645
BRACHYTARSOMYES. 40'
BRACHYUR(;MYS, 22. 40. 57. 602. 64s
BR,\CHYUR()MYKS, 40
brachyurus, Allactaga, 586
brachyurus, Eurjzygomatomys, 125
brachyurus, Geocapromys, 132
brachyurus, Hystrix, 200, 211, 212, 213. 214.
217
brachvurus group, Hvstrtx, 212, 217
BRASIINAK, 14
BRAMUS, 645
brandti. Coendou, 185, 187
branickii, Dinomys, 171
brasiliensis, Cavia, 242
brasiliensis, Ctenomys, 168
brauneri, Citellus, 442. 444
brauni, Heliosciurus, 401, 404
braziliensis, Echimys, iii, 112
brevicauda, Citellus (Brandt), 442, 443
brevicauda, Proechimys, 118, 120
brevicaudata, Chinchilla, 229
brevicaudus, Citellus (Merriam), 454
breviceps, Geomys, 527
breviceps group, Geomys, 526
brevidens, Thomomys, 513
brevinasus, Dipodomys, 501
brevinasus, Perognathus, 485
brevirostris, Geomys, 527
brevispina, Hystrix, 217
bridgemani, Paraxerus, 406, 408
bridgeri, Thomomys, 522
bridgesii, Octodon, 159
brochus, Syntheosciurus, 321
brockmani, Graphiurus, 607, 609
brodiei, Funambulus, 378, 380
brooki, Callosciurus, 351, 353, 374
BROTOMYS, 169
broweri, Marmota, 459
brownii, Geocapromys, 132, 133
browni, Microsciurus, 319
brucei. Allactaga, 584
bruneri, Erethizon, 181
brunneoniger, Sciurus, 328, 344
brunnea, Sciurus, 330
brunneus, Citellus, 447
bryanti, Perognathus, 492
bryanti, Sciurus, 338
buccatus, Citellus, 450
buckleyi, Citellus, 450
budini, Abrocoma, 154, 155
budini, Ctenomys, 163, 167
buechneri, Pteromys, 294
buffonii, Geor\'chus, 86
bulana, Ratufa, 386
bulbivorus, Thomomys, 510, 524
bulbivorus group, Thomomys, 510
bullata, Allactaga, 58?, 584
bullatus, Glaucomys,'297
bullatus, Thomomys, 522
bulleri, Liomys, 477, 479
buUeri, Pappogeomys, 527, 528
bulleri, Tamias, 430, 434
bunguranensis, Ratufa, 384, 387
bungei, Marmota, 459
bunkeri, Marmota, 458
bursarius, Geomys, 526
bursarius group, Geomys, 526
burrowsi, Atherurus, 206, 208
burrus, Proechimys, 120
burti, Thomomys, 519
butleri, Graphiurus, 607, 609
butieri, Jaculus, 595, 596
buxtoni, Citellus, 446
byatti, Paraxerus, 400, 406, 409
cabezonae, Dipodomys, 503
cabezonae, Thomomys, 517
66o
INDEX
cacodemus, Tamias, 431
caecator. Castor, 4A8
caedis, CalloscJurus, 367
cafer, Pedetes, 551, 552
cagsi, Callnsciurus, 374
calamophayus, Thr\'ononi\'s, 148
calidior, Drcmom>s, 381, 2^2
calidior, Procchimys, 118, 120
californica, Aplodonti-J, 259
californicus, Dipodomys, 49S
cajifornicus, IVIicrodipodops, 493
californicus, Glaucomys, 297
californicus, Perognathus, 491
californicus group, Peroynathus, 4S2, 491
californicus, Tamiasciurus, 347
caliyata, Marmota, 453. 45", 459
caligata group. Marmota, 455, 459
caliginosus, Thomomys, 320
callida, Dasyprocta, 195
callipcplus. Tamias. 430. 433
callistus, Perognathus, 4S4
calliurus, Protoxerus. 413, 416
CALLOSCIURUS. 3o.'49. 52, 58, 65, 263,
266, 267. 270, 311, 312, 348, 377, 380
CALLOSPERMOPHILUS. 266, 268, 437.
439. 441. 442, 453
CALOMYSCUS. 39. 51
calotus. Sciurus, 333
campestris, Zapus, 571, 572
campioni. Marmota. 458
camtschatica, Marmota, 457, 459
canadensis, Castor, 464, 466, 467
canadensis group, Castor. 467
canadensis, Geomys, 526
canadensis, Marmota, 457
canadensis, Zapus, 571
canaster. Heliosciurus. 401
candelensis. Das\'procta. 196
candelensis, Sciurus, 341
candidulus, Petaurista, 282, 284, 2S7
canescens, Citellus. 449
canescens, Dactyloniys, 136, 137
canescens, Georychus, 86
canescens, Glaucom\s, 296
canescens, Perognathus, 490
canescens, Tamias, 433
canfieldae, Citellus, 452
canicaudus, Tamias, 432
caniceps, Callosciurus, 351, 363
caniceps group, Callosciurus, 332, 363
caniceps, Diplomys, 115
caniceps, Protoxerus, 416
caniceps, Petaurista, 2S2, 284, 288
caniceps, Tamias, 431
canigenus, Callosciurus, 3()3
canicollis, Proechimys, 117, 118, 122
canipes, Tamias, 434
CANNOMYS, 37, 44, 48. hb, 645, 646, 651
cantWL'lli, Perognathus, 485
canus, Citellus, 446
canus, Thomomys, 516
canus, Liomvs, 479
"CAPROMYIDAK," 134
CAPROMYINAE, 25, loi, 102. 103, 128
169
CAPROMYS, 21, 23, 23. 42, 54. 73. 128,
132, 154
capensis, Georychus, 86
capensis, Graphiurus, 608
capensis, Hystrix, 219
capensis, Pedetes, 552
capensis, Xerus. 420, 422
capistratus, Sciurus, 339
capitis, Paraxerus, 406. 408
captorum, Spalax. 642, 6|3
capvbara, H\drochoerus, 253
CARDIOCRANIINAE, 33. 363, 574
CARDIOCRANIUS. 33, 43, 31. 59, 493,
561, 562, 575, 576
careyi, Callosciurus, 331, 360
carimatae, Callosciurus, 366
carimonensis, Callosciurus, 351, 366
carimonensis, Ratufa, 384, 386
caroli, Callosciurus, 351, 367
caroli, Hylopetes, 299
carolinensis, Castor, 467
carolinensis, Dasyprocta, 195
carolinensis group, Sciurus, 333
carolinensis, Sciurus, 324, 327, 328, 333
carpathicus, Dvromys, 618
CARPOMYS,'38, 48
carrikeri, Echimys, 109, 113
carruthersi, Citellus, 442, 444
carruthersi, Funisciurus, 412, 414
CARTERODON, 25, 53, 73, 107, 125
caryi, Citellus, 453
caryi, Perognathus, 484
car\i, Tamias, 431
car>'i, Thomomys, 522
cascadensis, Marmota, 459
cascadensis, Tamiasciurus, 347
casensis, Callosciurus, 364
caspicus, Glis, 623
castancoventris, Callosciurus, 351, 358
castaneus, Cannomys, 652
castaneus, Echimys, 112
castaneus, Lariscus, 392
castaneus, Petaurista, 282, 286
castanops, Cratogeomys, 528, 329
castanotus, Sciurus, 337
castanurus, Citellus, 433
CASTOR, 3, 23, 31, 52, 33. 5H. 62, 70. 97,
140. 465
CASTORIDAE, 31, 25^. 464
CASTOROIDAE, 30, 77, 464
CASTGROIDES, 466
castoroides, M\ocastor, 144
CASTOROIDIDAE, 14, 4 6
castus, Sciurus, 328, 343
catalinae, Sciurus, 340
catahnae, Thomomys, 518
catavinensis, Thomomys, 513
c;itellus, I'^uryzypomatomys, 125
catemana, Ratufa, 387
catrinae, Dasyprocta, igi, 194
cattoiri, CJraphiurus, 608
INDEX
66i
Caucasia, AlIactaRa, 584, 585
caucasica, Sicista, 568
caucasicus, Dyromys, 619
caucasicus, Sciurus, 333
caucensis, Sciurus, 343
caudata, Dasyprocta, 194
caudata, Marmota. 455, 457, 460
caudata Rroup, Marmota, 455, 457, 460
caudata, Sicista, 566, 568
caurinus, Heliosciurus, 401, 403
caurinus, Tamias, 432
cavator, Macroseomys, 534
CAVIA, 13, 23, 29, 55, 71, 171, 189, 224, 238,
239, 240
CAVIAE, 29, 239
CAVIELLA, 29, 71, 238, 239, 240, 243, 246
CAVIIDAE, 28, 96, 97, 170, 189, 221, 237
CAVIINAE, 29, 238
CAVIOIDAE, 28, 77, 237
cayanus, Dasyprocta, 193
cayennac, Dasyprocta, 191, 193
cayennensis, Proechimys, 118, 120
caycnnensis group, Proechimys, 120
cavmanum, Dasyprocta, 196, 197
CELAENOMYS, 39, 48
celaenopepla, Ratufa, 384, 3S9
celsus, Dipodomys, 504
CENTETIDAE, 8
centralis, Atherurus, 206, 20S
centralis, Callosciurus, 351, 359
centralis, Coendou, 185, 186
centralis, Dolichotis, 248
centralis, Jaculus, 595, 596
centralis, Marmota, 457, 460
centralis, Proechimys, 118, 120
centralis, Ratufa, 384, 385
centralis, Thomomys, 516
centricola, Dolichotis, 248
centricola, Protoxerus, 415, 416
ccpapi, Paraxerus, 406
cepapi group, Paraxerus, 406
CEPHALO.MYS, 169
CERATOGAULUS, 258
CERCOLABES, 182
CERCOMYS, 25, 72, 107, 108, 123
certus, Citellus, 454
cervicalis, Sciurus, 335
cervinus, Thomomys, 517
ceylonica, Ratufa, 384
chaclovi, Allactaga, 586
chadensis, Xerus, 420, 421
CHAETODIPLS, 480, 4S2, 487
CHAETOMYINAE, 27, 174
CHAETOMYS, 3, 23, 27, 55. 74. I74. 175
CHALICOMYIDAE, 14, 466
CHALICOMYS, 466
chapensis, Typhiomys, 631
chapini, Anomalurops, 542
chapmani, Dipodomys, 502
chapmani, Sciurus, 328, 341
CHELEMYSCUS, 39
cherriei, Macrogeomys, 534
cherriei, Proechimys, 118, 121
chiapensis, Dasyprocta, 194
chiapensis, Heteroceomys, 533
chiapensis, Sciurus, 335
chihliensis, Sciurus, 332
chihuahuae, Thomomys, 520
chilensis, Coendou, 186, 188
chilcnsis, Lagidium, 231
chilensis, Ctenomys, 169
chilensis, Mvocastor, 141
CHIl.OMYS, 39
CHINCHILLA, 23, 28, ^5, 72, 171, 224, 226,
227
chinchilla, Chinchilla, 229
CHINCHILLAE, 28, 227
CHINCHILLIDAE, 28, 98, 99, 170 226
CHINCHILLULA, 39, 55
chintalis, Dremomys, 381, 382
chinensis, Callosciurus, 351, 376
chiricahuae, Sciurus, 339
chiricahuae, Thomomys, 521
chirindensis, Heliosciurus, 402
chiriquensis, Sciurus, 328, 340
chiriquinus, Proechimys, 11 8, 120
CHIROMYSCUS, 3,'36, 38
CHIROPODOMYS, 3, 36, 38, 48, 635
chlorus, Citellus, 453
chobiensis, Paraxerus, 406
choco, Sciurus, 341
chocoensis, Dasyprocta, 195
CHOEROMYS, 144, 145
christvi, Graphiurus, 607, 6n
CHROTOMYS, 39, 48
chrysaeolus, Proechimys, 118, 120
1 hr\seola, Aplodontia, 258
chrysippus, Funisciurus, 412, 415
chrysodeirus, Citellus, 442, 453
chr\'sogaster, Sciurus. 334
chrysonotus, Callosciurus, 363
chr\-sonotus, Thomomys, 516
chr\'sophaenus, Anomalurus, ^40
chr^sothrix, Petaurista, 288
chr>'suros, Sciurus, 343
chr\surus, Echimys, iii, 113
chrysurus group. Echimys, 109, 113
chuscensis, Sciurus, 338
cicur, Petaurista, 285
cilicicus, Spalax, 642
cincticauda, Eliomys, 616
Cinderella, Petaurista, 282, 287
cineraccus, Graphiurus, 610
cinerascens, Citellus, 442, 453
cinereicollis, Tamias, 430, 434
cineraceus, Petaurista, 2S2, 284, 287
cinerascens, Perognathus, 487
cinerea group, Abrocoma, 154
cinerea, Abrocoma, 154, 169
cinerea, Geomys, 526
cincreus, Anomalurus, 540, 541
cinereus, Eupetaurus, 304
cinereus, Rhizomys, 650, 651
cinereus, Sciurus, 328, 330, 333, 33S
cinereus, Tamias, 434
cinereus, Thomomys, 512
662
INDEX
cinereus, T\phlomys, 63 1
cinereus, Zapus, s72
cineris, Perognatlius, 4S6
cmnamomeus, Callosciurus, 351, 361
cinnamoineus, Citcllus, 452
cinnamomeus, Proechimys, 119
CITELI.US. 23, 30, 51,' S3. sS, 61, 70, 262,
26ft, 267, 268, 269, 272, 307, 328, 437,
440. 455
citellus, Citellus, 428, 442, 444
citellus uroup, Citellus, 440, 444
citrinus, Anomalurops, ^42
clarencei, Dipodomys, 500
clarkei. Callosciurus. 351, 355
clarkei. Petaurista, 282^ 288"
clarkii, Cratogeomys, 520
clarus. Pcrognathus, 487
clarus. Tamias, 431
CLAVK;LIS. 604. 607. 60S
cleomophila, Dipodom\s. S02
cleomophila, Perognathus. 484
CLETHRIONOMYS, 40. 50, 52
cliftoni. Marmota. 457, 460
clivorum, Octodon, isg
clusius, Thomomvs, s22
CLYOMYS, 25, 55, 72. 107. 108. 125
cobaya. Cavia, 242
cockerelli, Callosciurus. 351, 352, 361
cocalis. Sciurus. 328, 34s
cocos, Sciurus, 32S, 335
coecutiens. Crvptomys, 87, 90
COELOGEXU.S. 221. 224
COELOGENYS. See Coclogenus
COELOMYS, 38
COENDOU, 3. 23, 27, 54. 55. 74. 145, IS4,
173. 174. 177. 178. iSi, 182
coenosus, Heliosciurus, 401
COETOMYS, 86
coibae, Dasyprocta. 191, 195
colburni, Ctenomys, 168
colimensis. Sciurus, ^^^
collaris. Graphiurus, 610
colliaei, Sciurus, 328, 335
collinus, Thomomys, 518
COLOBOTIS. 437. 439
COLOiMYS, 38, 56
colonus, Geomys. 526
coludo, Ctenomys. 163, 166
Columbiana, Aplodontia. 259
Columbiana, Dasyprocta, 193, 195
columbianus, Citellus, 447
columbianus, Cynomys, 463
columbianus, Dipodomys, 496, so2
columbianus, Perognatlius, 487 '
columbianus, Proechimys, 118, 120
columbianus, Thomom\s, 522
columbiensis, Glaucomys, 296
columbiensis, Tamiasciurus, 347
comes. Galea, 242
comonnus, Funambulus, 378, 380
compactus, Dipodomys, 497, ^03
compactus group, Dipodomys, 498, 503
conca\us, Heterogeomys, 533
concinnus, Nannosciurus, 31 s, 316
concisor, Thomomys, 513
concolor, Callosciurus, 331, 365
concolor, Citellus, 442, 443
concolor, Rattus. 47
concolor, Sciurus, 338
concolor, Sicista, 566, 567
concolor group, Sicista, 566, 567
conditi, Perognathus, 488
condurensis, Callosciurus, 351, 366
condurensis, Ratufa, 3S4, 386
confinalis, Thomom\s, S14
confinis, Ratufa, 386
confinis, Tamias. 431
congicus. I*"unisciurus. 412. 4n
congicus group. E'unisciurus, 412, 413
congicus, Thryonomvs, 148
CONILURUS, 38, 47
conipus, Callosciurus, 372
connectens, Citellus, 453
connectens, Thomom\s, 514
conradsi, Hystrix, 220
conscius, Hetcromys, 475
consitus, Cratogeomys, 529
consobrinus, Sciurotamias, 426
consobrinus, Tamias, 430, 431
conspicua, Ratufa, 384. 386
consularis, Menetes, ^90. 391
contuma.x. Callosciurus. 351. 360
convergens, Thomomys, 513
convexus, Cratogeomys, ^30
cooperi, Tamias, 434
copei, Perognathus, 483
coreae, Sciurus, 328, 332
corilinum, Muscardinus, 625
corybantium, Spalax, 642, 643
costaricensis, Alacrogeomvs, 534
cothurnata, Ratufa, 384, 386
couchii, Citellus, 4^0
couesi, Erethizon. 181
couiy, Coendou, 185, 188
coupei, Graphiurus, 610
coypus, Myocastor, 141
cradockensis, Cryptom\s, 92
crassicauda group, -Salpingotus, S75
crassicauda, Salpingotus. 57s
crassicaudatus, Graphiurus, 607, 608
crassicaudatus group, CJraphiurus, 608
crassicaudatus, Isothrix, 114
crassidens, Lagidium, 231
crassidens, Thomom>s, 520
crassirostris. Heteromys. 476
crassispinis, Thecurus. 199, 200. 211
crassispinis group, Thecurus, 211
cratodon, Dipodomys, 499
crassus, Lagostomus, 236
CRATEROMYS, 36, 38, 48 602
CRATOGEOMYS, 32, 63, 71, so6, :;o7, 528
"CRICETIDAE," 18 " '
CRICETINAE, 39
CRICETODIPLS, 4'-o
CRICETOMYS, 38, 56
CRICETULUS, 19, 51
INDEX
663
CRICETUS, 39. 51. 562, 603
crinicer, LaKitlium^ 231
criniKtT, Lat,'ostniTius, 236
crinigcrum, Layidium, 231
crinilus, Peropnathus, 490
crinitus, Petinomys, 301, 302
crispus, Liomys, 478
crispus group, Liomys, 477, 478
cristata, Dasyprocta, 191, 196
cristata. Hystrix, 98, 148, 200, 201, 215, 216,
217, 219
cristata Rroup, Hystrix, 212
cristatus. Echimys, 113
croaticus, Sciurus, 328, 331
croconota, Dasvprocta, 191, 193
CROSSOMYS, 36, 39, 47
crotalius, Callosciurus. 351, 359
crumpi, Callosciurus, 351, 359
CRUNOMYS. 38, 48
CRYPTOMYS, 24. 44. 57. 67, 81. 82, 86
cryptosrilotus, Citellus, 450
CTENODACTYLIDAE. 34, 96, 102, 553
CTENODACn'LOIDAE, 33, 78, 553
CTENODACT\'LUS. 23, 34. 50, 56, 58, 68,
553. 554. ^>5. 556, 603
ctenodact\'lus, Paradipus. S90
CTENO^IYS, 3. 23. 26. 55". 73, 102, 155. 156,
161, 240, 554, 603
cuandu, Coendou. 186
cucutae, Sciurus, 342
cumberlandianus, Geomys, 526
cumingii, Octodon, 159
cunealis, Petromus, 149, 151
cuneiceps, Hystrix, 217, 218
cunicularius, Cercomvs, 124
CUNICULIDAE, 28. 99. 220
CUMCULUS. 13, 23, 28, 54, S5, 74, 97.
100, 170, 171, 189, 190, 221
cuniculus, Orthogeomys, 531
cupidineus, Dipodomys, 502
currax, Pedetes, 552
curtatus, Thomomys, 512
curtatus. Zapus, 572
cuscinus, Sciurus, 328, 344
cuscus, Lagidium, 230, 231
cutleri, Cavia, 242
cuvieri, Abrocoma, 154
cuvieri, Hystrix, 217, 219
cuvieri, Lagidium, 231
cvunus, Spalacopus, 161
CYLINDKODONTIDAE, 15
CVNOMVS, 30. 43, 53, 62, 70, 262, 266, 267,
268, 273, 307. 461
cyrenaicus, Eliomys, 615, 616
dabagalia, Xerus, 420, 421
dacota, Marmota, 458
DACNOMYS, 38. 48
Jactylinus, Callosciurus, 351, 360
daci\ii .us, Dactvlomvs, 134. 136
DACTYLOMYINAE, 25, 98, 102, 105. 134,
169
DACTYLOMYS, 21, 23. 25, 55, 73, 102, 123,
129, 135. 136, 137
dagcstanicus, Dyromys, 619
dakotensis, Tamiasciurus, 346
damarensis, Cryptomys, 87, gi
damarensis, Pedetes, 552
dandolena, Ratufa, 384, 385
dariensis, Dasyprocta, 194
daricnsis, Macrogeomys, 534
darlingi, Cr\'ptomys, 87, 91
darlingi, Diplomys, 115
darricarrerei, Jaculus, 597
DASYMYS. 38. 56
DASYPROCTA, 13, 23, 27. 54. 55. 74. 170,
189. 190, ig6
DASYPROCTIDAE, 27, 97, 99, 100, 170,
173, 189, 221
dasythrix, fCchimys, 109, iii, 112
dasythrix group, Echimys, 109, 112
datus, Callosciurus, 371
daubentoni. Hystrix, 219
daucinus. Heliosciurus, 401, 402
dauricus, Citellus, 442, 445
davidi, Rhizomys, 650
davidianus, Sciurotamias, 426
daviesi, Zapus, 571
davisoni, Callosciurus, 351, 363
davisoni, lomys, 303
deaibata, Ratufa, 384, 385
deaysi, Dipus, 591
decolorata, Ratufa, 384, 389
decoratus, Menetes, 390, 391
decumana, Allactaga, 584, 586
decumanus, Proechimys, 118, 120
degus, Octodon, 159
dekan, Rhizomys, 651
delesserti. Funambulus, 379
demissa. Galea, 242, 243
DENDROMUS, 39. 56
DENDROMYINAE. 39
dentatus. Pedetes, 552
DEOMYINAE, 38
DEOMYS, 17, 34, 38, 44, 45, 46, 56, 564, 599
depauperatus, Thomomys, 516
deppei, Sciurus, 324, 327, 328
deppei group, Sciurus, 337
depressus, Thomomys, 512
derbianus, Anomalurus, 540
deserti, Dipodomys, 496, 497, 504
deserti group, Dipodomys, 497, 504
deserti, Jaculus, 595, 597
desertorum, Thomomys, 518
desitus, Thomomys, 514
desmarestianus. Heteromvs, 474, 475
DESMODILLISCUS, 36, 40, 56
UKSMOUILLUS, 40, 56
detumidus. 'I'homomys, 514
dextraiis, Callosciurus, 351, 362
diabolj, Thomomys, 511
diana, Lagostomus, 236
diardi, Callosciurus, 376
dichrous, Marmota, 457, 461
dickeyi, Microdipodops, 493
6&4
INDEX
UlCROSTONYX, 40. 50. 52
didflphoides, Mesoniys, 127
dilutus, Caliosciurus, 371
dilutus, Citi-llus, 445
dimidi:itus. I'rocchimys, lid, il7. llS, 122
dinniki, Alactagulus, sS?, sSS
DINOMVIDAE, 26. yo, 170, 173
DINOMYS, n, 26, s^. 72. i|S. 171
DIPl.OMYS, 2i, S4. 72, ic'i. loS, 115. 603
DIl'OUHS. 3:, S64. 'i'^'^
DIPODIDAi:. 34. 560
DIPODINAK, 3S, q6i, 363. 579
DIPODIPCS. sqo
DIPODUIDAK. u. 7^. 560
DIPCDOMYI-S. 32, 471. 4Q4
DII'CIDOMYINAE, 31. 47i. 479
DIPODOMYS, 3, 22, 31. 32, 53. 54. '>2. 71.
460, 470, 471. 4^3. *9*.. 583
DIP<1DC)PS, 494
DIPUS. 35. 51, sy. 562, 589. 590
director, Caliosciurus, 371
dispar, PeroRnathus, 49 1
dissimilis, Thomomys, 516
dissonus, Protoxerus, 41 s, 4'6
DISTAECHURUS, 543
di\ergens, Thomomys, 513
diversus, Lariscus, 392
dixoni, Dipodomys, 498
dolbrogeae, Spalax, 642, 643
dolichoccphalus, Macrogeomvs, S34
DOLICHOTIDES, 29, 239, 247
DOLICHOTl.S, 3, 13, 23, 29, 55, 71, '71,
189, 238, 240, 247, 248
dollmani, Graphiurus. 609
DOLOMYS, 41, 50
dolosus, Heliosciurus, 402
dolphoides, Caliosciurus, 376
domelicus, Caliosciurus, 351, 363
domensis, PcroKnathus, 4S8
doppelmayri. Marmota, 460
dorothcae, Graphiurus, 607, 609
dorsalis, Ctenomys, 163. 166
dorsalis, Sciurus, 324, 328, 336
dorsahs, Tamias, 429, 430, 435
dorsalis, Xerus, 420, 421
dorsatum, Ercthizon, 178, 181
douglasi, Citellus, 442, 451
douglasi, Tamiasciurus, 346, 347
douglasi, Thomom>"s, 510, 523
douglasi group, Thomomys, 510, 523
dravidianus, Funambulus. 379
DREMOMYS, 30, 49, 58, 65, 263, 267, 271,
311, 312, 349, 380
dr\'as, Dvromvs, 618
DRYOMYS, '616
dschmschicus, Xtrus, 422
duida, Sciurus, 344
dukciti, Sciurus, 331
dulittnsis, Caliosciurus, 351, 371
dulitt-nsis, Ratufa, 387
dunni, Hctcroccphalus, 95, 9'J
durangae, Tamias, 434
durangi, .Sciurus, 328, 338
durangi, Thomom>s, 320
dussumieri, I-unambulus. 379
OYROMYS, 37, 51, 60, 602, box. 612, 613,
616
tlxungariac. Allactaga, 585
eastwoodac, Graphiurus, 608
cbii, Epixerus. 417, 418
cbori\orus. Protoxerus. 415. 416
ecaudata. Hystrix. 217
ecaudatus, Mesomvs, 127
ECHIIMYIDAE, iq, 98, 100. 101, 17?
ECHIMYINAE. 25. 106, 169
IX'HIMYS. 25, 55, 72, 106. 108, 114. 123.
134. 174
ECHINOMVS. loS
IXHINOPROCTA. 3, 27, ■;=;. 74, 173. I77,
i7,S, 181, 1 86
ECHINOSCIL'RL'S. 26s, 267, 208, 321, 324
ECHIUTHRIX, 38, 48
edax, Thrinacodus, 133, 136
edithae, Trogopterus, 279, 280
effugius. Sciurus, 335
ehrenbergi, Spalax. O42
elassodontus, Hylopetes, 300
elater, Allactaga. 561, 583, 584, 585
elater group. Allactaga, 584, 585
elator, Dipodomys. 500
elbertae. Caliosciurus, 375
electilis, Petinomys, 301
electus, Paraxerus, 406, 407
elegans, Capromys (Phloeomys), 131
elegans, Citellus, 442, 447
elegans, Graphiurus, 608
elegans, Glirulus, 620
elegans, Heliosciurus, 401
elegans, Petaurista, 2S2, 285
elegans, Proechimys, 119
elephantinus, Dipodomys, 503
elibatus, Perognathus, 483
ELIGMODONTIA, 39,"55
ELIOMYS, 22, 37, 46. 51, 56, 60, 69, 601,
602, 603, 612, 613
ELIURI, 38
ELIURUS, 38, 37, 64^
ELl.UBIUS, 3, 36, 41, 50, 81, 64:
elphinstonei, Ratufa. 385
emiliae, Lonchothrix, 128
emiliae, Petaurillus, 302, 303
emilianus, Ctenomys, 163, 164, 167
emmi, Heliophobius, 85
emmi, Paraxerus, 406, 409
emini, Funisciurus, 413
emissus, Heliosciurus, 401, 403
emotus, Thomomys, 519
empetra, Citellus, i, 448
empetra, Marmota, 457
engelhardti, Marmota. 458
enixus. Thomomvs. 320
ENTUPTYCHINAE. 13
EOGLAUCOMYS, 30. s2. 37, 27O. 297, 298
EOMYINI, 18
INDEX
66s
EOSCIURUS, 262. 263. 383
EOTHENOMYS. 40, 48
EOXERUS, 263. 263. 376
EOZAPUS. 35. 51. 59.568
t'phippium, Ratufa, 384, 386
KPKiAULUS. 2s8
KPIXICRL'S, 30, 67. 264. 272, 308. 327, 417
cpomophorus, Callosciurus, 351, 364
cpixanthum, Krcthizon, 178. 181
ercbus, Callosciurus, 351, 367
eremicus, Perognathus, 489
KREMODIPUS, 35. 59, 597, 598
eremonomus, Citcllus, 4^2
ERETHIZON, 23, 27. 53. '^3, i73. i74, i77.
178
ERETHIZONTIDAE, 26, 98, 99, 173
ERETHIZONTINAE, 27, 174, 177
ERIOMYS. 227
EROPEPLUS, 38
CTubcscens, Callosciurus, 365
erythraeus, Callosciurus, 351, 352, 357
enthraeus group, Callosciurus, 352, 357
er\ throbronchus, Graphiurus, 610
er>'throgaster, Callosciurus. 351, 358
cryihrogcnys, Citellus, 442, 443
erythrogenys, Funisciurus, 412, 414
erythrogcnys, Rhizomys, 651
erythrogluteius, Citellus, 447
erythromelas, Callosciurus, 367
er>thronotus, Anomalurus, 540
erythrops, Funisciurus, 413
erythropus, Xerus, 420, 421
escuinapae, Liomys, 477
esculentus, Glis, 622
estor, Cratogeomys, 528, 529
EUCHOREUTES. 35, 51, s9, 562, 563. 577
EUCHOREUTINAE, 35, 561, 563, 576
EUNEOMYS, 39
EUPETAURUS, 30, 52, 58, 262, 263, 268,
274, 304
euphratica, Allactaya, 584, 585
eureka, Zapus, 573
europaea, Hystrix, 219
europaeua, Sciurus, 330
EURYZYGOMATOMYS, 25, 72, 107, 108.
124, 140
EUTAAUAS, 266, 267, 268, 426. 427, 42S,
EUTiTOMYlDAE, 14
EUXERUS, 264, 267, 418. 420, 421, 441
cveretti, Drcmomys, 381, 382
everetti, Hylopetes, 299, 300
eversmanni, Citellus, 442, 445
eversmanni group, Citellus, 440, 445
eversmanni, Perognathus, 492
evexus, Dipodomys, 502
evexus, Thomomys, 520
evidens, Sciurillus, 318
exalbidus, Sciurus, 332
excelsus, Cratogeomys, 529
txi:eanus, Paraxerus, 406, 409
exiguus, Liomys, 478
exilis, Dipodomys, 496, 501
exilis, Myoprocta, 197
exilis, Nannosciurus, 315
exilis group, Nannosciurus, 315
eximius, Dipodomys, 49S
eximius, Thomomys, 521
extenuatus, Thomomys, 514
extimus, Citellus, 452
extimus, Sciurus, 333
extimus, Thomomys, 521
falcifer, Thomomys, 523
fallax, Callosciurus, 364
failax, Geomys, 527
fallax, Perognathus, 490
falzfeini, Scirtopoda, 592
famatina, Abrocoma, 154, 155
faniatinae, Lagidium, 230, 232
famosus, Ctenomys, 163, 166
famulus, Callosciurus, 351, 370
fasciatus, Perognathus, 483
fasciatus group, Perognathus, 482, 483
fasciculata, Athcrurus, 206
favonicus, Funambulus, 378, 380
favonicus, Jaculus, 595, 597
fedjushrini, Sciurus, 332
felicia, Dasyprocta. 194
felipensis, Orthogeomys, 532
felix, Tamias, 432
felli, Ratufa, 384, 388
FELOVIA, 34, 68. 554, 555, 559
femoralis, Perognathus, 483, 491
femoralis, Ratufa, 388
ferminae, Sciurus, 341
ferreus, Sciurus, 338
ferrugineiventris, Sciurus, 334
ferrugineus, Callosciurus, 351, 352, 361
ferrugineus, IVIesomys, 126, 127
festina, Cavia, 240, 241
fiber, Castor, 466, 468
fiber group, Castor, 468
filchncrinae, Petaurista, 289
fimbriatus, Eoglaucomys, 298
finlaysoni, Callosciurus, 351, 352, 361
fisheri, Citellus, 442, 451
fisheri, Tamias, 436
tisheri, Thomomys, 523
rtammifer, Sciurus, 328, 344
flavescens, Allactaga, 586
Havescens, Citellus, 442
flavescens, Dasyprocta, iqi. 193
flavescens. Perognathus, 4S3
flavidus, Echimys, 109, 113
flavidus. Galea. 242, 243
flavidus, Thomomys, 518
flavimanus, Callosciurus, 351, 352, 359
flavinus, Funisciurus, 412, 413
flavinus, Marmota, 457, 460
flavior, Drcmomys, 381
tlaviventer, Microsciurus, 319, 321
flaviventris, Glaucomys, 297
flavivcntris, Marmota, 455, 457, 458
flaviventris group, Marmota, 455, 458
666
flavivittis group, Paraxerus, 406, 40S
flavivittis, Paraxerus, 406, 40S
fiavus, Castor, 468
fla%us, Perognathus, 483, 484
flavus, Sicista, 568
flavus. Xerus, 421
fleminci, Hystrix, 217
rtorenciae, Microsciurus, 320
florentiae, Jaculus. 595, $gy
floridanus, Geomys, 526
floweri, Callosciurus, 351. 362
fluminalis, Callosciurus, 351, 363
tochi, Ctenomys, 163, 165
fodax, Ctenomys, 163, 164, 167
folletti. Callosciurus. 351, 361
formosanus, Callosciurus, 351, 354
formoso\i, Sciurus, 332
formosus, Peroenathus, 4S3, 4S7
formosus i^roup, Perof^nathus, 482, 4S7
FORNARINA, 80. 94
fornicatus, Lariscus, 392
fornicatus. Perognathus, 48H
forresti, Callosciurus. 351, 355
forresti, Sciurotamias, 426
fortior, Anomalurus. 540
fortirostris. Mamiota. 458
fossor, Sciurus, 337
fossor, Thomomys, 522
fossor group, Thomomys. 310, 522
fosteri, Cercomys, 123, 124
fournieri, Capromys, 129
foxi, Cryptomys, Ny, 91
foxi, Graphiurus, 607, 609
frandseni, Callosciurus, 351, 361
franklinii, Citellus, 441, 442, 450
franziusi, "Octodon." 159
fraseri, Anomalurus. 540
fraseri group. Anomalurus, 540
fraseri, Sciurus. 328, 343
frater, Ctenomys, 163, 167
frater, Tamias. 430, 433
fraterculus. Callosciurus, 351, 357
fremonti, Tamiasciurus, 346, 348
frenatus, Dipodomys, 501
frerei, Paraxerus, 406, 408
fretensis, Ratufa. 3S4. 389
frondator. Castor, 467
frontalis, Ratufa, 388
frumentor, Sciurus, 328, 334
frutectanus, Napaeozapus, 574
fr\anus, Callosciurus. 351, 360
fueginus, Ctenomys. 163, 167
fulgens, Anomalurops, 542
fulgida, Cavia, 240, 242
fuliginosa. Dasyprocta, 191, 193, 195
fuliginosus, Glaucomys, 29(5
fuliginosus, Perognathus, 484
fuliginosus, Proechimys, 119
fuligmosus, Sciurus, 333
fulminatus, Sciurus, 32S, 345
fulvescens, Cratogeomys, 530
tulvinus, Petaurisla, 282, 288
fulvior, Xerus, 420, 422
INDEX
fuKus, Castor, 468
fuhus, Citellus, 442, 443
fulvus group, Citellus, 440, 443
fulvus, Ctenomys, 163, 167
fulvus, Cuniculus, 225
fulvus, Sciurus, 333
ful\'us, Thomomys, 517
fulvus group, Thomomys, 510, 517
fumigatus, Callosciurus, 363
fumigatus, Sciurus, 344
fumosus, Platygeomys, 530, 531
fumosus, Thomomvs, 512
FL'NAMBULUS,'3o. 49, 58, 65. 263, 266,
267. 271, 3ii, 312, 349, 376, 380
FUNISCIURUS. 30. 57, 67. 174, 264, 267,
269, 272, 309, 410
fusca, Geomys, 526
fuscatus, Heteromys, 47')
fuscoater, Sciurus, 328, 330
fuscocapillus, Petinomys, 300, 301, 302
fuscocapillus group, Petinomys, 301, 302
fusconigricans. Sciurus, 328, 331
fuscorubens, Sciurus. 328. 331
fusco\ariegatus, Sciurus, 336
fusculus, Microsciurus, 320
fuscus, Aconaemys, 158
fuscus, AJlactaga, 586
fuscus, Dremomys, 381, 382
fuscus, Glaucomys, 297
fuscus, Thomomys, 524
fuscus group, Thomomys, 510, 524
fuscus, Xerus, 420
gabrielsoni, Dipodomys, 499
GALEA, 29. 71, 23S, 239, 240. 242
galeata, Hystrix, 200, 216, 217, 220
galliae. Castor, 468
gallicus. Castor, 468
gambianus, Heliosciurus, 401
ganana, Paraxerus, 406, 408
garbei, Sciurus, 343
garonum. Dremomys, 381
gaumeri, Heteromys, 474, 476
gazellae, Paraxerus, 406. 409
geayi, Procapromys, 133
genibarbis, Petinomys, 301
genibarbis group, Petinomvs, 301
GEOCAPROMVS, 23. 54', 73, 128, 131, 134
GEOMVIDAK. 32, 2=;=?. 469, 505
GEOMYOIDAE, 31, 78, 468
GEOMYS, 22, 32, S3, 63, 71, 1S9, 506, S07,
524
GEORYCHUS, 22, 24, 67, 80. 81. 82, 83
GEOSCiURL'S. 264, 267, 418, 420
GERBILLINAE, 40
GERBILLUS, 22, 40, 50, 56
gerboa, Jaculus, 596
germaini, Callosciurus, 351, 352, 362
gerrardi, Sciurus, 327, 329, 341
getulus, Atlantoxerus, 423
GIDLEY. See Miller
gigantea, Ratufa, 384, 388
INDEX
667
giganteus, Spalax, 642, 644
gigas, Dinomys, 171
giglis, Glis, 622
gilvigularis, Sciurus, 342
gilvus, Perognathus, 484
gingianus, Callosciurus, 368
ginninianus, Xcrus, 422
glaber, Hetcrocephalus, f)$
GLAUCOMYS. 30. 52. 53. 54, 61. 70, 263,
266, 268, 276. 293, 294, 297, 299, 335
giaucinus. Sciurillus, 317, 318
glirinus, Zenkerella, 547
gliroides, Octodontomvs, i sy, I'io
GLIRISCUS, 604. 606. 608
GLIRULUS, 37, 60, 601, 603, 612. 613. 620
GLIS, 22, 37, 46, 51, 60, 601, 602, 603, 612,
613,620
glis, Glis, 333, 622
gloveri, Callosciurus, 351, 358
GLYPHOTES, 30, 49, 65, 263, 271, 305. 393
goeldii, Proechimys, 118, 121
goelthasi, Hoplomys, 123
goldmani, Citellus, 451
goldmani, Cratogeomys, 529
goldmani, Dipodomys, 498
goldmani, Glaucomys, 295
goldmani, Heteromys, 474. 475
goldmani, Perognathus, 489, 490
goldmani, Sciurus, 328, 336
goldmani, Thomomys, 521
GOLUNDA, 38, 48. 50
goodfellowi, Ctenomys, 163, 164, 168
gordoni, Callosciurus, 351, 358
gordoni, Jaculus, 595, 596
gorgonae, Proechimys, 118, 120
gorkhali, Petaurista, 282, 284, 288
gossei. Funambulus. 379, 380
gothardi, Sciurus, 330
gracilis, Cavia, 242
gracilis, Tamia^, 433
gradojevici, Citellus, 442, 444
graeca, Sciurus, 330
graecus, Spalax, 643, 644
grahamensis, Thomomys, 518
grahamensis, Tamiasciurus, 348
GRAMMOMYS, 38, 602
grammurus, Citellus, 4^2, 450
grandis, Echimys, iii. 113
grandis group, Echimys, iii, 113
grandis, Orthogeomys, 531, 532
grandis. Petaurista, 282, 286
GRAOMYS, 39
GRAPHIL'RINAE, 36. 603
GRAPHIURL'S. 34. 36, 44, 45, 46, 56, 69,
564, 599, 601, 604
gravipes, Dipodomys, 499
grayi, Callosciurus. 351, 374
gregorianus, Thr\onomys, 148
gregorianus group, Thr\onomvs, 144, 145,
148
grinnelli, Tamias, 435
grisea, Aplodontia, 258
griseicauda, Callosciurus, 351, 367
griseicollis, Ratufa, 387
griscifrons, Glaucomys, 297
griseimanus, Callosciurus, 351, 352, 363
griseimembra, Sciurus, 341
griseiventer, Callosciurus, 368
griseiventer, Petaurista, 286
griselda, Anomalurus, 540
grisclda, Dremomys, 381
griselda, Graphiurus, 607, 611
griseocaudatus, Sciurus, 335
griseoflavus, Sciurus, 328, 335
griseogena, Sciurus, 328, 340
griseopectus, Callosciurus, 351, 358
grisescens, Allactaga, 587
grisescens, Tamias, 431
griseus, Graphiurus, 610
griseus, Heteromys, 475
griseus, Sciurus, 325, 327, 328, 337
griseus, Tamias, 437
grotei, Hystrix, 217
grutei, Callosciurus, 362
guadalupensis, Thomomys, 514
guairae, Proechimys, 118, 121
guanta. Cunicuius, 224, 225
guardiae, Perognathus, 492
guayanus, Sciurus, 328, 343
gubari, Pteromys, 294
guentheri, Trichys, 205
guerrerensis, Liomys, 479
guerrerensis, Orthogeomys, 532
GUERLINGUEIUS, 265, 266, 268, 322.
325, 326, 340
guianae, Cavia, 240, 241
guianae, Echimys, in, 112
guianensis, Sciurus, 342
guiara, Eury-zygomatomys, 125
guillemardi, Callosciurus, 369
gularis, Dremomys, 381, 383
gularis, Proechimys, 118, 120
gundi, Ctenodact>'lus, 558
gundlachi, Capromys, 129
gunnisoni, Cynomys, 462, 463
gunong. Callosciurus, 351, 356
guttatulus, Citellus, 445
guttatus, Citellus, 445
g>-mnesicus, Eliomys, 615, 616
gy-mnicus, Tamiasciurus, 346
GYMNOBELIDEUS, 543
GYMNUROMYINAE, 39
GYMNUROMYS, 39, 57, 603, 645
g>mnurus, Hoplomys, 122, 123
g>mnurus, Plats'geomys, 530
GYOMYS, 38 '
g\-psi, Perognathus, 484
HADROMYS, 38, 48
HADROSCIURUS. 265, 270, 322, 323, 326.
327. 344
haedulus, Graphiurus, 607, 611
haemobaphes, Callosciurus, 359
H.AEROMYS, 38
hageni, Petinomys, 301, 302
668
INDEX
hageni ^Toup. Pt'tinom\s, 301, 302
haiKJ. Ctcnomys, 163, 1(15
hainana, Pt-taurista, 289
hainana, Ratufa, 384, 3SS
hainanus, Atherurus, 206
hainanus, Callosciurus, 354
halli. Dipus, i^tji
HALTiCUS, 591
halticus, AUactatja, 5S6
halticus, Dipodomvs, 500
HALTOMYS. 59^
hamiltoni, Kliomvs, Oi"^
HAPAI.OMYS. viii, 3-. 36. 3S, 48. 63^
HAPTOMVS, r6i, 164
hardyi, Htliosciurus, 401, 403
haidyi, Zapus. 571
harmandi, Callosciurus, 351, 362
harquahalae, Thomomys, 517
harrinctoni, Callosciurus, 351, 360
harrisi, Citellus, 442, 451
harrisoni, Callosciurus, 366
harrisoni, Hylopetes. 299
harrisoni, Thryonomys, 14S, 149
hartcrti, Massouticra, 559
hastilis, Callosciurus. 331, 364
heemianni, Dipodomys. 4^7, 498
heermanni ^'roup, Dipodomys, 497, 498
heermanni, Sclurus, 337
heinrichi, Hyosciurus, 398
helyci, Callosciurus, 3^1 "i
HELIOPHOBICS, 24, s7, 67. 80. 81, 82.
84, 555
HELIOSCIURUS, 30. 57, 67. 264, 266, 267.
269, 271. 310, 319, 321, 327. 346, 399
hellenicus, Spalax. 642, 643, 644
htlleri, Dipodomys, 504
htlleri, PeroKnathus, 4S9
hulleri, Thomomys, 524
helveolus, Sciurus. 337
helvina, Ahrocoma, 154
helvus, Callosciurus, 365
hemachalanus, Marmota, 460
hendcei, Callosciurus, 351, 359
hendt-ei, Proechimvs, 118. 121
HEPTAXODONtlDAK, 17
herberti. Callosciurus, 3*)i
herbicola, Citcllus, 442, 443
hercego\inensis, Spalax, 643
hernandezi, Sciurus, 329, 334
HERPETOMVS. 41
herreranus, Glaucomys, 295
HESPEROMYS, 39. S5
HESPEROSCIURUS, 2(m. 268, 321, 325,
337
hcspcrus, Thonionivs, ^24
HETEROCKPHAIJ, 24. 81, o^
HETEROCEPHALCS. 24, 57. 57. 80, Si, 94
heterodus. Macrogeomys, 534
HETEROGEOMYS, 32, 71^ sob, 507, 532
HETEROMYIDAE. n, 470
HETEROMYINAE, ii.471
HETEROMYS, 31, 54. 55, 70, 4^9. 47°. 47'.
472
HETEROSCIURUS, 348
heterothrix, Liomys, 478
hicmalis, Sciurus, 333
hilda, Proechimys. 118, 121
hinialaicus, Tro^opterus, 279
himalayana, Marmota, 457, 460
hindei, Lophiomys, 636
hindsi, Tamias. 430, 434, -135
hintoni, Petaurista, 282, 289
hippurellus, Callosciurus, 351, 374
hippurosus, Callosciurus, 351, 374
hippurus, Callosciurus, 351, 353, 374
hippurus group, Callosciurus, 353, 374
hirsutirostris, Hystrix, 200, 216, 217, 218
hirsutus, Sigmodon, 114
hirtipes, Jaculus, 596
hirtus, Cratogeomys, 529
hirtus, Sciurus, 335
hispidus, Heterogeomys, 533
hispidus, Liomys, 477
hispidus, Mesomys, 126. 127
hispidus, Perognathus, 480, 483, 488
hispidus group, Perognathus, 483, 4S8
historicus, Sciurus, 333
HISTRIOSCIURUS,^322, 325
hodgsoni, Marmota, 460
hodgsoni, Hystrix, 200, 201, 209, 214, 217
HODOMYS, 39, 54
hoffmani, Sciurus, 340
hoffmani group. Sciurus, 327, 340
hollistcri. Citcllus, 449
HOLOCHILUS, 39, 55
holoserictus, Cr\'ptomys, 87, 90
holti, Callosciurus, 351, 355
hoodi, Citellus, 448
hopicnsis, Perognathus, 484
hopiensis. Tamias, 433
HOPLOMYS, 2s. S4. 72, 102. 107, loR, 117.
122
horsfieldi, lomys, 303
hortualis, Eliomys, 615
hosei. Lariscus, 391, 392. 393
hosei group, Lariscus, 393
hosei, Petaurillus. 302
hotsoni, Allactaga, 582, 584, 585
hottentotus. Cr\ptomys, 87, 90
hottentotus group, Cr\ptomys, 90
howulli, Dremomys, 381, 382
howelli, Thomomys, 514
huachucha, Sciurus. 339
hualpaiensis, Thomomys, 314
hudsomcus, Tamiasciurus, 340
hudsonis, Erethizon, 181
hudsonius, Glaucomys, 296
hudsonius, Zapus, 571
hueti, Graphiurus, 601. 606, 607, 608
hueti group, Graphiurus, boS
hueyi, Perognathus. 488
humboldtiana, Aplodontia, 259
humei, Callosciurus, 351, 365
humeralis. Ratufa, 385
humilis, Callosciurus, 357
hungaricus, Spalax, 642, 643
INDEX
669
HYBOMYS, 38
hvdrochacris, Hydrochoerus, 253
HVUROCHOEklNAE. 29. 238. 249
HYDROCHOERUS. 23, 29, S4. 55, 72, 189,
250, 466
HYDROMYINAE, 39
HYDROMYS, 3. 36. 39, 47, 602. 627
hvlaeum, Plagiodontia, 137, 134
HYLENOMYS, 38
HYLOPETKS, 30, 49, 52, 64, 263, 276, 293,
298
HYOSCIURUS, 30, 65, 271. 305, 398. 423
HYOMYS, 38, 47
HYPER.ACRILTS, 40
hypervthrus, Callosciurus, 358
HYPOGEOMYS. 39. 57
hypoleuca, Ratufa, 387
hypoleucus, Citellus, 443
hypophaeus, Sciurus, 334
hypopyrrhus, Sciurus, 324, 334
hyporrhodus. Sciurus, 328, 340
hvpoxanthus, Sciurus, 334
HYSTRICES, 27, 208
HYSTRICIDAK. 27. 98, 100, 173. 197
HYSTRICOGNATHI, 23
HYSTRICOMORPH SERIES, 24
HYSTRICOIDAE, 24. 77, 96, 237
HYSTRIX, 23, 27, 49. S2. 57, S9, ^6, 69, 14s,
198, 208,209, 212,215, 218
ibeanus, Lophiomys, 636
ibeanus, Paraxerus, 406, 409
ICHTHYOMYS, 3, 36, 39, 55, 602
ictericus, Callosciurus, 351, 371
ICTIDOMOIDES. 437
ICTIDOMYS, 268, 437, 440. 441, 448
idahoensis, Citellus, 447
idahoensis, Perognathus, 486
idahoensis, Thomomys, 522
idahoensis, Zapus, 572
IDIURINAE, 33, 542
IDIURUS, 33. 57. 68, 537, 542. 543
ignava, Marmot.), 457
Ignitus, Sciurus, 344
igniventris, Sciurus, 328, 344
iheringi, Proechimys, 116, 117. irS, 122
iheringi group, Proechimys, 122
ileile, Hyosciurus, 399
illinoensis, Geomys, 526
imarius, Callosciurus, 351, 373
imhausi, Lophiomys, 636
imitator, Callosciurus, 351, 363
immoUis, Lagostomus, 236
imperator, Anomalurus, 540
imperator, Zapus, 573
imus, Dremomys, 381. 382
inauris, Xerus, 422
inca, Lagidium, 230, 232
incanus, Pteromys, 294
inconstans, Callosciurus, 351, 355
inconstans, Sciurus, 329, 341
incultus, Rhinosciurus, 398
indica, AUactaga, 584, 585
indica, HystrLx, 218
indica, Ratufa, 350, 384, 385
indicus, Funambulus, 378
INDOMALAYAN RODENTS, 64, 65, 66
inermis, Cercomys, 124
inexpectatus, Callosciurus, 363
inexpectatus, Perognathus, 487
infraluteus. Perognathus, 483
infrapallidus, Thomomys, 512
infuscatus, Sciurus, 328, 330
ingens, Dipodomys, 499
ingens, Thomoniys, 513
ingrahami, Geocapromys, 132
ingrami, Sciurus, 328, 343
innae, Dipus, 591
inopinus, Perognathus, 491
inornatus, Callosciurus, 373
inornatus, Petaurista, 282, 284, 288
inornatus, Perognathus, 483, 486
inquinatus, Callosciurus, 351, 374
insidiosus, Coendou, 185, 186, 188
insignis, Lariscus, 392
insignis group, Lariscus, 392
insignis, Napaeozapus, 574
insignis, Ratufa, 384, 386
insignis, Zenkerella, 547
insularis, Callosciurus, 359
insularis, Citellus, 452
insularis, Dipodomys, 501
insularis, Glis, 622
insularis, Perognathus, 488
insularis, Rhizomys, 650, 651
insularis, Spalax, 642, 643
intercessor, Tamias, 430, 436
interior, Funisciurus, 412, 413
intermedius, Bathyergus, 84
intermedius, Callosciurus, 358
intermedius, Citellus, 443
intermedius, Dyromys, 618
intermedius, Glis, 622
intermedius, Perognathus, 490
intermedius group, Perognathus, 483. 490
intermedius, Sciurus, 336, 358
intermedius, Spalax, 642
intermedius, Thomomys, 518
intensus, Xerus, 420, 421
internatus, Thomomys, 514
internus, Graphiurus, 607, 609
interpres, Citellus, 452
interposita, Ratufa, 388
interstriatus, Sicista, 567
interzonus. Sicista, 567
inyoensis, Tamias, 430, 433
lOMYS, 30, 49, 64, 263, 274, 275, 303
IRENOMYS, 39, 55
irolonis, Cratogeomys, 528
irroratus, Liomys, 477, 478
irroratus group, Liomys, 476, ^78
irroratus, Sciurus, 328, 344
isabella, Funisciurus, 412
isabellinus, Heliosciurus, 401, 403
ISCHYROMYIDAE, 14
670 INDEX
ISOBOLODON. 169
isolatus, Graphiurus, 610
ISOTHRIX. 23, 25. 72, 106. loS. 113
isthmica. Dasyprocta, iqi, 195
isthmicus. Heterogeomys, 533
isthmius, Hydrochoerus, 253
isthniius, Liomys, 477
isthmius, Microsciurus, 319, 320
istrandjae, Sciurus, 331
istricus, Citellus, 444
istricus, Spalax, 644
italicus, Glis, 622
italicus, Sciurus, 328, 330
jacinteus, Thomomys. 515
jacksoni, Anomalurus, 540, 541
jacksoni. Paraxerus. 406. 408
jacksoni, Pero^jnathus, 486
jacksoni. Tamias, 432
JACULUS, 22, 3s,' SI. 56. S9, 68, s=;s. S62,
589, 593. 598, 638
jaculus, Allactaga, 584, 586
jaculus, Jaculus, 593, 595, 596
jaculus group, Jaculus, 595, 596
jacutensis, Citellus, 442, 446
jacutensis, Sciurus, 332
jacutensis, Tamias, 436
jaliscensis, l>iomys, 479
jalorensis, Lariscus, 392
janifsoni, Cnptomys, 92
janett.i, Bathyergus, 84
janetta, Callosciurus, 351, 373
japonicus. Glirulus, 620
javanicuni, Hystrix, 199, 209, 211, 213, 214,
217
javanicus, Rhizomys, 65!
javanus, Lariscus, 392
javcnsis, Ratufa, 385
jenissejensis, Sciurus, 332
jentinki, Callosciurus, 350, 351, 356
jesupi, Heteromys, 475
joannia, Caviella, 246
joannius, Octomys, 157
johannis, Ctenomys, 163, 166
johnstoni, Graphiurus, 607, 610, 611
johorensis, Callosciurus, 368
johort-nsis, Ratufa, 384, 387
joleaudi, Ctenodactylus, 559
joloncnsis, Dipodomys, 498
jordani, Anomalurus, 540, 341
jordani, Graphiuru?, 611
jorisseni, Cryptomys, 87, 90
juglans, Citellus, 450
junipcrus, Tamias, 433
junodi, Cryptomys, 92
juraiis, Sciurus, 328, 345
juris, Ctenomys, 163, ibh
juvencus, Callosciurus, 351, 353, 375
kaffcnsis, Heliosciurus, 401, 402
kahari, Paraxerus, 407
kaibabensis, Sciurus, 338
kalaharicus, Paraxerus, 407
kalbinensis, Sciurus, 332
kaleensis, Belom\s, 277, 279
kalinowskii, Dasvprocta. 196
KANNABATKOMYS, 25, 73. 102. 135, 137
kapiti, Heliophobius, 85
karclini, Scirtopoda, 592
kathlecnae, Funambulus, 380
kazakstanicus, Citellus, 444
kelaarti, Funambulus, 378, 380
kellcni, Graphiurus, 611
kemmisi, Callosciurus, 360
kenaiensis, Tamiasciurus, 347
keniae, Heliosciurus, 401, 403
kennicotti, Citellus, 448
keraudreni, Callosciurus, 361
kermiti, Proechimys, 119
kernensis, Dipodomys, 500
kernensis, Tamias, 43 s
KERODON, 29, 55, 71, 224. 238, 239, 246
kessleri, Sciurus, 332
kinneari, Callosciurus, 351. 358
kingii, Caviella, 246
kinlochi, Petaurillus, 302, 303
kinoensis, Perognathus, 485
kirgisorum, Spalax, 642
kivuensis, Idiurus, 54b
kizljaricus, Allactaga, 585
klagcsi, Sciurus, 328, 340
klamathensis, Glaucomys, 297
klossi, Callosciurus, 368
klossi. Hystrix, 200, 213, 214, 217, 218
knekus, Perognathus, 488
knighti, Ctenomys, 163, 167
kodiacensis, Citellus, 442, 448
komatiensis, CrA'ptomys, 92
kongensis, Callosciurus, 351, 354
kootenayensis, Zapus, 572
koratensis, Menetes, 390
kozlo\i, Salpingotus, 575
kozlo\i group, Salpingotus, 575
kubangensis, Cryptomys, 91
kuchingensis, Callosciurus, 351, 366
kuhli, Sciurillus, 318, 342
kurdistaniLus, Dyromys, 619
kummi, Cr\ptomys, 87, 91
labaumei, Spalax, 643
labilis. Diplomas, 115
labradorius, Zapus, 571
LACHNOAIYS. 136, 137
lachupuilla, Thomomys, 519
lacnmalis, Cratogeomys, 529
lacr\'malis, Thomom\s, 512
lacustris, Xerus, 420, 421
ladas, Zapus, 37 1
lademanni, Hystrix, 220
laenata, Ratufa, 384, 38O
laenatum, Coendou, 145, 1S5, 187
laetus, Paraxerus, 40(7
LAGJDICM, 23, 28, 55, 72, 226. 229
INDEX
671
LAGOMORPHA. i
lagopus, Dipus, SQi
LAGOSPEDIUS, 247
LAGOSTOMI. 28, 227, 232
LAGOSTOMUS, 23. 28. 55. 72. 97» t7i
224. 226. 233
LAGOTIS. 229
LAGURUS, 41, 50. 52
laingi, Perognathus, 487
lalandianus, Graphiurus, 610
lamarum. Echimys, 109, iii, H2
Iambi, Perognathus. 492
lamucotanus, Callosciurus, 371
lancavensis, Callosciurus, 351, 364
langi. Cr\'ptomys, 92
langi, Idiurus, 546
langsdorffi, Sciurus, 328, 345
laniger, Anomalurops, 542
laniger, Chinchilla, 229
lanius, Heterogeomys, 533
lanka, Petaurista, 282, 287
lanuginosus, Tamiasciurus, 347
laomache, Dremomys, 381, 383
LAOMYS. 38
laotum, Callosciurus, 351, 355
laotum, Hvlopetes, 299, 300
LARIA, 391
LARISCUS, 30, 49, 65, 260, 263, 267, 271
306, 349, 380, 391
Iar\alis, Pedetes, 552
lascivus, Glaucomvs, 297
LASIOPODOIVn'S. 41
lasiotis, Glirulus, 620
LASILROMYS, 113
lasiurus, Platacanthomys, 629
lastii, Paraxerus, 406, 408
lateralis, Citellus, 442, 453
lateris, Heliosciurus, 401
laticaudatus, Rhinosciurus, 396
laticeps, Allactaga, 584, 585
laticeps, Anomalurus, 540
laticeps, Clyomys, 125
laticeps, Thomomys, 511
latifrons, Orthogeomys, 507, 532
latijugularis, Perognathus, 492
latimaxillaris, Dipodomys, 503
latipes, Glaucomys, 296
latirostris, Perognathus, 488
latirostris, Thomomys, 517
latouchei, Rhizomys, 650, 651
latrans, Cynomys, 462
latro, Ctenomys, 163, 165
latus, Thomomys, 514
laurentius, Cercomys, 123, 124
lautensis, Callosciurus, 371
layardi, Funambulus, 377, 379, 380
layardi group, Funambulus, 379
layardi, Petinomys, 301, 302
leathemi, Sicista, 566, 568
lechei, Cr>ptomys, 87, 91
lectus, Tamias, 431
legerae, Pectinator, 556
LEGGADINA, 38, 47
43
LEMMI, 40
LEMMUS. 40, 50. 52
lemniscatus, Funisciurus, 412
k-mniscatus group, Funisciurus, 412
LEMNISCOMYS, 38
Icna, Petaurista, 282, 286
leniceps, Mcsomys, 126, 127
LENOMYS, 38
LENOXUS. 39
lentulus, Ctenomys, 163, 165
lentus, Dremomys, 383
leo, Rhinosciurus, 396, 398
leonardi, Hylopetes, 299, 300
leonensis, Heliosciurus, 401, 403
leonis, Funisciurus, 412, 413
leonis, Sciurus, 328, 341
leonurus, Zapus, 571
lepidus, Hylopetes, 299
lepidus, lomvs, 303
LEPORIDAE, I, 18
LEPORILLUS, 38, 47
leporina, Aplodontia, 258
leporina, Dasyprocta, 194
leporinus, Sciurus, 337
leptodactjius, Spermophilopsis, 425
LEPTOMYS, 39. 47
LEPTOSCIURUS, 265, 322, 325
leptosoma, Proechimys, 119
leptura, Myoprocta, 196
lepturus, Heteromys, 475
lerotinus, Eliomys, 615, 616
leschenaulti, Ratufa, 385
lessonii, Xerus, 421
leucoblephara. Galea, 243
ieucocephalus, Callosciurus, 362
leucocephalus, Petaurista, 286
LELCOCROSSUROMYS, 268, 461, 462
leucodon, Ctenomys, 163, 164, 167
leucodon, Spalax, 644
leucodon, Thomomys, 511
leucodonta, Castor, 467
leucogaster, Sciurus, 334, 344
leucogenys, Dipodomys, 499
Icucogenys, Funisciurus, 414
leucogenys, Petaurista, 282, 285, 288
leucogenys, Ratufa, 384, 389
leucombrinus, Xerus, 420, 422
leucomus, Callosciurus, 306, 350, 351, 353,
375
leucomus group, Callosciurus, 353, 375
leucomystax, Proechimys, 119
leucopictus, Citellus, 445
leucops, Geor\'chus, 86
leucops, Sciurus, 334
Icucopus, Callosciurus, 351, 363
leucopyga, Cavia, 241
leucostictus, Citellus, 446
leucostigma, Funisciurus, 412, 413
leucotis, Callosciurus, 351, 354
leucotis, Dipodomys, 504
leucotis, Sciurus, 328, 334
leucourus, Sciurus, 328, 330
Icucura, HystrLx, 200, 212, 213, 217, 218
67=
INDEX
leucura group, Hystrix, 212. 216
leucurus, C'itcllus, 442. 452
ieucurus, Cynomys, 462
leurodon. Citcllus, 446
levaillanti, Xerus, 422
levipes, Dipodomys, 496. 504
lewisi, Ctcnomys, 163. 164, 167
lewisii, Cynomys, 462
liantis, Callosciurus, 355
libtricus, Heliosciurus, 401, 404
lichicnsis, Dremomys, 381, 3S2
lichtinsteini, Eremodipus, 598
liebnianni, Coendou, 187
lighti, Callosciurus, 372
lilacus, Sciurus, 328, 330
limanus, Myoprocta, 19ft, I97
limhatus, Heliosciurus, 401
limitancus, Xerus, 420, 421
limitaris, Thomomys, 519
limitis, Sciurus, 339
limosus, Thomomys, 523
lineatus, Sicista, 567
lineatus, Tamias, 430, 436
lingungensis, Callosciurus, 357
LIOMYS, 31, •;4. 62. 7°. 469. 47°. 47 1. 47^,
476
lipura, Trichys, 199, 203, 205
lis, Sciurus, 324, 328, 333
littledalei, Marmota, 457, 460
littoralis, Galea, 242, 243
littoralis, Graphiurus, 611
littoralis, Sciurus, 335
littoralis, Tamias, 434
litus, Perognathus, 483
llanensis, Geomys, 527
loandac, Protoxerus, 415, 4>6
loandius, Heliosciurus, 401, 402
lockwoodi, Lagidium, 230, 231
locusta, Jaculus, 596
loftusi, Jaculus, 595. 597
logonensis, Thryonomys, 149
lokriah, Dremomys, 380, 381
lokriah group, Dremomys, 381
lokroides, Callosciurus, 351, 353. 372
lomitensis, Heteromys, 475
LON'CHERES, 108
LON'CHOTHRIX, 25, 73. 102. i°7. 127
longicauda, Hystrix, 200, 217
longicaudatus, Coendou, 186
longicaudatus, Heteromys, 474, 475
longicaudatus, Proechimys, 118, 121
longimcmbris, Perognathus, 485
longimembris group, Perognathus, 482, 485
longior, Allactaga, 587
longipes, Dipodomys, 502
longipes, Zapus, 571
longipilis, Cavia, 242
longirostris, Echimys, 109, 113
lonnbergi, Cannomys, 652
lonnbergi, Hvstrix, 220
LOPHIOMYIDAE, 37, 600, 632
L(JPH1<)IMYS, 22, 36, 37, 56, 69, 599, 632
LOPHUROMYS, 38, 46, 5b
loquax, Tamiasciurus, 346
lordi, Perognathus, 483, 487
loriger, Sicista, 566, 567
loringi, Thomomys, 524
lorraineus, Graphiurus, 607, 610
lowei, Callosciurus, 351, 357
lowei group, Callosciurus, 351, 357
lualabae, Heliosciurus, 401, 404
lucas, Callosciurus, 364
lucidus, Microdipodops, 493
lucidus, Thomomys, 513
lucifer, Dasyprocta, 191, 193
lucifer, Heliosciurus, 400, 401, 405
lucifer group, Heliosciurus, 405
lucifugus, Glaucomys, 297
ludibundus, Tamias, 432
ludovicianus, Cynomys, 462
iudovicianus, Sciurus, 339
ludwigi, Cryptomys, 90
lugardi, Cryptomys, 87, 91
lugens, Petinomys, 301, 302
lumhoitzi, Petaurista, 2S6
lumholtzi, Ratufa, 387
luncefordi, Nannosciurus, 316
lunaris, Callosciurus, 372
lunaris, Dasyprocta, 191, I94
lunaris, Paraxerus, 406, 409
lusitanicus, Eliomys, 615, 616
lutea, Lagidium, 230, 231
luteiventris, Tamias, 432
luteolus, Dipodomys, 502
luteola, Marmota, 458
luteolus, Ctenomys, 163, 164, 168
lutescens, Callosciurus, 371
lutescens, Funambulus, 379, 380
lutescens, Geomys, 526
lutescens, Lagidium, 231
luteus, Zapus, 573
lutrina, Ratufa, 384, 388
Ivchnuchus, Tamiasciurus, 348
Ivlei, Callosciurus, 351, 355. 362
lylei, Petaurista, 282, 284, 287
lylei, "Tamiops," 351, 355
lyratus, Citellus, 448
lysteri, Tamias, 430, 436
macconnelli, Sciurus, 328, 342
maclellandi, Callosciurus, 351, 354
maclellandi group, Callosciurus, 353
macmillani, Dremomys, 381
MACROGEOMYS, 32, 54, 7"
macrorhabdotes, Tamias, 435
MACROSPALAX, 638, 641
macrospilotus, Citellus, 449
macrotarsus, Jaculus, 596
macrotis, Allactaga, 586
macrotis, Glaucomys, 296
macrotis, Idiurus, 544, 546
macrotis group. Idiurus, 546
macrotis, Rheithrosciurus, 395
macrotis, Thomomys, 522
macrotis, Trichys, 203, 205
506, 507, 533
INDEX
macroura, Ratufa, 378, 384
macroura, Sciurus, 339
macrourus croup. Athcrurus, 2o5
macrourus, Athcrurus. 199, 206
macrourus. t.'itcilus. ^42. 4S0
macrourus, Procchimvs, 119
.MACROXUS, 322
macrura. Echimys, 112
macruroides. Ratufa, 384, 388
MACRL'ROMVS, 38, 47
maculatus, Hciiosciurus, 401, 403
madogac, Hciiosciurus, 401, 402
madrensis, Citellus, 454
madrensis, Glaucomys, 296
madrensis, Thomomys, 520
madsoci, Callosciurus, 368
madurae, Callosciurus, 351, 369
macnas, Caviella, 246
maerens, Petinomys, 302
macstus, Xcrus, 420, 421
magdalcnac. Ferot'iiathus, 491
niagdaicnac, Sciurus, 342
magdalenae. Thomomys, 511;
magellanica, Dolichotis, 248
magcllanicus, Ctcnomys, 163, 164, 165
magnicaudatus, Sciurus, 339
magnificus, Petaurista, 282, 284, 288
magrudercnsis, Perognathus, 487
mahali, Cryptomys, 92
major, Allactaga, 584, 586
major Kroup. Allactaga, 584, 585
major, Aplodontia, 258, 259
major, Citellus, 449
major, Ratufa, 385
major, Zapus, 572
makkovikensis, Glaucomys, 296
malabarica, Hystrix, 218
malabarica, Ratufa, 385
malaccanus, Petinomvs, 301
MALACOMYS, 38 '
MALACOTHRIX, 36, 39, 56
malawali, Callosciurus, 369
MALLOMYS, 38, 47, 48
managuensis, Sciurus, 336
manarius, Microsciurus, 319, 321
manavi, Sciurus, 340
mandingo, Funisciurus, 412, 414
manipurensis, Callosciurus, 3SI, 354
mansalaris, Callosciurus, 356
mantchuricus. Sciurus, 328, 331
maporensis, Callosciurus, 3^1, 370
mapravis, Callosciurus, 351, 364
marabutus, Xerus, 421
marana, Ratufa, 384, 389
maraxica, Dasyprocta, 191, 194
marchio, Petaurista, 282, 285
marcoscnsis, Perognathus, 492
margaritae, Dipodomys, 501
margaritae, Perognathus, 492
marginatus, Citellus, 449
marica, Petaurista, 282, 284, 286
mariposae, Tamias, 435
marinsularis, Callosciurus, 371
673
maritimus, Bathyergus, 84
maritimus, Callosciurus. 3^1, 3154
marjoriae, Petromus, i^i
MARMOTA, 23, 30. 40. 51, 53, 54, ,8, 62,
66, 262, 265, 267, 268, 273, 274, 307, 454
marmota, Marmota, 455, 457, 461
marmota group, Marmota. 455. 461
" Marniotiiiae," 261, 262
MARMOTOPS, 267, 268, 454, 455, 494
martensi, Sciurus, 328, 331
martinensis, Thomomys, 520
martirensis, Dipodomys, 503
martirensis, Thomomys, 515
marungcnsis, Heliophohius, 85
marwitzi, Heliosciurus, 402
masae, Ratufa, 384. 388
massonii. Ctenodactvlus, 558
MASSOUTIERA, 34, 50, 56, 58, 68, 554,
MASTACOMYS, 38, 47
matagalpae, Macrogeomys, 534
matagalpae, Sciurus. 337
matthaeus, Callosciurus, 363
matugamensis, Funambulus, 379, 380
maulinus, Ctenomys, 169
maunensis, Paraxerus, 406
mauritanicus, Jaculus. 596
maurus, Sciurus. 334
maximus, Perognathus, 488
maxima, Ratufa, 384, 385
maximus, Lagostomus, 236
mayensis, Dipodomys, 501
mayumbicus, Funisciurus, 412
mazama, Thomomys, 524
mearnsi, Perognathus, 483
meamsi, Tamiasciurus, 348
mearnsi, Thomomys, 519
mearsi, Callosciurus, 351, 373
mechowi, Cryptomys, 87, 90
mechoui, group, Cryptomys, 90
medellinensis, Sciurus, 328, 343
media, Allactaga. 586
medius, Echimys, 109, 1:1, 112
medjianus, Heliosciurus, 404
megacephalus, Microdipodops, 493
MEGAI.OMYS, 39
mcgalops, Zapus, 571
MEGASCAPHEUS, 507, 510, 524
melania, Sciurus, 328, 337
melanochra, Ratufa, 384. 385
melanogaster, Callosciurus, 351, 353, 374
melanoleucas, Heteromys, 474, 475
melanopipla, Ratufa, 384, 389
mclanops, Callosciurus, 365
melanops, Thomomys, S23
melanopterus, Petaurista, 285, 289
melanoticus, Cryptomys, 92
melanotis, Nannosciurus, 315, 316
melanotis group, Nannosciurus, 316
melanotis. Perognathus, 484
melanotis, Thomomys, 517
melanotis, Sciurus, 328 339,
melanotis, Petaurista, 282, 285
674
INDEX
mflaiiunis, Capruiiixs, 129
inelaiuirus, Coendou, 185, 1S6, 18S
nielaniirus, Dipodoniys, 4g(>, 501
melnnurus, Kliomys, 615, Uih
nielaniirus. Tamias. 41,1
mcllandi. C'r\ptomys, 44. .S7, 90
melli. Drt'momvs, -^H-^
MKLOMVS, 38
mclonii. Glis. 622, 623
menamiciis, C'aIloscii.irus, 351, 361
mendanaiius. C'nllosciurus, 366
niendociTia, C'tcnoniss, 1(13, 16=;
MENKTKS. 30, 4(K (»s, 2<-M. 267, 271, 308,
380. 390
mentawi. Callosciurus, 351, 374
mentosus. Drcmomys, 38 1. 382
mt'nzbieri. Mamiota, 461
mergulus, Pctaurista, 2S2. 2S7
mendioccidt-ntalis, Citt-llus, 445
meridcnsis. Sciurus. 32S. 340
meridionalis, Lariscus, 392
mt-ridionalis, Pectinator, 55^1
meridionalis, Sciurus. 330
meridionalis, Tamias, 415
MERIONKS, 40. 50
merriami, Crato^^com>s, 528
merriami, Dipudunus, 496, 497, 500
merriami tiroup, Dipodomys, 497, 500
merriami, Peroj:inathus, 4S3
merriami, Tamias, 430, 435
mersinae, Hystrix, 218
mesatia, Dasyprocta, iQf>
mesemhnnus. Pc'rot,niathus, 4S7
MESKMURIOMYS, 3S, 47
MESOCRICETUS, 22, Vh si
MESOMYS. 25, 55. 73, 102, 107. n?, »26
mesopolius, Perognathus, 491
mesopotamica, Hvstrix, 219
MESOSCIURl'S, 26=;, 268, 322, 32=;. 326
MESOSPALAX. (138, 640, 641, 642
meticulosus, Callosciurus, 351, 365
mewa, Thomomys. 311
mexicana, Dasyprocta, 194
mexicanum, Coendou, 145, 182, 1S5, 187
mexicanum group, Coendou, 185, 187
mexicanus. Castor, 4O7
mexicanus, Citellus, 441. 442. 449
mexicanus, Cynomys, 4(12
mexicanus, Geom\'s, 52b
mexicanus, Pcroynathus, 484
mexianae, Cuniculus, 225
michianus, Callosciurus, 351. 359
michiganensis. Castor, 4^-
micklemi, Cryptom>s, 87, 91
microctphalus, Zapus, s7i
MICR()DII.LL!S, 40 ■
MICRODIPODOPS, 1, II, =1^,62.469.470,
471,479. ^^92
microdim. Xerus, 420. 422
MICR(.)MYS. 38, 48. 50
microphthalmus, Spalax, 642, 644
microps group, Dipodom\s, 497, 504*
microps, Dipodoniys, 5C4
micrnrhvnchus. Callosciurus, 3!;i. 368
MICROSPALAX. bjiH. 641
MICROSCICRUS. 30. S4, (IS. 69. 265. 26O.
268, 270, 310. 319
MICROTI. 40
MICROTINAK. 36. 40
microtis, Callosciurus, 369
microtis, Graphiurus, 607, 610
microtis, Jaculus, 596
MICROTUS. 41, 'so. 32, 54. 60:;
MICROXUS, 39
midas, Callosciurus, 351, 360
migratorius, Sciurus, 333
MILLARDIA, 38. 48
millardi, Callosciurus. 351, 360
MILLER, Classification of Order, 13, 14, 15,
16, 17
milleri, Callosciurus. 351, 364
niilleri, Dyrom\s, 619
milleri, Myoprocta, 196, 197
milleri, Sciurus, 328, 341
millsi. Hystrix, 214. 217, 218
mimax, Octom\'s, 157
mimellus, Callosciurus, 367
mimicus, Petaurista, 286
mimiculus, Callosciurus. 367
niimuius, Microsciurus, 319, 320
mimus. Sciurus, 338
minax, Trogopterus, 279, 2S0
mincae, Proechimys, 118, 120
mmdanensis, Callosciurus. 351, 374
mmiatus, Callosciurus, 351. 372
minimus, Galea, 243
minimus, Perognathus, 48<>
minimus, Tamias, 430
minimus group, Tamias, 430
minnesota, Tamiasciurus, 346
minor, Alactagulus, 587
minor, Cannomys, 652
minor, Liom\'s, 478
minor. Thomomys. 511
minor, Zapus, 572
minutulus, Myosciurus. 313
minutus, Alactagulus, 587
minutus, Ctenom\s, ibS
minutus, Glis, 623
minutus. Myosciurus, 313
mira\'allensis, Sciurus, 340
missouriensis. Castor, 467
missouriensis, Cynomys, 462
mitchelli, Dipodomys, 301
mitratus, Citellus, 454
mohileiisis, Geomys, 326
moco, Kerodon. 247
modestus, Callosciurus, 336
modestus, Dact\lomys, 137
modestus, Dremomys, 381, 382
modicus, Thomom>s, 518
moerens, Protoxerus. 413, 416
moerescens, Menetes. 390. 391
mogollonensis, Tamiasciurus. 348
inohavensis, Citellus, 442, 432
niohavensis, Dipodom\s, 49i(
INDEX
675
mohavensis, Thomomys, 515
mohcius, Callosciurus, 351, 365
mohillus, Callosciurus, 351, 365
moi, Callosciurus, 351, 355
mollcndorffi, Callosciurus, 375
molliac, Isothrix, 114
mollipilosus, Pcroutiathus, 486
mollipiiosus, Tamiasciurus, 347
mollis, Citc'llus, 442, 446
molyneauxi, Cr>'plomys, 87, yi
momonga, Ptcromys, 293, 294
monardi, Graphiurus, 607, 609
monardi uroup, Graphiurus, 6og
monax, Mamiota, 455, 457
monax group. Marmota. 455, 457
mongolica, AllactaRa, 584, 587
mongolicus, Citellus, 442, 445
monoensis, Dipodomys, 502
monocnsis, Tamias, 432
monocnsis, Thomomys, 523
montana, Sicista, 566, 567
montaiius, Cryptomys, 92
montanus, Dipodomys, 502
montanus, Zapus, 573
MONTICAVIA, 238. 239, 243. 244. 246
monticola, Citellus, 449
monticola, Perognathus, 486
monticola, Spalax, 642
monticola, Thomomys, 508, 510, 523
monticola group, Thomomys, 510, 523
monticolus, Callosciurus, 351, 354
montosus. Graphiurus, 607, 608
morda\, Trogoptcrus, 279
mordosus, Ctcnomys, 163, 167
moreni, Lagidium, 230, 232
moricandi, Chaetomys, 172
morio, Sciurus, 334, 344
moroccana, Hystrix, 219
morroensis, Dipodomys, 499
morli\'allis, Dipodomys, 500
morulus, Sciurus, 341
mossambicus, Paraxerus, 406, 409
mottoulei, Hcliophobius, 86
mouhotei, Menetes, 390
mowewensis, Callosciurus, 375
mugosaricus, Citellus, 442, 443
mulleri, Hystrix, 200, 212, 217
multicolor, Heliosciurus, 401
munbyanus, Kliomys, 615, 616
mural is, Thomomys, 515
MURKS, 38
MURICCLUS, 38
Ml'RIDAi:, 38, 599. 600
MURINAK, 38
murinus, Graphiurus, 607, 610
murinus group. Graphiurus, 609
murinus, Sciurillus, 49, 317, 318
murinus group, Sciurillus, 318
MUROIDAE, 35, 78. 599
murravi. Abrocuma, 154
MUS.'22, 38, 47, 4X, 50, 52, 56
ML'SCARDIMDAK, 36, 599. 600
MUSC.\RDININAE. 37, 603. 612
MUSCARDINUS, 22, 37, 51, 60, 601, 602,
612, 613, 623, 627
muscardinus, Muscardinus, 625
musculinus, Heliosciurus, 404
musculus. Thomomys, 521
musicus, Citellus, 444
mustelinus, Sciurus, 334
musteloides, Galea. 242
mutabilis, Heliosciurus, 401, 402
mutabilis, Paraxerus, 40')
mutabilis, Thomomvs, 519
MYCTEROMYS, iS
MYLAGAULIDAE, i^, 17. 2^8
MYLAGAULUS, 258
MYLOMYS. 38, 56
MYOCASTOR, 3, 23, 26, 42, 55, 73. 97. 98,
103, 123, 129, 134, 140, 145
MYOCASTORINAE, 25, loi, 102, 104, 139
MYOMIMUS, 36, 37, 60, (03, 612, 613, 626
MYOMORPH SERIES, 32
MYOPOTAMUS, 140
MYOPROCTA, 27, 55. 74, i^JO. 196
myops, Erethizon, 178, 181
myops, Thomomvs, 524
MYOPUS, 40, 50
MYOSCALOPS, 84
MYOSCIURUS, 30, 57. 67, 260, 264, 267,
269. 305. 312
MYOSPALACINAE. 40
MYOSPALAX. 3. 10, 36, 40, 51, 81, 637
myosuros, Proechimys, 119
MYOXUS, 620
MYRSILUS, 30, 67, 264, 272, 30H, 416
mvstax, Eunisciurus, 412, 414
MYSTROMYS. 39, 56
mzabi, Massoutiera, 559
nadymensis, Sciurus, 328, 331
nagarum, Callosciurus, 351, 358
nagtgiasi, Graphiurus, 6c8
nakanus, Callosciurus, 351, 364
namaquensis, Xerus, 420, 422
nana, Capromys, 128, 129, 131
nana, Cavia, 240, 242
Nannosciurinae, 13, 262
NANNOSCICRUS, 30, 49. 55, 57, 64. 262,
263, 267, 269, 306, 312, 313, 317
NANNOSPAL.A>:, 638. 640, 641, 642
NANOCAVIA, 238, 239, 243, 244, 246
nanogigas, Ratufa, 384, 387
nanus, Graphiurus, 607, 611
nanus, Thomomys, 513
napaea, Sicista. 566
NAPAEOZ.APUS, 35, 53, 63, 568, 573
napi, Microsciurus, 319, 320
narynensis, Hystrix, 219
nasicus, Thomomys, 524
naso, Euchoreutes, 577, 579
nasutus, Thomomys, 513
natalcnsis. Cr>plomys, 92
nattereri, Ctenomys, 169
natuncnsis, Callosciurus, 351, 357
676
INDEX
navigator. Callosciurus, 351. 367
na\us, Thomornys, 511
na\aritensis, Sciurus, 32S, ^vj
NHACOMVS. 4. 39
XEARC-TIC RODENTS. (,o. 61. 62, 63
nea\'ci. Anomalurus, 540, 541
nebouxi, Sciurus, 328, 343
nebrascensis, Glaucomys, 295
nebulicola, Citellus. 44S
nebulosus, Thomomys. 522
necopinus, Sciurillus, 31S
NECTOMYS. 39. 54 "
neglectus, Citellus, 453
neglectus, Ctenomys. 165
neglfctus, Ptrounathus. 48(1
neglectus, Platygeomys, 331
neglectus, Sciurus, 338
neglectus. Tamias. 430. 431
neglectus, Thomomys, 515
negligens, Sciurus, 337
negrensis. Galea, 242, 243
negrensis, Isothnx, 114
nehringi, Spalax. f'4^
NELOMYS, loS, 109
nelsoni, Citellus, 442, 452
nelsoni, Cuniculus, 225
nelsoni, Dipodomys, 500
nelsoni, Heteromys, 474, 476
nelsoni, Pappogeomys, 52S
nelsoni. Orthogeomys, 332
nelsoni. Perognathus, 490, 491
nelsoni, Sciurus, 32N, 335
nelsoni, Thomom\'S, <^2i
NELSONIA, 39. '54'
nemo, C'r>'ptomys, 93
nemoralis, Sciurus, 328, 334
nemoralis, Zapus, s7i
NEOCTODON, 1^9
NEODON, 41. 4S, 30
NEOFIBER, 41, s^'
neomexicanus, Tamiasciurus, 34S
NEOSCIL'RUS, 264, 267, 26S. 321. 324, 333
NEOTAMIAS, 268. 426. 428. 429, 430
NEOTOMA. 36. 39, 52, :;4
NEOTOMODON, 39
NEOTOMYS, 39. 3=;
NEOTROPICAL R(.)DENTS. 69, 70, 71.
72. 73. 74
nesaeus, Sciurus, 32S, 341
nesiotes, Callosciurus, 331. 370
nesioticus, Citellus, 431
NESOKIA, 38. 43, 4S, so
"NESOMYIDAE," 41
NESOMYS, 39, 57. 643
NESOROMYS, 38, 47
ne\adensis, Citellus, 447
nevadensis. Dipodom>s. 300
ne\;Klcnsis, Perognathus, 48O
ne\adensis, Tamias, 433
nevadensis, Thomom\s, 511
nevadensis, Zapus, 572
nexilis, Dipodomys, 503
iiexu^. Tamias, 434
niata. Caviella, 246
niapu, Funisciurus, 414
nicotianae, Callosciurus, 369
nicoyana, Sciurus, 336
niger. Castor, 468
nigeriae, Protoxerus, 415, 416
niger, Sciurus, 328, 330, 338
niger, Thomomys, 523
nigra, Aplodontia. 239
nigra, Dasyprocta. 191, 105
nigra, Marmota, 461
nigratus, Sciurus, 344
nigrensis, Anomalurus, 340
nigrensis, Funisciurus, 412, 414
nigrescens, Erethizon, i8i
nigrcscens, Liomys, 477, 478
nigrescens, Ratuta. 388
nigrescens, Sciurus, 32S, 330, 334
nigriana, Caviella. 246
nigricans, Allactaga, 386
nigricans, Cavia, 242
nigricans, Coendou, 186, 188
nigricans, Dasyprocta, 195
nigricans, Thomomys. 312
nigncaudatus, Petaurista. 283
nigriceps, Ctenomys, 163, 164, 168
nigriclunis, Dasyprocta, 193
nigridorsalis, Callosciurus, 359
nigrimontis, Perognathus, 490
nigripes, Hylopetes, 29Q, 300
nigripes, Sciurus, 328, 337
nigrispina, Echimys, 112
nigrovittatus, Callosciurus, 331. 332. 36S
nikkoms, Petaurista, 282, 28S
nikolskii, Citellus, 444
nimrodi, Cr\'ptomys. 87, 90
ningpoensis, Callosciurus, 331, 338
niobe, Lariscus, 392
nitedula, Dyromys, 61S, (>i9
nitida, Petaurista, 283
nitidula, Petaurista, 282, 283
nitratoides, Dipodomys, 496. 497, 501
nitratus, Dipodomj's, 300
ni\aria, Marmota. 459
niveatus, Funisciurus. 412, 413
niveus, Muscardinus, 625
nobilis, Petaurista, 282, 288
nobliei, Dasyprocta, 193
noctivagus, Spalacopus, 161
nogai, Dipus, 391
nordhotfi, Protoxerus, 415
nordmanni. Sicista, 567
norvegica, Sicista, 566, 567
norvegicus, Rattus, 47
nosophora, Marmota, 438
notabilis, Protoxerus, 413, 4i<>
notabilis, Ratufa, 386
notatus, Callosciurus, 349, 331, 332, 368
notatus group, Callosciurus, 352, 368
notialis, Hvdrochoerus, 253
XOTIOMYS, 3, 36. 39. 55
notioros, Marmota, 438
NOTOCITELLL'S,"268, 437. 440. 441, 451
INDEX
677
NOTOMYS. 3. 36. 38. 47
NOTOSCIURUS. 265, 322, 325. 326, 344
novachispaniae, Coendou, 187
novcmlineatus, Callosciurus, 354
nox, Callosciurus, 351, 362
nuchalis, Dasyprocta, 195
nuchalis, Sciiirus, 335
nudipes, Citellus, 451
numantius, Sciurus, 328, 330
numarius, Funambulus, 379, 3H0
nyansac, Hcliosciurus, 401, 404
nvcthemera, Coendou, 185, 187
NYCTOCLKPTES, 645, 646, 649, 651
NYCTOMYS. 39, 54
nyx, Callosciurus, 367
obfuscatus, Hcliosciurus, 401, 403
obolenskji, Dyromys, 6iy
obscura, Echimys, 109, 113
obscura, Marmota, 458
obscurus, Citellus, 445
obscurus, Dipodomys, 502
obscurus, Funambulus, 380
obscurus, Galea, 243
obscurus, Lariscus, 392
obscurus, Liomys, 477
obscurus, Mesomys, 127
obscurus, Perognathus, 490
obscurus, Tamias, 435
obsidianus, Citellus, 450
obsolctus, Citellus, 450
occasius, Echimys, 109, iii, 112
occidanca, Hystrix, 217, 219
occidentalis. Callosciurus, 375
occidentalis, Dinomys, 171
occidcntalis, Eliomys, 615, 616
occlusus, Cr\'ptomys, 90
occultus, Ctenomys, 163, 164, 166
occultus, Perognathus, 491
OCHOTONIDAE, i
ochracea, Marmota, 457
ochraceocinereus, Cryptomys, 91
ochraceus, Paraxerus, 406, 407
ochraceus, Proechimys, 119
ochraccus, Tamias, 432
ochraspis, Pctaurista, 282, 286
ochrescens, Sciurus, 328, 344
ochrogaster, Funisciurus, 412, 414
ochrogenys, Tamias, 430, 434
ochrus, Perognathus, 491
ocius, Thomomys. 522
o'connulli, Proechimvs, 119
OCTODON. 23,26,'55,73. i^s. 156,158,^^4
OCTODONTINAE, 26, gS.^ioi. 104. 15*5,
169
OCTODONTOMYS, 26. 73. i55. 156, 159
OCTOMYS. 26, 55, 73, 155, 156, 161, 554
ocularis, Graphiurus, 606, 607, 608
oculatus, Sciurus, 328, 339
OENOMYS, 38, 56
oenone, Funisciurus, 412, 413
ognevi, Dyromys, 619
o^'nevi, Pteromys, 294
ognevi, Sciurus, 332
okadae, Tamias, 436
okanagana, Marmota, 457, 459
olga, Graphiurus, 607, 609
olivaceus, Callosciurus, 351, 355
olivaccus, Citellus, 449
olivaceus, Hcliosciurus, 404
olivaceus, Perognathus, 487
olivellus, Funisciurus, 412, 413
oliviae, Funisciurus, 412, 414
oiympica, Aplodontia, 258
olympicus, Glaucomys, 297
olympius, P'unambulus, 378, 380
olympus, Mannota, 459
omensis, Hcliosciurus, 401, 402
ONDATR.A, 3, 41, 52
ONYCHOMYS, 39, 52. 53, 54
operarius, Tamias, 431
operarius, Thomomys, 517
ophinsae, Eliomys, 615, 616
opimus, Ctenomys, 163, 164, 168
opimus, Dremomys, 381, 383
optabilis, Thomomys, 514
opulentus, Thomomys, 514
oral, Pctaurista, 282, 287
oralis, Jaculus, 597
orangiae, Cryptomys, 93
orangiae, Pedetes, 551, 552
orarius, Tamiasciurus, 347
orarius, Zapus, 573
orbitalis, Liomys, 477
ordii, Dipodomys, 496, 497, 502
ordii group, Dipodomys, 497, 502
ordinalis, Tamias, 430, 436
oreas, Pctaurista, 282, 2S8
orcgonensis, Glaucomys, 296
oregonus, Citellus, 442, 447
oregonus, Microdipodops, 493
oregonus, Thomomys, 523
oregonus, Zapus, 572
oreocetes, Cratogcomys, 529
oreocetes, Tamias, 431
oreoecus, Thomomvs, 517
OREOSCIURUS, '322, 324
orestes, Callosciurus, 351, 36S
oresterus, Heteromys, 476
orienlalis, Anomalurus, 540, 541
orientalis. Athcrurus, 206
orientalis, Glis, 623
orientalis, Jaculus, 595, 596
orientalis group, Jaculus, 595, 596
orientalis, Tamias. 430, 436
orientis, Sciurus, 328, 332
orii, Pteromys, 294
orinoci, Isothrix, 114
oris, Proechimys, 118, 121
orizabae, Thomomys, 519
orlovi, Citellus, 444
omatus, Dipodomys, 500
omatus, Dremomys, 381, 382
ornatus. Paraxerus, 406, 408
orobinus, Graphiurus, 607, 609
678
INDEX
ORTHOGI-:OMYS, 32, 71, ^06, =;o7. 531
ORTHRIOMYS, -|i
ORVZOMVS. V). =;^. S4
ORYZORICTES. S
osgoodi, Citellus, 44S
osgoodi, Ctenom\"s, 16S
osgoodi, Thoniomys. 516
otinus, Microsciunis, 319, ^20
OTOMYINAK. ^g
OTOMYS, iQ. ^'b, 60:;
OTOSPKRMOPHILCs. 265. 26S, 4:^7, 43g,
440. 441. 450
OTOSCIURUS. 264. 26S. 322. 325, 337
OTOTYLOMYS. 39. 54
o\\ensi, Callosciurus, 351. 373
owstoni, Dremomya, 3S1, 3S2
owstoni, Sciurotamias, 426
oxianus, Citellus. 442. 443
OXYMYCTERL'S, 39. 46. 55
oxylona, IMarmota, 459
paca, Cuniculus, 224, 225
paca group, Cuniculus, 225
pacarana, Dinomys, 171
pachita, Proechim\s, iiS, 121
pachvura, Isothrix, 114
PACHYUROMYS. 22. 40, 50. 493
pacifica, Aplodontia. 25S, 259
pacificus. Castor. 467
pacificus, Perognathus, 486
pacificus, Zapus, 573
padangensis, Rhizomys, 651
pagurus, Isothrix. i 14
PALAK ARCTIC RODENTS, 57. 58, 59, 60
PALAEOCASTOR. 466
palatmus, Zapus, 572
paleacea. Echimys, iir, 113
palki, Cryptomys. 93
pailasi, Taniias, 435
pallasi, Spalax, 644
pallescens, Citellus, 449
pallescens, Sciurus, 328, 333
pallescens, Thom(jm\s, 512
palliata, Ratufa, 3S4, 386
palliatus, Paraxerus, 406, 40S
palliatus group, Paraxerus, 406, 40S
pallida, Cavia, 240
pallida, Sicista, 567
pallidicauda, Citellus. 443
pallidior, Cavia, 241
pallidior, Caviella, 246
pallidior, Kannabateornys, 137
pallidulus, Dipodomys, 49S
pallidus, Alactagulus. 5SS
pallidus. Capromys, 129
pallidus. Citellus. 442, 444. 449
pallidus, Coendou, 185, 186, 1S8
p.illidus, Lryptomys. 93
pallidus, Ctenomys, 169
pallidus, Dyromys, 619
pallidus, iiliomys, 615. 616
pallidus, Heliophobius, 85
pallidus, Microdipodops, 493
pallidus, Nannosciurus. 316
pallidus, Octodon, 159
pallidus, Perognathus, 490
pallidus, Tamias, 431
pallipes. Lacidium, 230, 232
palmarum, Fuiianihulus, 377, 37S, 3S0
palmarum group. Funambulus, 37S
palmeri, Dipodomvs, 502
palmeri, Microsciurus, 319, 320
palmed, Tamias. 433
palustris, Callosciurus. 3(16
palustris. Galea. 242. 243
pamparum, Cavia, 240. 241
pamparum, Lagostomus, 236
panamensis, Heteromys, 476
panamensis, Prnechiinys, iiS, 120
panamintinus, Dipodomys, 499
panamintinus, Tamias, 430, 432
panamintinus, Perognathus, 483, 4S5
pandora, Dasyprocta, 191, 195
panga, Idiurus, 546
panjioli, Callosciurus, 351. 364
panjius, Callosciurus. 351. 364
pannosus, Rhizomys, 630, 651
pannovianus, Callosciurus. 370
pansa, Ma;rogeomys, 534
papae, Hvstrix, 218
PAPPOGEOMYS. :!2, 71. S07. 527
PARADIFUS, 3s. si. S9. 589
PARADOLICHOTiS, 247, 24S
paradoxus, Cardiocranius, 576
paradoxus, Perognathus, 488
paraensis, Sciurus, 329, 343
parayayensis, Coendou, 182, 185, 1S6, 188
paragayensis group, Coendou, 186, 188
paraguaNensis, Dasvprocta, 191, 194
PARAHYDROMYS, 39. 47
paralius, Liomvs, 477
PARAMYIDAE. is
PARAMYS, 17
PARAPEDETES, 5s i
PAR.\SCIURL'S, 2f>5. 267, ^^S. 322, 325,
327. 33!^
PARAXERLTS, 30, 57, 67. 262, 264, 267, 271,
309. 3^0, 400, 405, 412
PAROTOAIYS, 39. 5t
parry'i, Citellus, 440, 442, 448
parryii group, Citellus, 447
parthianus. Citellus. 442. 443
par^a, Myoprocta. 197
parviceps, Liomvs, 477
par\'iceps, Thomomys, 521
parvidens, Citellus, 449
par\'idens, C)nomys, 463
parvula, Marmota, 458
parvulus. Citellus, 451
parvulus, Graphiurus. 612
par\us, Callosciurus, 3sf>
parvus, Citellus, 449
parvus, Dipodomys, 500
par\*us, Graphiurus, 607, 609
piir\us, Perognathus, 483, 486
INDEX
679
par\us group. Perognathus, 482, 486
pascalis, Thomomys, 512
pasha, Heliosciurus, 401, 403
patachonica, Dolicholis, 248
patagona, Dolichotis, 248
pater, Cannomys, 652
pauli, Paraxcrus, 407
pearsoni, Belomvs, 277
PECTINATOR, 23, 33, 34, 56, 68, 553. 554,
555, 560
pectoralis, Petaurista, 289
pectoralis, Thoniomvs, 519
PEDETIiS, 21, 22, 33. 43, 56, 68, 144, 548, 549
PEDETIDAE, 33, 547
PEDETOIDAE, 33, 78. 547
PEDO.MYS, 41
PEDIOLAGLS, 247
peiapis, Callosciurus, 366
pelii, .\nomalurus, 540, 541
pelii group, Anomalurus, S4i
PELOMYS, 38
pemangilensis, Callosciurus, 351, 370
pemangilis, Atherurus, 206
pembertoni, Callosciurus, 354
pembertoni, Eunisciurus, 412, 413
penangensis, Pc-taurista, 289
penangensis, Ratufa, 384, 389
penialius, Callosciurus, 366
penicillatus, Funambulus, 378
penicillatus, Perognathus, 488
penicillatus group, Perognathus, 483, 488
peninsulae, Citellus, 442, 452
peninsulae, Lariscus, 392
peninsulae, Perognathus, 483, 491
peninsulae, Ratufa, 389
peninsularis, Callosciurus, 351, 370
peninsularis, Dipodomys, 503
peninsularis, Menetes, 390, 391
pennanti, E'unambulus, 377, 379, 380
pennipes. Citellus, 452
peracer, Rhinosciurus, 396
pcramplus, Thomomys, 518
perblandus, Dipodom\s, 500
perci\ali, Paraxerus, 406, 407
perditus, Thomom\-s, 521
peregrinator, Sciurus, 334
peregrinus, Cratogeomys, 529
peregrinus, Thomomys, 519
pergracilis, Perognathus, 486
perhentiani, Callosciurus, 351, 372
pericalles, Perognathus, 485
pcridoneus, Cratogeomys, 530
perlutea, Lagidium, 230, 231
pemiger, Perognathus, 483
pemix, Ctenomys, 169
pemix, Perognathus, 483, 489
pernyi. Dremomys, 381
pemvi group, Dremonivs, 381
PERODIPLS, 494
PEROGNATHINAE, 470
PEROGNATHI, 31, 471, 479
PEROGNATHUS, 31, 53, 54, 62, 70. 469,
470. 47 1 , 479. *80, 482, 493
54
PEROMYSCLS, 39, 51
perotensis, Citellus, 450
perotcnsis, Cratogeomys, 529
perotensis, Dipodomys, 500
perpallidus, Citellus, 453
perpallidus, Thomomys, 510, 51 ^
perpallidus group, Thomomys, 510,
perpes, Thomomys, 516
perplanus, Cratogeomys. 529
perplexus. Dipodomys, 503
perrcnsi, Ctenomys, 163, 166
persicus (Glis), 623
persicus, Sciurus, 333
personatus, Geomys, 526, 527
personatus, Graphiurus, 610
personatus, Myomimus, 626
personatus, Protoxerus, 416
peruana, Octodon, 159
peruanum, Lagidium, 230, 232
peruanus, Dact\lomys, 136, 137
peruanus, Microsciurus, 320
perustus, Anomalurus, 540, 54!
penagus, Thomomys, 518
pessimus, Citellus, 446
PETAURILLUS, 30, 64, 263, 274
PETAURISTA, 23, 30, 49, 52, 57,
263, 274, 275, 281
petaurista, Petaurista, 282, 285
petaurista group, Petaurista, 284, 285
"PHTAURISTIDAE," 264
PETAUROIDES, 543
PETAURUS, 543
petilidpns, Lagostomus, 236
PETINOMYS, 30, 49, 64, 263, 275,
petrensis, Marmota, 457
PETROMUS, 13, 23, 26, 33, 42, 57
102, 103, 149, 554
PETRO.MYINAE, 26, 101, 104, 149
PETROMYSCUS, 39
petulans, Tamiasciurus, 347
phaea, Aplodontia, 2S9
PHAENOMYS, 39 "
phaeognathus. Citellus, 448
phaeomelas, Aeromys, 291
phaeopepla, Ratufa, 384, 389
phaeopus, Sciurus, 336
phaeurus, Liomys, 477
phaeurus, Sciurus, 338
phaeus. Castor, 467
PHAIOMYS, 41
PHALAXGERIDAE, 542
phalaena, Paraxerus, 406. 407
phanrangis, Callosciurus, 351, 361
phasma, Perognathus, 490
phasma, Thomomys, 518
phayrei, Callosciurus, 351, 353, 373
pha\Tei, Hylopetes, 299, 300
phelleoecus. Thomomvs, si7
PHENACOMYS, 41, '52 "
philippinensis, Callosciurus, 351, 353,
philippcnsis, Petaurista, 282, 284. 286
phillipsii, Dipodomys, 497, 49S, 500
phillipsii group, Dipodomys. 500
275, 302
64, 262,
93, 300
68, 98,
374
6So
INDEX
phillipsi. Hetcrocephalus, 94, 95, gO
phipsoni. Petinomvs, 301
PHLOKOMVS, 36. 38. 4S. 60'.
PHODOPUS. 22. 39. 51
phrvgius. Dvromvs, 61S, 611;
PHYLLOIVIYS. loS. log
PHVLLOTIS, 39. 55
picatus, Tamiasciurus, 347
piccus. Callosciurus. 351, 3'^>7
picinum. Erethizon, iSi
pictus, Dyromys, 610
pictus, Isothrix, 1 14
pictus group, Isothri\. j 14
pictus, Liomys, 477
pictus group. Liomys, 476, 477
pictus, Tamias, 431
piLTrei, Callosciurus, 351, 361
pilorides, Capromys. I2Q
pilosus americanus, Krcthizon, 181
pinetorum, Thomonns. 524
pinetis. Geomys, 526
pinicatae, Perognathus. 400
piniensis, Callosciurus, 357
piniensis, Ratufa, 3S4, 38S
piperi, Perognathus, 484
pipidonus. Callosciurus, 351. 364
piraia. Callosciurus, 151, 360
PITHECHEIR, 38. 48
PITVMYS. 41, 50. 52. 54
piutensis, Thomomvs, =; i s
PLAGIODONTIA. 2s, S4, 71, 102, 104, 133
PLAGIODONTIINAE, 25. loi, 133
planict'ps, Platygeomys, 531
planicola, Citellus, 442, 444
planifrons, Cratogeomys, 530
pla irostris, Thomomys, 516
plantani, Callosciurus, 36S
plantinarensis, Liomys, 477
plasticus, Callosciurus, 351, ^72
PLATACANTH{;MViNAE. 37.601, 603. 626
PLA'PACANTHOMYS. 4, 36. 37, 4S; h6.
601, 602, 603, 626, 627, 632
platyccntrotus, Coendou, 185, 187
platvcL'phaius, Dipodomvs. soi
PLATYCERCOMYS, 5S8 '
PLATYCRANIUS, 604, 606
PLATYGEOMYS, 3:, 71. 506. 507, 528, 530
platyops, Graphiurus, 607, (108
platyops group. CJraphiurus, O07
platyurus, Hylopetcs, 2yi>
piatyurus, Pygeretmus, 588
platyurus group, Pygcrctmus. 5S8
plesius, Citellus, 448
plumbcscens, Cannoni>s, 652
pluto, Orthogeomys. 532
pluto, Callosciurus, 351, 3()7
podolica. Spalax, 644
poensis, Hehosciurus, 400. 401, 404
poensis group, Heliosciurus, 404
poeppingi, Caproni>s. i2<j
poeppigi, Spalacopus. idi
pOLvi, Capromvs, 129
POGONOMYS, 38. 47
pohlei. Castor, 468
polia, Ratufa, T,Hy
POLIOCITELLCS. 26S, 437. 441, 450
polionotus. Microdipodops, 41)4
poliopus, C.'llosciurus, 372
poliopus, Proechimys, iiy
poliopus, Sciurus, 334
polonicus, Spalax, 644
pontifex, Ctenomys, if^-?. 165
pooli, Funisciurus, 412
popelairi, Myocastor. 141
porcellus, Cavia, 240. 242
porteri, Aconaemys, 158
porteousi, Ctenomys, 163, 1A6
portus, Callosciurus. 351, 361
postus. Glis, 622
potanini. AlactaHulus, 58S
potosinus. Thomomys, 520
prachin, Callosciurus, 37^
praetextus, Funisciurus, 413
pranis, Callosciurus, 363
pratensis, Citellus, 450
pratti, Myoprocta, iq6
preblici, Dipodomys. 504
prebliei, Tamiasciurus, 347
preblorum, Marmota. 457
prehensilis, Capromys. 129
prehensilis, Coendou, 185, 186
prehensilis group. Coendou. 182, 185, 18^)
premaxiliaris, Thoniom\'s, =124
prestigiator, Xerus, 421
pretiosus, Callosciurus, 369
pretiosus, Liomys, 478
pretoriae. Cryptomys, 93
pretoriae, Graphiurus, 611
PREVIOUS CLASSIFICATIONS OF
SCIURIDAE, 261, 262, 263, 264, 265.
266, 267, 26S
prevosti. Callosciurus, 321, 349, 351, 365
prevosti group, Callosciurus, 352, 365
pricei, Perognathus. 4S3, 489
pricei, Tamias, 430, 435
prietac, Perognathus, 492
primrosei, Petaurista. 282, 287
princeps, Xerus, 420, 422
princeps, Zapus. 572
PKIONOMYS. 39, 56
pnttwitzi. Hystnx, 219
probus, H^'lopetes, 299, ^00
PROCAPROMYS. 25, 73. 132
procerus, Callosciurus, "^56
PROECHIMYS. 25,54^55.72. 107. loS. 115,
123
PROEOROMYS. 41
progrediens, Heterocephalus. 95
PROMETHEOMYS. 3. 36, 4!, 50. 81
propinquus, Tamias, 432
proprius, Castor, 46S
proserpinae, Callosciurus, 366
proteus, Callosciurus, 351, 372
PROTOPTYCHINAE, 16
PROTOXERUS, 30, 44. 57. f.7. 262. 264.
2(17, 272, 308, 415, 417
INDEX
68 1
providentialis, Thomomys, 517
proximus, Scirtopoda, 592
proximus, Thomomys, 519
pruinosus, Coendou, 185, 186, 188
pruinosus, Marmota, 459
pruinosus, Rhizomys, 650, 651
pryeri, Callosciurus, 351. 353, 374
prymnolopha, Dasyprocta, 191, 193
pryori, Thomomys, 522
psakastus, Heturornvs, 475
PSAMMOMYS, 40, 50
PSEUDOCHIRUS, 543
PSEUDOSCILRIDAK, 15
PSEUDOMVS. 38
"PTKRDMVINAE," 13. 262
PTEROMVES. 30, 260, 261, 273
PTEROMYS, 30. 45, 52. 57, 262, 263. 268.
274. 276, 291, 293, 298
PTEROMY'SCUS. 30. 64. 263. 275. 280
puchcrani, Sciurus. 329, 343
puchcrani yroup, Sciurus, 327, 343
puertae, Thomomys, 512
pulcher, Muscardinus, 625
pulcher, Nannosciurus, 315, 316
pullus, PtTognathus, 492
puiverulcntus, Pteromyscus, 281
pumilio, Alactagulus, 587
pumilio, Myosciurus, 313
pumilis, Thccurus, 199, 211
pumilis yroup, Thecurus, 21 1
pumilus, Callosciurus, 356
punctata, Dasyprocta, 191, 194
punctata, Petaurista, 282, 284, 286
punctalissimus. Callosciurus, 351, 358
punctatus, Echimys, 109, iii, 113
punctatus, Heliosciurus, 401, 402
pundti, Ctcnomys, 169
punensis, Lagidium, 230, 232
puralis, Myoprocta, 196, 197
purpurcus, Ratufa, 385
pusillus, ,\nomalurus, 540, 541
pusillus group, Anomalurus, 541
pusillus. Sciurillus, 317, 318
pusillus croup, Sciurillus, 318
pusillus, Thomomys, 518
pusillus, Thryonomys, 148
pvgacanthus, Orthogeomvs, 532
PYGERETMLS, 3 = . 5i\ 59. 276, s8o, ^87.
588
pygery-thrus, Callosciurus, 351, 353, 373
pyger>thrus group, Callosciurus. 352. 372
pygmaeus, Alactagulus, 587
pygmacus, CitcHus. 442, 443
pygmaeus group, Cilellus, 440, 443
pygmaeus, Thomomys, 522
pyladei, Sciurus, 336
pvrcnaicus, Glis, 623
PYROMYS, 38
pyrrhinus, Sciurus, 329, 342
pyrrhomerus, Dremomys, 381, 383
pyrrhonotus, Sciurus, 329, 345
pyrrhopus, Funisciurus, 412, 413
pyrrhopus group, Funisciurus, 412, 413
pyrrhotrichus, Cynomys, 462
pyrrocephalus, Menctcs, 390
pyrsonota, Ratufa, 384, 388
quadratus, Thomomys, 523
quadrimaculatus, Tamias, 430, 435
quadrivittatus, Tamias, 430, 433
quadrivittatus group, Tamias, 430, 433
quantulus, Callosciurus, 351, 360
qucbradensis, Sciurus, 341
quelchi, Sciurus, 329, 342
quemi, Capromys, 129
qucrccti, Glaucomys, 295
qucrcinus, Eliomys, 615
qucrcinus, Sciurus, 329, 334
quercinus, Thomomys, 519
quichua, Coendou, 185, 187
quindianus. Sciurus, 329. 340
quinqucstriatus, Callosciurus, 351, 373
quinquestriatus group, Callosciurus, 253* 373
quotus, Paraxerus, 406, 407
raceyi, Marmota, 459
rafflesi, Callosciurus, 351, 365
rafiventer, Sciurus, 334
rainieri, Aplodontia, 259
rajah, Petaurista, 282, 285
rajasima, Callosciurus, 376
ramosus, Citellus, 442, 445
raptor, Graphiurus, 607, 610
raptorum, Funisciurus, 412, 414
raptorum, Thnonomys, 148
rattinus, Proechimvs, 118, 122
RATTCS, viii, 3,8, 36. 38, 47. 48. 50. 52.
56. 603
rattus, Rattus, 47
RATUFA, 30, 49, 65, 263, 267, 271, 309, 319,
349. 377, 383
recessus, Ctenomys, 163, 164, 165
redimitus, Callosciurus, 368
regalis, Tamisciurus, 347
reguli, Petaurista, 282, 287
REITHRODON, 39. 55
REITHRODONTOMYS, 39. 53. 54
relictus, Capromys, 129
relictus, Citellus, 444
relictus, Thomomys, 511
rcpens, Heteromys, 474, 476
repentinus. Castor, 467
retectus, Nannosciurus, 315, 316
RHABDOMYS, 38
RHAGOMYS, 39
RHEOMYS, 39, 54
RHEITHROSCICRUS, 30, 49. 65. 260, 262.
263, 267. 271. 305, 327, 393
RHINOSCILRLS, 30,4^65. 263.267,271.
305. 349. 3S0. 395
rhionis. Rhinosciurus, 396, 398
rhipidurus, Echimvs, iii, 113
RHIPIDOMYS, 39, ^4
RHIZOMYIDAE, 37", 48, 600. 644
6R2
INDEX
KHIZOMYS, 10. :;7. 44, 4S, ^o. 60, (^b, 5<)(,,
637, 645. 646
rhoadsi. Sciurus, 344
rhodesiiic, Ht-liosciurus. 401, 402
rhodopt-nsis, Sciurus, i-?i
RHOMHOMVS. 36. 40. ^o
RHYNCHOMYINAE. 39
RHYNCHOMYS. 36. 39, 4S. 396
nchardsoni, Citeilus, 442. 447
nchardsoni proup, Citt-IIus. 447
nchardsoni, Coendou, 1S7
nchardsoni, Dipodomys, 4<j6. 502
nchardsoni, Tamiasciurus. 34(1
richniondi, Dasyprocta, i(>4
nchmondi. Sciurus. 340
ngidus, Sciurus, 329, "^37
npanus. Thomomys. 517
nudonensis, Dremomys. 383
riudoni, Callosciurus, 354
roancnsis. Napaeozapus, 574
roberti, Callosciurus, 351, 359
roberti, Coendou, 185, i86,'i8.S
roberti, Proechimys. 118, 121
roberti, Sciurus, 343
robertsoni, Funambulus, 378, 3S0
robinsoni, Callosciurus, 351. 357
robinsoni, Rhinosciurus, 396. 39S
robusta, Cavia, 242
robusta, Marmota, 457, 460
robustus, Ctenomys, 168. 1^)9
robustus, Dyromys, 619
rubustus, Heliophobius, S=i
RODHNTIA, definition oV Order, 1
rodolphei, Callosciurus, 351, 3^4
rundoni, Ctenomys, 16S
rosa, Proechimys, 118, 120
rosenbergi, Callosciurus, 350. 351, 353, 37=;
rosida, Cavia, 240, 241
rostratus, Lariscus, 393
rostratus, Liomys, 477
rostratus, Perognathus, 4S9
rothschildi, Coendou, 185. 187
rothschildi, Massoutiera, 559
rotundus Perognathus, 48b
ruatanica, Dasyprocta. 191, 195
rubeculus, Callosciurus, 35S
rubellus, Cratoeeoniys. 530
rubellus, Proechimys, 120
ruber, Sciurus, 339
rubex, Callosciurus, 351, 360
rubicaudatus, Sciurus, 339
rubicundus. Petaurista, 289
rubidiventns, Callosciurus, 371
rubrata, Dasyprocta, lyi, 193
rubricatus, Heliosciurus, 404
rubripes, Kunisciurus, 413
rubnrostns, Microsciurus, 319, 320
rubru enter, Callosciurus, 350, 351, 375
rubriventer group, Callosciurus, 353, 375
rubrolineatus, Tamiascmrus, 346
ruckbeili, Allactaga, 584. 587
rudinons, Peropnathus, 488
ruta, Aplodontia. 258
rufa. Euiyzygomatomys, 125
rufescens, Ca\ia, 242
rufescens, Citellus, 442. 443
rufescens, Echinoprocta, 181
rufescens, Marmota, 457
rufescens, Menetes. 390, 391
rufescens, Perognathus, 492
rufescens, Thomom>s, 521
ruficaudus, Cit llus, 447
ruficaudus. Tamias, 434
rufifrons, Xerus. 420, 421
rufigenis, Dremomys, 381, 382
rufigenis group, Dremomys, 381, 382
rutipes. Petaurista, zSb
rufiventer. Sciurus. 329, 339
rufobrachiatus, Heliosciurus. 40-;
rufobrachium, Heliosciurus, 401, 403
rufodorsalis, Diplomys, 115
rufogaster, Callosciurus, 374
rufogularis, Callosciurus, 365
rufoniger, Callosciurus. 351, 367
rufoniger, Sciurus, 340. 343
rufuius, Cn'ptomys. 93
rufus, Sciurus, 330
ruidosae, Thomomvs. sn
RURAIA, 383
rupatius. Callosciurus. ^si. "^70
RL PKSTES, 272. 425.'426 "
rupest i<;, Citellus, 450
rupestris, Kerodon. 247
rupestris, Perognathus, 490
rupestris, Sciurus, 329, 332
rupicoia. Graphiurus, 604, 607, 608
rupinarum, Citellus. 451
russatus, Sciurus, 333
russeolus. Thomomys, 512
russeolus, Callosciurus, 355
russicus, Pteromys, 293
russus, Sciurus, 329, 330
rutilans, Sciurus, 330
rutiliventris, Callosciurus, 371
rutilus, Xerus, 420
rutschuricus, Thryonomys, 148, 149
ruwenzorii group, Heliosciurus, 404
ruwenzorii, Heliosciurus. 400. 401, 404
sabanillae, Microsciurus, 321
sabrinus, Glaucomys, 295, 296
sabrinus group, Glaucomys, 295, 296
saccatus, Geomys, 526
saccharalis, Cratogeomvs, 528
SACCOMYS, 472
SACCOSTOMUS, 38, 56
sagitta, Dipus, 591
sagitta, H\lopetes, 299
sagitta group, Hylopetes, 299
sagittalis, Geomys, 327
sagittatus, Castor, 467
salae, Myrsilus, 417
salaquensis, Sciurus, 342
salentensis, Sciurus, 343
saliens, Allactaga, 584, 586
INDEX
683
salinac, Pedetes, 551, 552
salinia, Caviell:(, 246
salinicola, Dolichotis, 248
SALPINGOTUS, 35, 51, 59, 493, 562, 574,
576
saltarius, Ctenomys, 163, 166
saltatnr, Allactaga, 584, 587
saltator, Zapus, 573
saltuensis, Sciurus, 342
salutans, Paraxcrus, 407
salvini, Liomys. 477, 478
samarensis, Callosciurus, 351, 353, 374
samaricus, Nannosciurus, 315, 316
samuiensis. Callosciurus, 364
sanctacmartae, Cocndou, 185, 1S7
sanctituciac, Dipodomys, 503
sandakanensis, Ratufa, 384, 386
sanggaus, Callosciurus, 366
santaccruzac, Myocastor, 144
sarae, Lagidium, 230, 232
sarasinorum, Callosciurus, 375
sarawakensis, Callosciurus, 351, 366
sardus, Eliomys, 615, 616
satunini, Citdlus, 444
satunini, Hystrix, 218
saturatus, Callosciurus, 369
saturatus, Citdlus, 454
saturatus, Glaucomys, 295
saturatus, Graphiurus, 607, 610
saturatus, Lagidium, 230, 232
saturatus, Lariscus, 392
saturatus, Rhinosciurus, 396
saturatus, Thomomys, 524
saturatus, Xerus, 420, 421
satumus, Echimys, iio, iii, 113
sauicri, Callosciurus. 354
savannius. Heliosciurus, 403
saxicola, Citellus, 451
saxatilis, Dipodomys, 499
saxatilis, Galt-T, 243
saxatilis, Thomomys, 513
sayii, Sciurus, 339
scalops, Orthogeomvs, 531, S32
SCAPTKROMYS, '39, 54
scapterus. Thomomvs, 516
SCARTL'RCS, 494. 561. 580. 583
schistacea, Abrocoma, 154, 155
SCHIZODON, 157
schlfgeli, Callosciurus, 367
schlueteri. Jaculus, 595, 597
schmidti, Allactaga, 585
schmidti, Citdlus. 445
schmitzi, Hystrix, 217. 219
schumakovi, Spcrmophilopsis, 425
schvvabi, Graphiurus, 612
scirti:ti:s. sSo
SCIRTOMYS.'sSo
SCIRTOPODA. 35. 51. 59, 276, 589, 591.
SCICHAVIDAE. 15
sciureus, Kcrodon, 247
SCICRI, 30, 261, 304
SCIURIDAK, 30. 255, 259
SCIURILLUS, 30, 49, 55, 64, 69, 265, 269,
306, 317
SCIUROGNATHI, 29
SCIL'ROIDAK. 30. 78. 259
SCIUROMORPH SKRIES, 29
SCIUROPTERL'S. 262. 263, 291
SCIUROTAMIAS, 30, 52. 58, 66, 263, 272.
309, 425
SCIURUS, 23, 30, 49. 52. 53. 54, 55. 58. 61,
69, 261, 262, 263, 264, 266, 267, 268, 270,
312.321,377, 380
sclateri, Thryonomys, 145, 148
scorteccii. Hetcrocephalus, 96
SCOTINOMYS, 39. 54
scotti. Callosciurus, 351. 372
scrutator, Tamias, 432
securus, Proechimys, 118, 121
scfrius, Jaculus, 595, 597
SL-gurae, Sciurus, 329, 331
scimundi, Callosciurus, 351, 357
semipallidus, Dipodomvs, 501
semipalmatus. Thr\onomys, 148
semispinosus, Proechimys. 118, 120
semivillosus, Echimys, 112
semliki, Hcliosciurus, 401. 404
senegalica, Hystrix, 216, 217, 219
senescens, Hcliosciurus, 401
senescens, Tamias, 430, 436
senex, Dremomys, 381, 382
senex, Petaurjsta, 282, 288
senex, Rhizomys, 649, 650
senex, Sciurus, 334
senex, Tamias, 430, 434
sennetti, Dipodomys, 503
seorsus. Perognathus, 492
scptenlrionalis, Citellus, 444
septentrionalis, Microsciurus, 320
sequoioensis, Tamias, 433
seraiae, Callosciurus. 371
serbicus, Spalav, 642. 643
seri, Perognathus, 489
sericeus, Coendou, 186, 188
s;'riceus, Ctenomys, 163, 165
sertonius, Proechimys. 1 8, 122
serutus, Callosciurus, 371
setchuanus, Eozapus, 569
setosus. Liomys, 478
setosus, Petinomys. 300, 301
setosus group, Petinomys, 301
setosus, Proechimys, 117, 118, 122
setosus, Xerus, 422
severtzovi, Allactaga. 584, 585
severtzowi, Sicista, 567
shanicus, Callosciurus, 351, 362
sharpei, Funisciurus. 412
shasiensis, Castor. 468
shawi, Thomomys. 523
shddoni. Marmota. 459
shddoni. Thomomys, 521
shitkovi, Pygeretmus, 588
shitkovi group, Pygeretmus, 588
shindi, Hcliosciurus, 402
shiptoni, Caviella, 239, 246
6S4
shircnsis, Heliosciurus, 402
short r id LjL-i. C;iIlosciurus, 35 t, j^'io
siamensis. Cullosciiirus, 351. ^fu
siantanicus. Callosciurus, 356
sibcTU, Callosciurus, 357
sibt-ru, L riscus, 392
sibil-, Marmota, 451;
sibirica, Allactaga, 5S6
sibirica group. AUactapa. 5S3. 5S4, 5S6
sibirica, Marmota, 457, 460
sibirica, Sicista. 567
sibiricus. Ptcromys, 293
sibiricus. Tamias, 430, 43s
SICISTA. ^, 22, 34, 46, ^'i. su,
SICISTINAE, 34. 563, 564
siccus. Citellus. 445
siccus, Perognathus, 489
sierrae, Cuniculus, 224, 235
sierrae, Marmota, 458
SIGMODON, 39, 5^ 54. ss. boT.
SIGMOMYS, 39
signatus, F"unambulus, 379. 3N0
signatus, Protoxerus, 415, 4 if'
silanus, Sciurus, 331
silus, Glaucomys. 295
silvifugus, Thomom>s, 514
siinilis, Callosciurus, 351, 373
siniilis, Dipodomys, 501
similis, Microsciurus, 319. 320
siniiolus, Dipodomys, 496, 501
simonsi, Coendou, 185, 187
simonsi, Alicrosciurus, 319, 320
simonsi, Octodontomys. it>o
simonsi, Proechimvs. 118, 121
SIMOSCICRCS,'265, 322, 325, 326
simplex, Xerus, 421
simulans, Dipodom\'s, 503
simulans, Tamias, 434
siniulus. Thomomys, 521
simus, Glyphotes, 393
smaloae, Thomomys, 517
sinaloensis, Sciurus, 329, 335
sindi, Paraxerus, 406, 407
sinensis, Rhizomys, 650
sinensis group, Rhizomys. C150
singapurensis, Callosciurus, 351. 370
smhala. Ratufa, 384, 385
sinistralis, Callosciurus, 351, 362
SINOETHERCS. 1S2
sinus. Ratufa, 384. 389
sipora, Ioni>s, 304
sirhasstnensis, Ratufa, 3S4, 387
siriensis, Callosciurus. 371
siskiyou, Tamias, 434
sladeni. Callosciurus, 3s i, ^^2., ;6o
SMINTHCS, 564
smithi, Graphiurus, '107, 610
smithi, Lophiomys, 63b
smithi, Ratufa, 384, 3S6
soccatus, Paraxerus, 406, 407
socialis, Cynomys. 462
socialis, Sciurus, 335
sodalis, Cavia, 241
INDEX
561. 562, 564
soderstromi, Sciurus, 340
soleatus, Graphiurus. 607, Gog
solitarius. Castor. 4*18
solitarius, Thomomys, 517
solivagus, Tamias, 434
solutus. Callosciurus, 360
somaliensis. Hystrix, 217, 220
sonomae, Tamias, 435
sonoranus. Lionns, 477
sonoriensis, Citellus, 453
sonoriensis. Dipodomys, 504
sonoriensis, Perognathus, 484
sonoriensis. Thomom\s, 521
sordidus, Callosciurus, 351, 356
sordidus, Nannosciurus. 316
soricinus, Nannosciurus, 31b
soricinus, Zapus, 571
sowerbyi, Dipus, 591
spadiceus, Hvlopetes. 298, 2uu
SPALACIDAE, 37, boo, 636
SPALACOPUS, 3,2b, ss. 73. 10^. iss, is6,
160, 554
SPALAX, 3, 10, 22, 3b. 37. 43, 45, 51, 60,
81, 599, G37, 638, 640, b4i, b43
spalax, Heliophobius, 85
speciosus, Muscardinus, b2s
speciosus, Tamias, 430, 433
spectabilis. Dipodomys, 49b. 497. 499
spectabilis group, Dipodomys. 497. 499
spekei, Pectinator, 556
spencei. Callosciurus, 353
SPERiMOPHILOPSIS. 23, 70, 32, s8, 272,
306, 423
SPEUMOPHILUS, 262, 268, 4^,7
SPHIGGCRUS, 182, 185, 187
spiciitus. Mesomys, 126, 127
spiculum. Allactaga, 586
spilosoma, Citellus, 441, 442, 449
spilosoma group, Citellus, 449
spilotus, Perognathus. 488
spinatus, Perognathus, 491
spinatus group, Perognathus, 483, 491
spinosus, Coendou, iSb, 18S
spinosus, Euryzygomatomys. 125
spixii. Galea, 242, 243
splendcns, Callosciurus, 351, 3bi
splendidus, Sciurus, 342
spoliatus, Glis, b22, 623
sponsus, Paraxerus, 40b, 408
spurrelli, Graphiurus, 607, bii
squamicaudatus, Anomalurus, 540
stangeri, Protoxerus, 41s
STEATOlMYS, 39, 56 *
steerei, Callosciurus, 351, 353, 374
steerei, Proechimys, 119
stegmanni, Hvstrix, 216, 219
steinbachi, Sciurus, 329, 345
steinbachi, Ctenomys, ib3, 1^4, ibS
stejnegen. Citellus, 44b
stellaris, Callosciurus, 372
stillatus, Crvptomvs, 93
STENUCEPHALEMYS, 38
stephanicus, Xerus, 420, 421
INDEX
68s
stephensi, Citellus, 446
stephcnsi, Dipodomys, 499
stephensi, Glaucomys, 297
stephensi, Pcrognathus, 489
stevensi, Athcrurus, 206
stevensi, Caliosciurus, t;i, 'i^i. -177
STICTOMYS, 22, •■'SJ'"3
stigmosa, Ratufa, 384, 388
stimulax, Mcsomys, 126, 127
stirlingi, Mamiota, 457, 461
stocklcyi, Pctaurista. 287
stolida, Cavia, 240, 241
stont-i, Citellus, 448
stramineus, Citellus, 446
straminous, ,Sciurus, 323, 329, 343
stramineus group, Sciurus, 327, 343
strandi, Allactaga, 585
strandi, Sicista, 567
streatori, Dipodomys, 496, 498
streatori, Tamiasciurus, 346
streeteri, Graphiurus, 611
stresemanni, Caliosciurus, 369
striatus, Tamias, 430, 436
styani, Caliosciurus, 351, 359
st\'gius, Heterocephalus, 95
STYI.ODIPUS, 591, 592
suahelicus, Paraxerus, 406, 408
subalbidus, Protoxerus, 416
subauratus. Castor, 468
subauratus, Sciurus, 339
subcristatus, Hystrix, 200, 214, 216, 217,
subcristatus group, Hystrix, 212, 213, 21
subflaviventris, Dremomys, 381
sublaevis, Cuniculus, 225
sublineatus, Funambulus, 377, 379, 3S0
sublineatus group, I*\inambulus, 379
sublucidus, Perognathus, 489
subluteus, Caliosciurus, 351, 370
sublutcus, Cratogeomys, 530
subniger, Cuniculus, 225
subnubilis. Cratogeo.iiys, 530
suboles, Thomomys, 518
subrosea, Lagidium, 230, 232
subrufus, Graphiurus, 610
subsimilis, Thomomys, 518
subsimus, Cratogeomys, 529
subspinosus, Chactomys, 177
substriatus, Funisciurus, 412, 414
subtilis, Sicista, 566, 567
subtilis group, Sicista, 566, 567
subviridescens, Heliosciurus, 404
sucklcyi, Tamiasciurus, 347
suffusus, Caliosciurus, 351, 367
suillus. Bathyergus. 84
sulcatus, Pctaurista, 282, 289
sulcatus group, Pctaurista, 285, 289
sulcidens, Cartcrodon, 126
sullivanus, Caliosciurus, 351, 363
sumaco, Sciurus, 341
sumatrae, Thecurus, 199, 211
sumatrana, Caliosciurus, 368
sumatrana, Pctaurista, 289
sumatranus, Nannosciurus, 316
218
■44. 145.
sumatrensis, Rhizomys, 650, 651
sumbawae, Hystrix, 214, 217
supcrans, Ivliomys, 6i5
supcrans. Ratufa, 3S4, 385
superciliaris, Paraxerus, 406
supcrnus, Thomomys, 520
surdaster, Pedetes, 551. 552
surdus, Caliosciurus, 351, 356
surdus, Graphiurus, 602, 607 609
surdus group, Graphiurus, 609
surrutilus, Nannosciurus, 316
SUS, 47
suschkini, Allactaga, 586
suslicus, Citellus, 428, 442, 445
suslicus group, Citellus, 440, 445
swarthi, Dipodomys, 499
swinderianus, Thryononiys, 148
swinderianus group, Thrvononivs,
148
swinhoci, Caliosciurus, 351, 355
su-ynnertoni, Paraxerus, 406, 408
syhilla, Pctaurista, 282, 286
sylvanus. Ctcnomys, 163, i56
Sylvester, Caliosciurus, 351, 373
SYNAPTOMYS, 40, 52
SYNTHEOSCIURLS, 30, 69. 265, 266, 268,
270, 282, 312, 321
SYNTHERES. See Sinoetherus
syriacus, Sciurus, 333
syrius, Jaculus, 595, 597
syrmiensis, Spalax, 643
tabandius, Caliosciurus, 351, 364
tabanus, Spalacopus, 161
taborae. Pedetes, 552
tachardi, Caliosciurus, 351, 361
tachin, Caliosciurus, 362
TACHYORYCTAE, 40
TACHYORYCTINAE, 40
TACHYORYCTES, 10, 22, 3S, 40, 56, 602,
637. 645
tacomensis, Thomomys, 523
tacopius, Caliosciurus, 351, 364
taczanowskii, Cuniculus, 224, 225
taczanowskii group, Cuniculus, 224, 225
taedifcr, Sciurus, 329, 344
taeniurus, .Sciurus, 337
taguan, Pctaurista, 285
tahan, Caliosciurus, 351, 356
talarum. Ctcnomys, 163, 165
talboti. Funisciurus, 412, 413
talpoides, Cryptomys, 87, 90
talpoides, Thomomys, 510, 521
talpoides group, Thomomys, 510, 521
tamae, Sciurus, 340
tamansari, Caliosciurus, 351, 369
tamaulipcnsis, Cratogcomvs, 529
TA.MIAS, 23, 30, 44, 45,' 51. 52, 53, 58, 61,
70, 262, 263, 266, 267, 268, 272, 308, 425,
426, 439
TAMIASCIURUS, 30, 53, 54, 61, 264, 266,
267, 268, 269, 270, 310, 345, 399
5S6
INDEX
TAMIODKS, 267. 37(1, 377
TAMIOPS, 263. 2fi7. ^^>o, 34S. 340. 350, 1,^},
354
TAMISCL'S, 2(hj. 405
tanae, Paraxurus, 406, 408
tangan>ikac, Funisciurus, 412. 415
tanganyikae, Paraxerus, 401)
tanaiticus, Dyroinys, 619
tapanulius, Callosciurus, 351. 370
taparius. Sciurus. 329, 345
tarussanus, Callosciurus, 36(>
tasmani, Graphiurus. 611
tataricus, Marmota, 460
TATF.RA. 40, 48, 50, 56
TATKRILLUS. 40
taylori. Pttaurista, 2S2, 2SS
taylori. Perognathus, 486
tcdoneus. Callosciurus, 372
telibius, Callosciurus, 351, 3^^^
teluni, Scirtopoda, 5^2
temmincki, Protoxerus, 415. 416
temporalis, Hcteromvs, 476
TENES. 267. 322. 324, i,jji
tenellus. Zapus, 572
tt-nnanti, Ratufa, 384
tenuirostris, Callosciurus, 351, 370
tenuis, Callosciurus, 350, 351, 355
tenuis group, Callosciurus, 351, 355
tephrogaster, Sciurus, 337
tephromelas, Aerom\s, 291
tephromelas group, Aeromys. 291
tepicanus, Sciurus. t,}^
tereticaudus, Citellus, 442. 452
tersus, Citellus, 452
terutaus, Atherurus, 206
terutaus, Petaurista, 285
teruta\ensis, Callosciurus, 351, 365
tescoruin, Citellus, 453
tesludo, Lophiotnys, 636
tetradactyla, Allactaga, 583, 584
tetradactyla group, Allactaga, 584
texensis. Castor, 467
texensis, Citellus, 44H
texensis, Geoniys, 527
texensis, Glaucomys, 295
texensis, Liomys, 479
texensis, Thomomys, 51S
texianus, Sciurus, 33"^
thai, Callosciurus, 3fi2
thaiwanensis, Callosciurus. 3^1, 359
THALLOMYS. ^H
THAMNOMYS. 38, 56
thayeri, Sciurotamias, 426
THECURUS, 23, 27, 48, 66. 198, 208. 209
THERIDOMYIDAE, 13, 598
thermaicus, Spulax, 642, 643
THOMAS, (JLDEIELD, CLASSIFICA-
TION OF ORDER, 5, 6, 7
thoniasi, Aeromys, 291
thomasi group, Aeromys, 291
thomasi, Cuniculus, 225
thomasi, I^chimys, 109, iii, 112
thoniasi, Funambulus, 379, 380
thomasi, Eophiomys. 636
thomasi, Petaurista, 2S9
thomasi, Salpingotus, 574, 575
thomasi, Sciurus, ^37
TII(>MASOMYSr39. 54
THOMOMYS, 32, 53, 63, 71, soh, 507
thomsoni, lomys, 303
thoracatus, Geocapromys, 132
THRICHOMYS, 123
THRINACODUS. 2^, ss, 7^ 102, 135
THRYONOMYINAE, 26, 105. 144, i6q
THRYONOMYS, 23, 26, 42, 57. 68. oS, loi.
103, 139, 144
tianschanica, Sicista, 366, 568
tichomirowi, Dyrom\s, 618
tigrina, Marmota, 461
tingahi. Callosciurus, 375
tiomanensis, Ratufa, 384, 3S9
tiomanicus, Callosciurus, 351, 356
tionis, Atherurus, 206, 208
tobagensis, Sciurus, 329, 341
toltecus, Thomom>"s, 510, 318
lolucae, Sciurus, 339
tolucae, Thoniom\'s, 320
TOMEUTES, 263, 266, 267, 348. 349
tongensis, Paraxerus, 408
tonkeanus, Callosciurus, 375
tootalis, Lagidium, 230, 232
topapuensis, Callosciurus, 375
torquata, Hystrix, 217
torquatus, Ctenomys, 163, 164, 167
torrentiuni, Protoxerus, 416
torridus, Heterogeomys, 333
torridus, Liomys, 478
tortilis, Chaetomys, 177
tosae, Petaurista, 282, 288
townsendi, Citellus, 442, 446, 447
townscndi group, Citellus, 446
townscndi, Taniias, 430, 431
townsendi group, Tamias, 430, 434
townsendi, Thomomys, 511
to%vnsendi group, Thomom\s, 310, 311
transbaicalicus, Citellus, 446
transsyhanicus, Spalax, 642, 643
transvaalensis, Cn'ptomys, 93
trapezius, Muscardinus, 623. 626
TRECHOMYINI, 18
trepidus, Citellus, 433
trichodactylus, Lagostomus. 236
trichopus, Zygogeomys, 533
trichotis, Bclonivs, 277, 279
TRICHYS, 27.' 48. 66, 102, 106, 199, 202,
203, 20S
tricolor, Coendou, 185, 1S7
tricolor, Sciurus, 329, 344
tridecemlineatus, Citellus, 269, 441, 442. 448
tridccemlineatus group. Citellus, 448
tnhneatus, Funambulus, 3S0
tniiitatis, Citellus, ^f54
tnnitatis, Dipodomys, 49S
trinitatis, Proechim\s, 118, 121
TRINOMYS. 1 15," 116. 117, 122
trinotatus, Zapus, 371, 573
INDEX
6S7
tripartitus, Sicista, 567
tristriatus, Kunanibulus, 377, 370, 380
tristriatus. Sicista, 567
trivittatus, Allantoxerus, 423
trizona, Sicista, 566, 567
TROGONOTHERIUM, 466
TROGOPTKRUS, 30. 52, 57. 64, 263, 274.
275, 279. 304
tropicalis, Gcomys, 527
tropicalis, Petromus, 149, 151
trotteri, Callosciurus, 361
truei, Hoplomys. 122, 123
truci, Sciurus. 335
trumbullcnsis, Thomomys, 513
tschetschenicus, Glis, 623
tschudii, Cavia, 240, 241
tsinetauensis, Callosciurus, 358
tuconax, Ctenomys, 163, 164, 168
tucumanus, Ctenomys, 163, 165
tucumanus. Lauidium, 230, 231
tularensis, Dipodomys, 498
tularosac-. Citcilus, 450
tularosae, Thomomys, 519
tulduco, Clt;nonivs, 163, 166
TULLBKRG, CLASSIFICATION OF
ORDKR, 8, 9, 10
tunctae, F.Homys, 616
tupaioides, Rhinosciurus, 396
turcicus, Spalax, 643
turcomanus, Spermophilopsis, 425
turnbulli, Hylopetes, 300
turneri, Atherurus, 206, 208
turovi, Pteromys, 294
turovi, Scirtopoda, 592
tuza, Geomys, 526
tuza group. Geomvs, 525, 526
TYLOMYS. 39, 54
tylorhinus, Platvgcomvs, 531
TYPHLOMYS. 36, 37, 48. 66, 599, 601, 602,
626, 627, 629, 632
tj'phlus, Spalax, 644
typicus, Graphiurus, 608
typicus, Pedetes, 552
typicus, Petromus. 149, 151
typicus, Sciurus, 330
typus, Dactylomys, 136
tzaneensis, Graphiurus, 610
ubericolor, Callosciurus, 370
ugandae. Paraxerus, 409
uinta, Thomomvs, 522
UJHELYIANA. 638
ukrainicus, Sciurus, 332
umbrata, Ca\'ia, 240, 241
umbratus, Citcilus, 442, 445
umbriceps, Rhizomys, 650, 651
umbrinus, Tamias, 433
umbrinus, Thomomys, 510, 519
umbrinus group. Thomomys, 510. 519
umbrosus, Alenetes, 390. 391
umbrosus, Pcrognathus, 490
umbrosus, Tamias, 436
underwoodi, Dasyprocta, 194
underwoodi, Glaucomys, 296
underwoodi, Macrogcomys, 534
underwoodi, Sciurus, 336
undulatus, Citellus, 443
undulatus, Heliosciurus, 401, 402
ungae, Citellus, 443
unicolor, Kchimys, 112
uralensis, Sciurus, 332
L'RANOMYS, 38, 56
urichi, Proechimvs, 118, 121
L ROCITKLLUS, 437, 439
TROMYS. 38, 47
IROSCIL'HUS. 265, 322. 326, 327
uru^uma, Dasyprocta, 195
urucumus, Sciurus, 345
Utah. Citellus, 451
utahensis. Dipodomys, 502
utahensis, Tamias, 435
utahensis, Zapus, 572
uthensis, Tamias, 436
utibilis, Ctenomys, 163, 167
vaccarum, Abrocoma, 154, 155
vacillator, Proechimys, 117, 118, 121
vae, Felovia, 560
vagus, Sicista. 567
valdiviae, Sciurus, 342
vali, Ctenodactylus, 558, 559
vallicola, Tamias, 432
vanakeni, Callosciurus, 351, 357
vancouverensis, Marmota, 459
vancouverensis, Tamiasciurus, 346, 347
vandami, Cr\'ptomys, 93
vandami, Graphiurus, 612
VANDELEURIA, 3, 38, 48
vanheurni, Callosciurus, 369
vanrossemi, Thomomys, 513
variabilis, Sciurus, 329, 341
variegata, Dasyprocta, 191, 195
variegatoides, Sciurus, 329, 336
variciiatus. Castor, 468
variegatus, Citellus, 442, 450
variegatus. Thr\onomys, 148
varius. Castor. 468
varius, Sciurus, 329, 330
vassali, Callosciurus, 363
velUgera, Chinchilla, 229
venezuelae, Cavia, 241
venningi, Petaurista, 282, 287
vcntorum, Tamiasciurus, 346
venustulus. Microsciurus, 319
venustus, Dipodomys, 503
venustus, Pcrognathus, 485
venustus, Sciurus, 342
venustus, Tamias, 437
veraecrucis, Liomys, 477
verbecki, Callosciurus, 369
versicolor, Sciurus, 329, 341
vescus, Thomomys, 513
vestitus, Callosciurus. ;iS5
vcstitus, Coendou, 185, 186, 188
6SS
INDEX
vestitus, Rhizoni>s, 650
vtstitus proup. Rhizomys, 650
vetensis, Cr\ptomys, 03
vexillaris. Allacta^a. 586
vexiliarius. Hehosciurus, 400, 401, 405
viatorum. Lagidium, 230, 232
\icinus. Kozapus, 56Q
vicinus. Sciurus, 33S
vigilis. Citcllus, 446
vigilis, Marmota, 459
villosus, Belomys, 277
villosus. Coendou. 1S5, i8f), i SS
\il]osus, Isothrix, 114
vinulus. Citellus. 442, 452
viperinus. Ctenomys, 163, if>fi
virgatus. Cuniculus, 225
virgo. Callosciurus, 351, 373
virgulosus, Sicista, 567
VISCACCIA. 229
\iscaccia, Lagidium, 230, 231
viscaccia, Lagostomus, 236
viscaccica, Lagidium, 231
vistulanus, Castor, 468
vittata, Ratufa, 387
vittatus, Callosciurus, 34g, 351, t,s~' :>^9
vittatula. Ratufa, 387
vivatus. Microsciurus, 320
vivax. Sciurus, 337
VIZCACIA, Z3i
vocator, Jaculus, 595, 597
vociferans. Citellus, 452
volans, Glaucomys. 295
volans group, Glaucomys. 295
volans, Pteromys, 293
volucella, Glaucomys, 295
\ordermanni, Petinom>"s, 301
vryburgensis, Cryptomys. 93
vulcani. Dipodomys. 501
vulcani, Lagidium, 230, 231
vulcani, Liomys. 478
vulcani, Orthogeomys, 532
vulcanicus, Graphiurus, 612
vulcanius, Heliosciurus. 401. 404
vulcanius, Thomomys, 520
vulcanorum, Paraxcrus, 406. 409
vulcanus, Lanscus, 392
vulgaris, Glis, 622
vulgaris. Pteromys, 293
vulgaris, Sciurus, 324, 327, ^Zi). 330, 346
vulpinus, Sciurus, 33S
vulturous, Jaculus, 595, 596
wagneri, bciurus, 334
wallowa, Thomomys, 523
wardi, Rhizomys, 650
warreni, Marmota, 438
warreni, Proechimys, ii-S. 121
Washington!, Cittllus. 442, 447
washmgtom i^roup. Citellus, 447
washoensis, Citt-llus, 44b
watasei, Petaurista, 289
waterhousii, Heliosciurus, 403
watsoni, Callosciurus, 3=;i, "^72
WKBER, CLASSIFICATION OK ORDKR,
II, 12, 13
weberi, Callosciurus, 375
weigoldi, Sicista, 567
wellsi, Callosciurus, 351. 359
wellsi. Cialea, 243
WEVENBHRGHIA. 247
whiteheadi, Nannosciurus, 315, 316
whiteheadi group. Nannosciurus, 316
whytei, Cryptomys, 87, 90
williamsi, Allactaga. 5S4, 585
williamsi group, -Allactaga, 584, 585
wiUiamsoni, Callosciurus, 351, 361
wilsoni, Epixerus, 417, 41S
WINGE, CLASSIFICATION OFORDKR,
18, 19, 20. 21
wingei, Dyromys, 618, 619
wintoni, Funisciurus, 413
wolfTsohni, Lagidium, 230, 232
woodi, Callosciurus, 359
woosnami, Graphiurus, 607. 6o()
woosnami group, Graphiurus, 609
wortmani, Citellus, 442, 453
wrayi, Callosciurus, 351, 365
wroughtoni. Funamhulus, 379, 3S0
xanthipes, Trogopterus, 279
xanthodon, Spalax, 644
xanlhonotus, Perognathus, 487
xanthoprymnus. Citellus. 442, 444
xanthotis, Petaurista, 282. 283, 288, 289
xanthotis, Sciurus, 340
XEROMYS, 39, 47
XEROSPERMOPHILUS, 268. 437, 441,
442, 452
X1':RUS, 23, 30, 57. 68, 240, 2()0. 262. z(>4,
2by, 268, 272, 306, 377, 418
XYLOMYS. 472, 474, 476
yakimensis, Citellus. 446
yatesi, Georychus. 86
yelmensis, "^rhomomys, 523
youngi. Callosciurus, 351, 358
yucatanensis, Heterogeomys, 533
yucatanensis, Sciurus, 329, 336
yucataniae, Coendou, 183, 187
yucatanica, Dasyprocta, 194
yukontns.s, Glaucomys, 296
yulei, Paraxerus, 406, 407
yungarum, Dasyprocta. 191. 105
yunnanensis, Hystrix, 218
yunnanensis, Pt-taurista. 282, 287
zacatt'cae, Perognathus, 4S8
zacatecae, Thomom%*s, =;20
INDEX
689
zaissanensis, Dipus, 591
zamorae, Dasyprocta, 195
zamorae, Sciurus, 345
zaphaeus, Glaucomys, 296
ZAPODINAK, 34. 561, 562, 563. 568
ZAPUS, 35. 53, 63. 562, 564. 568, 569
zarumae, Sciurus, 343
zechi. Crvptomvs, 87, 91
ZELOTOMYS; 38, 56
ZENKEKE LLA, 3. 33, 57. 68, 537, 542. 543.
546
zenkeri, Idiurus, 544, 546
zenkeri group, Idiurus, 546
ZETIS, 26:, 380
zeus, Muscardinus, 625
zeylanicus, Katufa, 384
zeylonensis, Hystrix, 218
zimmeensis, Callosdurus, 351, 362
zonalis, Hcteromys, 475
zuliae, Sciurus, 342
zuluensis, Graphiurus, 612
zuluensis, Hystrix, 219
zuniensis, Cvnomvs, 463
ZYGODONTOMYS. 39
ZYGOGEOMYS. 32, 71, 506, 507, 534
zygomaticus, Atherurus, 206, 208
zvgomaticus, Dipodomys, 499
ZYZOMYS, 38, 47
2 8 JUN 1940
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