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Full text of "The families and genera of living rodents"

'j^A. 





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 -. 



(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 (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 CALIFORNI